Flora of the Venezuelan Guayana [7]

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Flora of the Venezuelan Guayana VOLUME 7 MYRTACEAE–PEDALIACEAE

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

MISSOURI BOTANICAL GARDEN PRESS St. Louis

Copyright © xxxx by the Missouri Botanical Garden Press All rights reserved. ISBN x-xxxxxx-xx-x Printed in U.S.A. Published on xxxxxxxxxxxx by Missouri Botanical Garden Press P.O. Box 299 St. Louis, Missouri 63166-0299, U.S.A.

Contents Preface and Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix Contributors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiii Myrtaceae (Bruce K. Holst, Leslie Landrum, and Francesca Grifo) . . . . 1 Najadaceae (Robert R. Haynes and Lauritz B. Holm-Nielsen) . . . . . . . 100 Nyctaginaceae (Julian A. Steyermark and Gerardo A. Aymard C.) . . 101 Nymphaeaceae (John H. Wiersema) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118 Ochnaceae (Claude Sastre) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124 Olacaceae (John M. MacDougal) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162 Oleaceae (James S. Miller and Gerardo A. Aymard C.) . . . . . . . . . . . . 186 Onagraceae (Elsa M. Zardini and Peter H. Raven) . . . . . . . . . . . . . . . . 188 Opiliaceae (James S. Miller) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198 Orchidacaeae (Germán Carnevali, Ivón M. Ramírez-Morillo, Gustavo A. Romero-González, Carlos A. Vargas, and Ernesto Foldats) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200 Oxalidaceae (Gerardo A. Aymard C. and Paul E. Berry) . . . . . . . . . . . 619 Passifloraceae (Stephen S. Tillett) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 625 Pedaliaceae (James S. Miller) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 667 Peridiscaceae (Bruce K. Holst) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 670 Phytolacacceae (Gerardo A. Aymard C. and Nidia L. Cuello A.) . . . . 672 Piperaceae (Julian A. Steyermark and Ricardo Callejas-Posada) . . . 681 Plantaginaceae (Gladys Rodríguez) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 739 Plumbaginaceae (Carlos A. Vargas) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 740

Appendix: List of new names published in this volume . . . . . . . . . . . . 742 Index (compiled by George Thornburgh) . . . . . . . . . . . . . . . . . . . . . . . . 743

v

Preface and Acknowledgments This volume continues the taxonomic treatments of all native and naturalized vascular plant families known to occur in the Venezuelan Guayana. It includes 18 families of flowering plants, from Myrtaceae through Plumbaginaceae. The remaining families will be published in alphabetical order in two 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. The University of Wisconsin–Madison has also supported this project in its later stages. This volume is based upon work supported by the National Science Foundation under Grants Nos. BSR-8717303, BSR-9045532, and BSR-9201044. The Foundation provides awards for research in the sciences and engineering. The awardee is wholly responsible for the conduct of such research and preparation of the results for publication. The Foundation, therefore, does not assume responsibility for such findings or their interpretation. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the authors and do not necessarily reflect the views of the National Science Foundation. The Julian A. Steyermark Fund, established by the late Dr. Steyermark at the Missouri Botanical Garden, provided major funding for the flora. The Lewis B. and Dorothy Cullman Foundation provided funding for the Gentianaceae, in part. The Armand G. Erpf Fund provided financial support for part of the plant illustrations in this volume. Besides the illustrations by Bruno Manara, Rossana Marrufo did the drawing of Eurystyles cotyledon (Fig. 336), and G. C. K. Dunsterville did the drawings of Huntleya meleagris (Fig. 357), Kegeliella orientalis (Fig. 367), and Ornithocephalus falcatus (Fig. 439). The Dunsterville drawings are courtesy of Orchid Herbarium of Oakes Ames. 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. We are grateful to Maria do Carmo Amaral, Peter Dodson, Peter Jorgensen, and Valery Malecot for their reviews of certain family treatments. 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 vii

viii P REFACE AND A CKNOWLEDGMENTS

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

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

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x INTRODUCTION

FAMILIES INCLUDED IN THIS VOLUME Genera Species Myrtaceae Najadaceae Nyctaginaceae Nymphaeaceae Ochnaceae Olacaceae Oleaceae Onagraceae Opiliaceae

17 1 6 1 12 9 1 1 1

183 1 43 7 120 31 2 17 2

Genera Species Orchidaceae Oxalidaceae Passifloraceae Pedaliaceae Peridiscaceae Phytolaccaceae Piperaceae Plantaginaceae Plumbaginaceae

157 2 3 2 1 6 2 1 1

732 8 53 2 1 10 124 1 1

221

1338

the numbers of genera and species currently recorded from the flora area appears in Appendix A of Volume 1. The 18 families that appear in Volume 7 and their numbers of native or naturalized genera and species in the Venezuelan Guayana are given above. Organization and Scope of the Floristic Treatments The floristic treatments in this flora are organized alphabetically by family, first within the ferns and fern allies, then within the seed plants. The gymnosperms, monocotyledons, and dicotyledons are not treated separately within the seed plant category. The authors for each family treatment are cited below the family heading; in multi-authored families, if different parts were contributed by different authors, the respective authors are cited separately under the parts they were responsible for (such as a particular genus or the key to the genera). Each family treatment contains a complete description of the family and the genera that are present in the flora; these descriptions are worldwide in scope. Within each family, genera are arranged in alphabetical order. At the end of each family or genus description, a separate paragraph lists the overall distribution of the family or genus. This is followed by the author’s estimate of the number of subordinate taxa worldwide, as well as the number of taxa present in the Venezuelan Guayana. For each genus, the estimated number of species in all of Venezuela is also given. All keys used in the flora are dichotomous and bracketed. In each couplet, the number in parenthesis refers to the couplet that led to it. The keys to the genera are designed primarily to cover the species that occur in the Venezuelan Guayana and not necessarily species found elsewhere. Slightly more than half the species in the flora are illustrated by black-andwhite line drawings. Volume 7 includes 646 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

INTRODUCTION

xi

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 “w” followed by the figure number appears at the end of the paragraph. A separate paragraph is sometimes used to provide taxonomic discussion and/or notes on the uses of the plant in the flora area. Any varieties or subspecies present in the flora area appear below their corresponding species entry, and they contain a similar level of information as species entries. A key is included if more than one subspecies or variety is present. Forms found in the flora area do not have separate entries, but they are discussed under the next higher rank. Throughout the flora, author abbreviations follow Authors of Plant Names (R. K. Brummitt and C. E. Powell, editors. Royal Botanic Gardens, Kew. 1992). The only exception is the use of the abbreviation “H.B.K.” for species described in Nova Genera et Species Plantarum, by A. von Humboldt, A. Bonpland, and K. S. Kunth (Librairie Grecque-Latine-Allemande, Paris. 1815–1825). This series of seven volumes was published in two versions (quarto and folio), each with different page numbers. Page numbers cited in this flora refer to the quarto version, which is more widely available than the folio version, except for volume four, which is the only volume in which the folio edition was distributed before the quarto. Book abbreviations follow Taxonomic Literature: A Selective Guide to Botanical Publications and Collections with Dates, Commentaries and Types, 2nd edition (F. A. Stafleu and R. S. Cowan. Bohn, Scheltema & Holkema, Utrecht, Netherlands. 1976– 1988), and journal abbreviations follow Botanico-Periodicum-Huntianum (G. H. M. Lawrence, A. F. G. Buchheim, G. S. Daniels, and H. Dolezal, editors. Hunt Botanical Library, Pittsburgh, Pennsylvania. 1968) or B-P-H/S: Botanico-Periodicum/ Supplementum (G. D. R. Bridson and E. R. Smith, editors. Hunt Institute for Botanical Documentation, Pittsburgh, Pennsylvania. 1991). When the effective publication date of a reference is different from the imprint date, the effective publication date is placed in brackets following the imprint date. Herbarium abbreviations follow Index

xii

INTRODUCTION

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 Gerardo A. Aymard C. Herbario Universitario PORT UNELLEZ–Guanare Mesa de Cavacas 3323, Portuguesa VENEZUELA Paul E. Berry Botany Department University of Wisconsin–Madison 132 Birge Hall 430 Lincoln Drive Madison, Wisconsin 53706 Ricardo Callejas Posada Instituto de Biología-Herbário HUA Facultad de Ciencias Exactas y Naturales Universidad de Antioquia AA 1226, Medellín COLOMBIA Germán Carnevali Herbário CICY Centro de Investigación Científica de Yucatán A. C. (CICY) Calle 43, No. 130, COl, Chuburná de Hidalgo 97200 Mérida, Yucatán MEXICO Nidia L. Cuello A. Herbario Universitario PORT UNELLEZ–Guanare Mesa de Cavacas 3323, Portuguesa VENEZUELA

Francesa Grifo Department of Ecology, Evolution and Environmental Biology, Columbia University 10th floor Schermerhorn Extension, MC 5557 1200 Amsterdam Avenue New York City, New York 10027 Robert R. Haynes Department of Biological Sciences University of Alabama 425 Scientific Collections Building Box 870345 Tuscaloosa, Alabama 35487-0345 Lauritz B. Holm-Nielsen [recheck] 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 Leslie Landrum Herbarium, Plant Biology Department Arizona State University Tempe, Arizona 85287 John M. MacDougal Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166

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xiv

CONTRIBUTORS

James S. Miller Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166

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

Ivón Ramírez Herbário CICY Centro de Investigación Científica de Yucatán A. C. (CICY) Calle 43, No. 130, COl, Chuburná de Hidalgo 97200 Mérida, Yucatán MEXICO

Stephen S. Tillett Herbário Ovalles, Facultad de Farmacia Universidad Central de Venezuela Apartado 40109, Nueva Granada Caracas 1040-A VENEZUELA

Peter H. Raven Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166

Carlos A. Vargas Environmental Consultant Urb. Palomino, Block B-3, Dept. 22 Lima 1, PERU

Gladys Rodriguez c/o Fundación Jardín Botánico de Caracas Apartado 2156 Caracas 1010-A VENEZUELA Gustavo A. Romero-González Oakes Ames Orchid Herbarium Harvard University 22 Divinity Avenue Cambridge, Massachusetts, 02138 Claude Sastre Herbier, Laboratorire de Phanérogamie Muséum Nationale d’Histoire Naturelle 16 rue Buffon F-75005 Paris FRANCE

John H. Wiersema Systematic Botany and Mycology Laboratory USDA/ARS, Bldg. 011A, BARC-West 10300 Baltimore Ave. Beltsville, Maryland 20705 Kay Yatskievych Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Elsa M. Zardini Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166

MYRTACEAE by Bruce K. Holst, Leslie Landrum, and Francesca Grifo Trees, shrubs, or rarely subshrubs, usually with abundant, small, circular glands embedded in the tissues; plants often aromatic, mostly evergreen. Leaves usually petiolate, exstipulate, opposite (in the flora area), alternate, or rarely trifoliate; blades coriaceous to membranous, entire or very rarely crenulate, pellucid-punctate. Inflorescences usually bracteate, axillary or subterminal, occasionally woody tissue present, paniculate, dichasial, racemose, cymose, corymbose, or 1-flowered. Flowers bisexual or rarely unisexual, actinomorphic, short-lived, often fragrant, subtended by a pair of bracteoles, 4- or 5(6)-merous; hypanthium adnate to ovary, occasionally prolonged above the summit of the ovary. Calyx of free lobes or lobes partially to completely connate, when connate opening by longitudinal or circumscissile tears; petals as many as sepals or rarely fewer or lacking, usually free, mostly white or rarely pinkish or bluish. Stamens numerous, rarely few, borne on ovary summit or hypanthium, occasionally in fascicles opposite the petals; filaments free or partly connate; anthers dorsifixed or rarely basifixed, dehiscing by longitudinal slits or rarely by apical pores, the connective often terminating in a gland. Ovary inferior, 2- or 3(–many)-locular; placentation axile, subapical, or basal; ovules 1–many per locule, usually biseriate or multiseriate; style simple; stigma small, usually capitate. Fruit a berry, capsule, or nut, usually crowned by the calyx remnants. Seeds one to many, the seed coat membranous, subcoriaceous, chartaceous, or hard and bony; embryo starchy or oily, variously shaped with the neotropical species falling roughly into three subtribes: (1) subtribe Myrciinae, with cotyledons membranous, folded, and hypocotyl well developed and elongate; (2) subtribe Eugeniinae, with thick, fleshy cotyledons, separate or fused, and hypocotyl very short or indistinct; and (3) subtribe Myrtinae, with very small and membranous or fleshy cotyledons, and well-developed, elongate, curved or spiral hypocotyl. Tropics and subtropics worldwide; ca. 100 genera and 3000 species, 17 genera and 183 species in the flora area. Myrtaceae are traditionally divided into two subfamilies, each with approximately the same number of species: Myrtoideae, concentrated in the New World but also found in Africa, southern Asia, and Australia; and Leptospermoideae, concentrated in tropical Asia and Australia, with a single American genus, Tepualia, found in temperate Chile and adjacent Argentina. Species of Myrtaceae are important floristic and ecological elements of many forests in the tropics, with the greatest diversity found in Australia (capsularfruited species) and the Neotropics (fleshy-fruited species). Many species are widely cultivated and appreciated for their edible fruit, such as guava, Psidium species; jaboticaba, Myrciaria cauliflora (Mart.) O. Berg; pineapple guava, Acca sellowiana (O. Berg) Burrett; araçá-boi, Eugenia stipitata McVaugh; and Suriname cherry,

1

2

M YRTACEAE

Eugenia uniflora L. Others are grown for spice, such as allspice, Pimenta dioica (L.) Merr., and clove, Syzygium aromaticum (L.) Merr. & L.M. Perry, or for their ornamental value (Eucalyptus species, Callistemon species, Eugenia species, Myrtus communis L.) Eucalyptus species, widely grown around the world for their timber and gum, are being planted in northern Bolívar state, increasing the likelihood that they will escape and become part of the naturalized flora. Besides their generally resinous sap and capsular fruits, Eucalyptus species can be easily distinguished from most native Myrtaceae by their alternate leaves. While many fleshy-fruited species are unpalatable, none are known to be poisonous, and a fair number are locally harvested and consumed for their uniquely spicy flavor. Flowers are generally fragrant, insect pollinated, and frequently open en masse. Fleshy-fruited species are typically bird-dispersed; capsular-fruited species are typically wind-dispersed. Myrtaceae are often considered difficult to identify to genus and species. While there are serious problems with generic delimitation in the genera centered on Eugenia and Myrcia, many species (outside of several notorious complexes) can be readily distinguished. Subtribal affiliation (in the flora area, subtribe Myrciinae: Calyptranthes, Marlierea, Myrcia; subtribe Eugeniinae: Calycorectes, Eugenia, Myrcianthes, Myrciaria, Plinia, Pseudananomis, Siphoneugena, Syzygium; and subtribe Myrtinae: Blepharocalyx, Calycolpus, Campomanesia, Myrteola, Psidium, Ugni) is often easily distinguished by observing mature seeds as well as examining the structure of the inflorescence. The embryo characters that aid in generic and subtribal placement are readily observed after extracting a seed from the fruit and removing the testa. Useful characters for species delimitation are found in the flowers, particularly the calyx (number, cohesion, shape, indument), as well as the inflorescence (type, peduncle and pedicel length, indument), and the leaves (midvein prominence on upper surface, indument). Leaf shape, however, can be variable, even on a single plant. A distinct species of uncertain generic affinity is known from the lower Río Caroní in northern Bolívar and Lara state, but not included in this treatment. It has small leaves (1.5–3.5 cm long), convex midvenin, tetramerous flowers, foliaceous calyx lobes, and solitary axillary flowers with a Myrcioid embryo. It may be a species of Myrcia with extreme inflorescence reduction, but would not key out there because of the lack of an inflorescence axis. Representative collections are Díaz 2487 (SEL), Liesner & González 11091 (MO, SEL), Rosales et al. 614 (SEL).

Key to the Genera of Myrtaceae by Bruce K. Holst and Leslie Landrum 1. 1.

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

Inflorescence paniculate or dichasial ........................................................ 2 Inflorescence racemose (sometimes with the axis greatly reduced and the inflorescence appearing glomerate or fasciculate), cymose, corymbose, or flowers solitary .................................................................................. 9 Fruits many-seeded, the seed coat bony; inflorescences of solitary flowers or dichasial, not paniculate ..................................................... 14. Psidium Fruits 1- or 2-seeded, the seed coat membranous, chartaceous, or coriaceous; inflorescences various ................................................................. 3 Inflorescences dichasial, these rarely with irregular branching .............. 4 Inflorescences paniculate ........................................................................... 6

M YRTACEAE 3

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

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

11(9). 11. 12(11). 12. 13(12). 13. 14(12). 14. 15(11). 15. 16(15).

16.

17(16).

Calyx lobes persistent .............................................................. 9. Myrcianthes Calyx lobes deciduous after anthesis ........................................................ 5 Calyx closed in bud, the flowers 4-merous; embryo crescent-shaped, cotyledons tiny ......................................................................... 1. Blepharocalyx Calyx open in bud, the flowers 4- or 5-merous; embryo globose or oblong, cotyledons large, plano-convex ................................... 13. Pseudanamomis Calyx completely closed in bud .................................................................. 7 Calyx open in bud, the lobes evident, often rounded and overlapping each other ....................................................................................................... 8 Calyx circumscissile, falling as a cap-like calyptra ............. 4. Calyptranthes Calyx opening longitudinally, usually resulting in 4 irregular, nonoverlapping calyx lobes ................................................................... 7. Marlierea Petals free and falling separately; embryo with cotyledons membranous, folded, and hypocotyl well developed and eleongate ................... 8. Myrcia Petals connate and falling as a unit; embryo with thick fleshy cotyledons, separate or fused, and hypocotyl very short or indistinct ..................... ............................................................................. 16. Syzygium (S. cumini) Calyx, androecium, and free portion of hypanthium deciduous as a circumscissile unit after anthesis; midveins usually convex ................. 10 Flowers not as above, flower parts individually deciduous or persistent .............................................................................................................. 11 Hypanthium gradually widened above ovary, the flower bud usually obovoid; embryo of fused cotyledons ..................................... 10. Myrciaria Hypanthium constricted above ovary, the flower bud with outline of an hour-glass; embryo divided, with two plano-convex cotyledons ......................................................................................... 15. Siphoneugena Flowers solitary........................................................................................ 12 Flowers two or more per inflorescence .................................................... 15 Calyx closed in bud or with a terminal pore; plants shrubs or small trees of mid- to low-elevation areas ............................................................. 13 Calyx open in bud, the individual calyx lobes evident; plants ericoid shrubs of high altitudes (1900–2800 m) ............................................. 14 Fruits many-seeded, the seed coats bony ................................... 14. Psidium Fruits 1-seeded, the seed coat chartaceous .......... 7. Marlierea (M. uniflora) Flowers nodding; stamens not exceeding the corolla; anthers introrsely dehiscent, > 1/3 the length of the filaments ................................... 17. Ugni Flowers spreading or erect; stamens longer than the petals; anthers laterally dehiscent, < 1/3 the length of the filaments .............. 11. Myrteola Ovary 3- or more-locular; calyx lobes (4)5 .............................................. 16 Ovary 2-locular; calyx lobes 4 .................................................................. 18 Locules 4–8, the locular wall becoming a glandular, warty, false seed coat as the fruit matures; 0 or 1 seed developing per locule, the true seed coat a thin membrane, usually not evident in the mature fruit; mature leaves membranous to chartaceous; lateral veins broadly arching; no clear marginal vein evident ............................................ 5. Campomanesia Locules 2–5, the locular wall rarely glandular, never becoming a false seed coat; usually 2–many seeds developing in each locule, the seed coat hard; mature leaves usually coriaceous; lateral veins broadly arching to straight; a lateral vein often clearly evident ............................ 17 Seed coat lustrous, smooth, not covered with a thin layer of pulpy or

4

M YRTACEAE

crusty tissue, 1–5 cells thick at narrowest point; calyx tearing between the lobes or not at all at anthesis; peduncles often in 1–3 decussate pairs on very short axillary shoots ........................................ 2. Calycolpus 17. Seed coat dull or rough, not lustrous, covered with a thin layer of pulpy tissue when wet (or a glaze or crusty layer when dry), the hard portion of the seed coat about 9–30 cells thick at narrowest point; calyx tearing irregularly or between the lobes at anthesis; peduncles not normally in decussate pairs on very short axillary shoots ........................ 14. Psidium 18(15). Hypanthium prolonged beyond the summit of the ovary; calyx completely closed in bud or the lobes evident only as reduced terminal lobes .... 19 18. Hypanthium not prolonged beyond the summit of the ovary, the calyx lobes free or partially to rarely completely fused ............................... 20 19(18). Embryo with 2 distinct, plano-convex cotyledons; ovules many per locule .............................................................................................. 3. Calycorectes 19. Embryo of fused cotyledons, homogenous; ovules 2 per locule ...... 12. Plinia 20(18). Embryo of fused cotyledons; inflorescences racemose ................... 6. Eugenia 20. Embryo of free cotyledons; inflorescences paniculate, or if racemose, then the petals rose or reddish purple .......................................... 16. Syzygium 1. BLEPHAROCALYX O. Berg, Linnaea 27: 348 (in key). Jan l856 [l854]; 4l2. Feb l856. [Subtribe Myrtinae]. Marlieriopsis Kiaersk., Bot. Tidsskr. 17: 281. 1890. by Leslie Landrum Shrubs or trees to 30 m tall; trichomes unicellular, simple or dibrachiate, to ca. 1 mm long. Inflorescence normally a dichasium or panicle of up to 35 flowers. Flowers 4-merous. Calyx open or closed in bud and tearing into 4 separate lobes ca. 1.5– 2 mm long that are deciduous at about anthesis. Ovary bilocular; ovules 4–17 per locule, aggregated near the center of the septum, aligned in a few irregular rows. Fruit a globose berry. Seeds 1–15, reniform, the testa membranous; embryo more or less crescent-shaped (in the flora area), C-shaped, spiral, or nearly straight, the hypocotyl thickened, the cotyledons relatively small, at times scarcely detectable. New World tropics, from southeast Caribbean to Chile; 3 species, 1 in Venezuela. Blepharocalyx eggersii (Kiaersk.) Landrum, Fl. Neotrop. Monogr. 45: 121. 1986. —Marlieriopsis eggersii Kiaersk., Bot. Tidsskr. 17: 282. 1890. Shrub or buttressed tree to ca. 23 m tall; leaves lustrous on upper surface; calyx closed in bud; inflorescence to 35-flowered; fruit purple-black at maturity, 1- or 2-seeded. Premontane semideciduous to evergreen montane forests, 300–1600 m; Delta Amacuro (Serranía de Imataca), Bolívar (La Escalera above La Danta), Amazonas (Cerro Aracamuni). Nueva Esparta; Guadeloupe, St. Vincent, Guyana, Peru, Amazonian Brazil. ◆Fig. 1.

Fig. 1. Blepharocalyx eggersii

Calycolpus

5

2. CALYCOLPUS O. Berg, Linnaea 27: 378. 1856. [Subtribe Myrtinae]. by Leslie Landrum Shrubs or trees to 10(–15) m tall; trichomes whitish or yellowish, unicellular, simple (in the flora area) or dibrachiate, to ca. 1.5 mm long. Leaves persistent, coriaceous, the venation brochidodromous, with several pairs of nearly straight laterals that are united by a marginal vein that parallels the leaf margin. Inflorescence an axillary, solitary flower or a very short axillary bracteate shoot with 1–3 decussate pairs of flowers. Flowers 5-merous (less often 4-merous). Calyx lobes often flared, often with an apical appendage, the calyx fused beyond the ovary’s summit and tearing between the lobes at anthesis or the calyx lobes free; petals slightly fleshy, white or tinged with red, often drying brown; bracteoles usually small, ± triangular, caducous at about anthesis or in C. legrandii leafy and persistent. Stamens 35–270, folded centerward or ± erect in the bud; anthers somewhat to markedly elongate, with ca. 4–40 glands. Ovary 2–6-locular, the locular wall sometimes glandular; ovules usually 8–32, the placenta a U-shaped pad or mound of tissue or an essentially round peltate structure. Fruit subglobose. Seeds few to numerous, ± reniform, the central portion usually soft, the outer C-shaped portion with a smooth, hard, lustrous seed coat 1–4 cells thick, the surface cells rounded to elongate; embryo oily, whitish, C-shaped, the cotyledons reflexed or straight, less than 1/4 the length of the embryo. Southern Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil; ca. 12 species, 6 in Venezuela, 5 of these in flora area. The foliage of most of the species in the flora area turns dark brown or blackish upon drying. The flowers tend to be large and showy, and a few species are worthy of cultivation (Calycolpus calophyllus, C. goetheanus). The one species in Venezuela outside the flora area, Calycolpus moritzianus (O. Berg) Burret, is known from Andean Venezuela and Colombia. Key to the Species of Calycolpus 1. 1. 2(1).

2.

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

Hypanthium densely covered with trichomes, these sometimes minute .... 2 Hypanthium glabrous or sparsely pubescent basally ............................... 4 Leaves sessile to subsessile, the petiole 0–1 mm long; lateral veins prominent on lower surface of leaf; peduncles borne in the axils of bracts, clustered in bracteate shoots at the tips of branches ......... C. bolivarensis Leaves petiolate, the petiole 1–6 mm long; lateral veins indistinct to scarcely visible below; peduncles borne in the axils of leaves, not clustered in bracteate shoots at the tips of branches ................................. 3 Leaves mainly over 4 cm long; calyx lobes 4 or 5, 3–4 mm wide, linguiform, blunt ........... C. alternifolius × C. roraimensis? [see C. roraimensis] Leaves mainly < 4 cm long; calyx lobes 5, 1.5–3 mm wide, triangular, sharp. .................................................................................... C. alternifolius Calyx lobes broadly rounded to obtusely triangular, 1–2 mm long, as wide or wider than long, the inner surface covered with appressed trichomes, at least in proximal half; calyx tearing between lobes at anthesis ................................................................................. C. calophyllus

6

M YRTACEAE

4.

5(4).

5.

Calyx lobes elliptic, narrowly elliptic, or subovate, 3–10 mm long, mostly longer than wide, the inner surface glabrous or sparsely villose at the base only; calyx not tearing between lobes at anthesis ....................... 5 Calyx lobes 5, elliptic to ovate, 3–6 mm wide, 0.8–1.7 times as long as wide, entirely glabrous within; young twigs terete, not 4-angled ............................................................................................... C. goetheanus Calyx lobes 4 or 5, narrowly elliptic to linear, ca. 2 mm wide, ca. 3.5– 5 times as long as wide, sparsely villose basally within; young twigs terete or weakly 4-angled .................................................... C. roraimensis

Calycolpus alternifolius (Gleason) Landrum, Madroño 36: 10. 1989. —Myrtus alternifolia Gleason, Brittonia 3: 173. 1939. Shrub to 2 m tall, sometimes with trailing stems to 5 m long, abundantly villous; leaf margins revolute; flower buds pinkish. Rocky areas in savannas on tepuis, 1500–2500 m; Bolívar (Auyán-tepui, Macizo del Chimantá), Amazonas (Cerro Cuao, Cerro Sipapo). Brazil (Amazonas: Serra Aracá). ◆Fig. 5. Collections similar to Calycolpus alternifolius from Cerro Guaiquinima appear to be the result of hybridization with C. roraimensis.

Calycolpus calophyllus var. angustifolius Steyerm., Ann. Missouri Bot. Gard. 71: 330. 1984. Shrub or tree to 15 m tall; leaves coriaceous, typically obovate, drying black or dark brown, the secondary venation obscure. Along periodically flooded river banks and in

Calycolpus bolivarensis Landrum, Ann. Missouri Bot. Gard. 76: 930. 1989. Shrub ca. 1 m tall. Lowland scrub forests, 100–300 m; Bolívar (between Guaniamo and Perro de Agua, km 145 on San Juan de Manapiare road). Endemic. Calycolpus calophyllus (H.B.K.) O. Berg, Linnaea 27: 381. 1856. —Myrtus calophylla H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 133. 1823. —Guayabito.

Fig. 2. Calycolpus calophyllus

Calycolpus 7

Fig. 3. Calycolpus roraimensis

Fig. 4. Calycolpus goetheanus

8

M YRTACEAE

Fig. 5. Calycolpus alternifolius

savannas, granitic outcrops, 50–400 m; widespread in western Amazonas. Amazonian Colombia, Peru, and Brazil. ◆Fig. 2. The fruit of Calycolpus calophyllus is edible. Calycolpus goetheanus (DC.) O. Berg, Linnaea 27: 381. 1856. —Myrtus goetheana DC., Prodr. 3: 240. 1828. —Piriko-so-yek (Arekuna). Shrub or tree to 20 m tall; leaves usually sharply acuminate, the venation inconspicuous. Near streams and lakes, gallery forests, woods bordering savannas, 50–1300 m; widespread in Delta Amacuro and Bolívar. Guyana, Suriname, French Guiana, Brazil (Amazonas, Maranhão, Pará). ◆Fig. 4.

Calycolpus roraimensis Steyerm., Fieldiana, Bot. 28: 1009. 1957. Calycolpus roraimense Kausel, Lilloa 33(6): 98. 1971 [1972]. Shrub or tree to 3.5 m tall, the stems usually 4-angled; leaves caudate-acuminate. Along montane streams and evergreen forests, 700–1200 m; Bolívar (Cerro Guaiquinima, Gran Sabana). Guyana, Peru, possibly Brazil (Rondônia). ◆Fig. 3. Calycolpus roraimensis is closely related to C. revolutus (Schauer) O. Berg, a species mainly of Guyana. Additional studies are needed to establish whether they represent discrete species or extremes of a continuum. Calycolpus roraimensis appears to hybridize with C. alternifolius on Cerro Guaiquinima.

3. CALYCORECTES O. Berg, Linnaea 27: 317. 1856. [Subtribe Eugeniinae]. by Bruce K. Holst Shrubs or trees; twigs terete or compressed. Leaves variable in size, shape, and venation. Inflorescence usually short-decussate-racemose and appearing fasciculate, or a solitary flower, axillary or subterminal; bracts persistent. Flowers 4– 6-merous; hypanthium prolonged above the summit of the ovary; bracteoles usually narrow and often deciduous before anthesis. Calyx closed in bud or with 4 small

Calycorectes 9

lobes, opening by irregular, ± longitudinal splitting. Anthers typically elongated, oblong to linear. Ovary bilocular, 3–many ovules per locule. Fruit globose or ellipsoid. Seeds 1 or 2; embryo eugenioid, the cotyledons fused. West Indies, South America east of the Andes; 10–15 species, 1 in Venezuela. A genus of disparate elements, Calycorectes is distinguished from other eugenioid genera solely by the prolongation of the hypanthium above the summit of the ovary. Several species of Eugenia, particularly in the Eugenia feijoi O. Berg species complex, however, show slight to marked hypanthial prolongation. Because of this, and other significant morphological variation, Calycorectes is unlikely to be maintained in the future as currently circumbscribed. Nevertheless, the type species (C. grandifolius O. Berg) has affinities with the one flora area species and several others of the Guianas region, and further study is needed on these taxa. The only other species of Calycorectes previously recognized for Venezuela, C. yatuae McVaugh, is here placed under Eugenia, due to its close affinities with the Eugenia feijoi species complex.

Calycorectes enormis McVaugh, Mem. New York Bot. Gard. 18(2): 223. 1969. —Terciopelo. Tree 8–20 m tall, the bark smooth; young vegetative growth, flowers, and fruits with dense, dark brown indument; leaves large, 20–40 cm long, the upper surface lustrous, the marginal veins at the very edge of the blade; inflorescence 2- or 4-flowered; flowers sessile, 4-merous; bracteoles deciduous; fruits ellipsoid, to 3.5 cm long, 1-seeded. Premontane to montane semideciduous to evergreen forests, cloud forests, 200–600 m; Delta Amacuro (El Palmar, Serranía de Imataca), Bolívar (Serranía de Imataca). Endemic. ◆Fig. 6.

Fig. 6. Calycorectes enormis

10

M YRTACEAE

4. CALYPTRANTHES Sw., Prodr. 79. 1788, nom. cons. [Subtribe Myrciinae]. Chytraculia P. Browne, Civ. Nat. Hist. Jamaica 239. 1756. by Bruce K. Holst Shrubs or trees; stem branching predominantly bifurcate, branchlets terete or compressed, occasionally narrowly bialate with the wings terminating distally between the petiole bases; indument when present of simple or more commonly of dibrachiate trichomes. Leaves decussate. Inflorescence usually of paired panicles arising from opposite leaf axils at the lowest node of an abortive branch, or the branch prolonged and leafy and the paired panicles from the lowest node, sometimes subtended by small or foliaceous bracts. Calyx completely closed in bud, circumscissile, hypanthium prolonged above the ovary, the operculum usually attached at one side in anthesis, deciduous or occasionally persistent; petals 0, 2, 3(–5), small and inconspicuous and often attached to the operculum and falling with it. Stamens numerous, borne on the hypanthium; anthers globose or elliptic. Ovary 2-locular; ovules 2 per locule. Fruits globose, crowned by the free portion of the hypanthium and often the calyx. Seeds 1 or 2; embryo contortuplicate, seed coat shiny, chartaceous. New World tropics and subtropics; ca. 100 spp., ca. 20 in Venezuela, 12 of these in the flora area. Calyptranthes, while easily recognized in Central America and the Antilles by the paired inflorescences with closed, circumscissile calyx, intergrades with Myrcia (4 or 5 distinct calyx lobes in bud) and Marlierea (calyx closed in bud or nearly so and with irregular longitudinal tearing) in South America. Key to the Species of Calyptranthes 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(1).

5.

Midvein of leaf blades convex, plane, or with 2 convex ridges on upper surface ......................................................................................................... 2 Midvein of leaf blades concave or sulcate on upper surface ..................... 5 Leaf blades shallowly cordate at base, sessile or short- and stout-pedicellate, drying blackish ............................................................... C. nigrescens Leaves rounded to acute at base, noticeably pedicellate ......................... 3 Leaves abruptly acuminate-caudate at apex; petioles 1.5–2 cm long ............................................................................................ C. conduplicata Leaves acute to obtuse at apex, petioles < 1 cm long ............................... 4 Inflorescence axes glabrous; flower buds ca. 3 mm long; leaf blades (9–) 10–12(–17) cm long; fruits 10–15 mm diameter ........................ C. forsteri Inflorescence axes pubescent; flower buds 1.5–2 mm long; leaf blades 4– 8.5 cm long; fruits 5–8 mm diameter ..................................... C. multiflora Flowers glabrous; leaves and young stems glabrous or with inconspicuous pubescence; branchlets narrowly winged, the wing perpendicular to the plane of the petioles .................................................................... 6 Flowers pubescent, usually densely so or at least densely sericeous at the base; leaves and young stems usually with noticeable pubescence (except C. pulchella and C. ruiziana); branchlets narrowly winged, compressed, or terete ................................................................................... 8

Calyptranthes 11

6(5).

Leaf blades obtuse to rounded at base; flower buds 2.5–3 mm long ........................................................................................................ C. pullei 6. Leaf blades cuneate to acute at base; flower buds ≤ 2.5 mm long ............ 7 7(6). Flower buds 2–2.5 mm long, obovoid to globose, not constricted above ovary .............................................................................................. C. crebra 7. Flower buds 1–1.5 mm long, obpyriform, slightly constricted above ovary ....................................................................................................... C. lucida 8(5). Leaves acute to broadly obtuse or rounded at apex ................................. 9 8. Leaves acuminate to acuminate-caudate ................................................ 10 9(8). Leaves and stems mostly glabrous ............................................... C. pulchella 9. Leaves and stems (particularly young ones) densely ferruginoustomentose ................................................................................ C. clusiifolia 10(8). Branchlets narrowly winged; young leaf blades sparsely sericeous on lower surface, glabrescent; mature fruits glabrous or sparsely sericeous ........................................................................................... C. ruiziana 10. Branchlets terete or compressed, young leaf blades densely and usually permanently sericeous on lower surface; mature fruits sericeous ..... 11 11(10). Leaf blades 7–13 cm long ........................................................... C. fasciculata 11. Leaf blades 20–30 cm long ...................................................... C. macrophylla Calyptranthes clusiifolia O. Berg in Mart., Fl. Bras. 14(1): 39. 1857, “clusiaefolia.” Tree 3–4(–17) m tall, young vegetative parts and inflorescences densely ferruginous-tomentulose; branchlets with smooth bark; leaf blades impressed-punctate, ovate to elliptic. Montane humid forests, 1300– 1500 m; Amazonas (Río Yudi headwaters, Sierra Parima). Aragua, Distrito Federal, Táchira; Brazil (Minas Gerais). Calyptranthes clusiifolia is similar in leaf size and shape to the type specimen from Brazil, but mature flowers and fruit are not known from the two known flora area populations. Indument on Venezuelan collections is rust-colored and that of Brazilian populations yellow-tan. Calyptranthes conduplicata B. Holst, Selbyana 23: 137. 2002. Tree 6–10 m tall; leaf blades 15–20 cm long, drying reddish brown, midvein plane or with two convex ridges on upper surface, apex abruptly acuminate and conduplicate, petioles 1–2 cm long; inflorescence 7–13 cm long, the branches strigose, many-flowered; flowers pedicellate, cream. Evergreen lowland forests, 100–200 m; Amazonas (along San Carlos-Solano road, Caño San Miguel). Endemic.

Calyptranthes crebra McVaugh, Fieldiana, Bot. 29(3): 181. 1956. Small tree to 3 m tall; branchlets narrowly winged; leaves impressed-punctate on upper surface; inflorescences many-flowered. Scrub forest on white sand, 100–200 m; Amazonas (Piedra de Cocui). Amazonian Peru and Brazil. Calyptranthes fasciculata O. Berg, Linnaea 27: 31. 1855. Calyptranthes sericea Griseb., Fl. Brit. W. I. 233. 1860. Thin shrub or small tree 3–6 m tall; lower surface of leaf blades densely coppery brownor beige-pubescent; flowers cream or yellow; inflorescence short-pedunculate, the clustered fruits dull purple at maturity. Riverbanks, evergreen or semideciduous forests, 100–900 m; Bolívar (northern areas, Gran Sabana, upper Río Caura), Amazonas (Río Orinoco, Río Siapa). Distrito Federal, Falcón, Sucre; Lesser Antilles, Trinidad, Guyana, Suriname, French Guiana, Amazonian Brazil and Bolivia. ◆Fig. 7. Calyptranthes forsteri O. Berg, Linnaea 27: 23. 1855. Glabrous small tree or shrub 2–7 m tall; leaves usually broadly rounded to obtuse or cuneate at base, drying pale greenish; inflo-

12

M YRTACEAE

rescences glabrous; fruits large, 1–1.5 cm diameter, thick-walled. White-sand scrub, lowland evergreen forests, along rivers, 50–200 m; Delta Amacuro (Caño Arature, Serrania de Imataca), western Amazonas. Lesser Antilles, Guyana, Amazonian Ecuador and Brazil. ◆Fig. 8. Calyptranthes lucida DC., Prodr. 3: 258. 1828. Shrub or small tree to 5 m tall, mostly glabrous; young stems narrowly winged; leaves impressed-punctate on upper surface. Evergreen forests along rivers, 100–1000 m; Amazonas (Cerro Duida, Río Cunucunuma, Río Mawarinuma). Suriname, Brazil. A rheophytic form with narrow leaves and reduced inflorescences from Río Mawarinuma at the base of Sierra de la Neblina may represent a distinct taxon (e.g., Gentry & Stein 46931, MO, SEL, VEN). Calyptranthes macrophylla O. Berg in Mart., Fl. Bras 14(1): 45. 1857. Small tree 3–7(–12) m, leaf blades large, > 20 cm long, lower surface covered with appressed dibrachiate trichomes. River banks, lowland evergreen forests, 100–200 m; Amazonas (widespread). Amazonian Colombia, Peru, and Brazil. ◆Fig. 11. The large-leaved, petiolate Calyptranthes species of the Guianas and Amazonia are in need of study. Described species are C. macrophylla, C. speciosa Sagot, and C. speciosa var. gigantifolia (McVaugh) McVaugh, but judging by the material from Peru and Brazil identified as these, there are probably several more. Calyptranthes multiflora O. Berg in Mart., Fl. Bras. 14(1): 42. 1857. Calyptranthes florifera McVaugh, Mem. New York Bot. Gard. 18(2): 72. 1969. Shrub 3–5 m, or tree to 10(–15) m tall; leaves mostly elliptic, midvein convex on upper surface; fruits 5–8 mm diameter. Along rivers, flooded forests, 50–400 m; Delta Amacuro (Piacoa), northeastern Bolívar (Río Caura, Río Chaviripa, Río Nichare basin, Río Paragua), Amazonas (Cerro Moriche, Río Casiquiare, Río Negro, Río

Orinoco, Río Ventuari). Anzoátegui, Apure; Amazonian Colombia, Peru, and Brazil (Amazonas, Roraima, Rondônia), Bolivia. Calyptranthes nigrescens B. Holst, Selbyana 23: 138. 2002. —Ojo de pavón. Shrub or tree 4–12 m tall; leaf blades subcordate, drying blackish along with inflorescence, flower buds, and fruits; mature fruits purple-black to black, 1.5–2.5 cm diameter, thick-walled. Periodically flooded margins of blackwater rivers, 100–200 m; Amazonas (Caño Yagua, Maroa, Río Atacavi, Río Autana, Río Baría, Río Cuao, Río Guainía, Río Emoni, Río Negro, Río Pasimoni, Río Yatua). Colombia (Guainía), Brazil (Amazonas: upper Rio Negro). ◆Fig. 10. Calyptranthes nigrescens is a species that is somewhat intermediate with Marlierea because of the calyptra that occasionally irregularly tears, instead of the typical cleanly circumscissile calyx in Calyptranthes. The fruits are described as edible but astringent. Calyptranthes pulchella DC., Prodr. 3: 257. 1828. —Maikanpi-miyuyek (Arekuna). Shrub or small tree 1–9 m tall, branchlets narrowly winged; leaves obovate (-rhomboid), rounded at apex; inflorescence unbranched, (1)3-flowered, buds abruptly apiculate. Along streams, Bonnetia forests on tepui slopes and summits, 1100–1900 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Gran Sabana, Macizo del Chimantá [Acopántepui, Sarvén-tepui]), Amazonas (Sierra de la Neblina). Guyana, Suriname, Colombia, Brazil (Minas Gerais). ◆Fig. 12. A single collection from low elevations at Cacurí [ca. 100 m] in Amazonas has the same small, obovate (–elliptic), apically rounded leaves as C. pulchella, but the collection (Colchester 2403, MYF) is sterile and cannot be positively identified. Calyptranthes pulchella belongs to a difficult complex consisting of several distinct populations. The type of the species, from Minas Gerais in Brazil, has inflorescences with 5–9 flowers and obovoid, scarcely apiculate buds. Collections from Amazonian Peru

Calyptranthes 13

and Brazil often identified as this species seem to represent a distinct species; the leaves are elliptic, stem wings prominent and prolonged into a stipule-like appendage at the node, and the buds ellipsoid. The Guayana Highland material may represent yet another species. Calyptranthes pullei Burret ex Amshoff, Rec. Trav. Bot. Neerl. 42: 4. 1950. Colombia, Venezuela, Guyana, Suriname; 2 varieties, 1 in Venezuela. C. pullei var. immaculata McVaugh, Mem. New York Bot. Gard. 10(1): 78. 1958. Shrub or tree 2–10 m tall, glabrous, branchlets narrowly winged, leaves ovate, broadly elliptic, to obovate, obtuse to rounded at the base, panicles three times compound. Periodically flooded areas along

rivers, 100–500 m; Bolívar (Río Caura, Río Orinoco, Río Paragua), Amazonas (junction of Río Orinoco and Río Ventuari). Anzoátegui, Apure; Colombia (Bolívar). ◆Fig. 9. Calyptranthes ruiziana O. Berg, Linnaea 27: 22. 1855. Tree 5–15 m tall, branchlets scarcely winged or not; leaves ovate to elliptic, apex acuminate; flower buds appressed-pubescent at base; inflorescence branched to nearly spicate. Humid, disturbed or riparian forests, 100–300 m; Amazonas (Río Cataniapo, Río Negro, Río Orinoco near junction with Río Atabapo). Peru, Brazil (Amazonas). A single collection from Sierra de la Neblina at 1100 m elevation, Maguire et al. 42185 (NY) is completely glabrous except for the vegetative buds, but otherwise matches collections from lower elevations.

Fig. 7. Calyptranthes fasciculata

14

M YRTACEAE

Fig. 8. Calyptranthes forsteri

Fig. 9. Calyptranthes pullei var. immaculata

Fig. 10. Calyptranthes nigrescens

Calyptranthes 15

Fig. 11. Calyptranthes macrophylla

Fig. 12. Calyptranthes pulchella

16

M YRTACEAE

5. CAMPOMANESIA Ruiz & Pav., Fl. Peruv. Prodr. 72, pl. 13. 1794. [Subtribe Myrtinae]. by Leslie Landrum Shrubs < 1 m tall to trees to ca. 15 m tall; trichomes whitish, yellowish, or light reddish brown, unicellular, simple, to ca. 1 mm long. Leaves persistent or caducous in less than one year, membranous to stiffly coriaceous, the venation usually brochidodromous with broadly arching lateral veins, a marginal vein not clearly differentiated; tufts of trichomes often present in the axils of the main lateral veins on lower surface of leaf. Inflorescence a solitary axillary flower or a 3-flowered dichasium, the peduncles subtended by leaves or reduced bracts, sometimes aggregated on short bracteate shoots. Flowers 5-merous; calyx open (in the flora area) or closed to various degrees; bracteoles linear, narrowly elliptic, or lanceolate, usually deciduous at or before anthesis. Stamens 60–700, whitish, folded centerward in the closed bud; petals whitish, suborbicular or obovate, 3–30 mm long. Ovary (3)4–18-locular, the locular wall becoming glandular, submembranous to slightly woody in the mature fruit, serving as a false seed coat; ovules 4–20 per locule, normally arranged in 2 longitudinal rows, normally all or all but one aborting per locule. Fruit a globose berry to ca. 7.5 cm diameter. Seeds normally 1 per locule or none, often ca. 1–4 per fruit, the seed coat a thin membrane, often not detectable in the mature fruit; embryo coiled, oily, reddish, the hypocotyl thickened, the cotyledons relatively small. Tropical and subtropical South America, from Trinidad to southeastern Brazil (where most diverse); ca. 30 species, 2 in Venezuela, both in the flora area. Key to the Species of Campomanesia 1.

1.

Style 4–7 mm long; disk 3–3.5 mm diameter; calyx lobes 1.2–3.2 mm wide; ovary 4–6-locular; stamens usually < 100; peduncle 0.3–0.5 mm wide ......................................................................................... C. aromatica Style 11–15 mm long; disk 6–7 mm diameter; calyx lobes 3–9 mm wide; ovary 7- or 8-locular; stamens 200 or more; peduncle ca. 1 mm wide ............................................................................................... C. grandiflora

Campomanesia aromatica (Aubl.) Griseb., Fl. Brit. W. I. 242. 1860. —Psidium aromaticum Aubl., Hist. Pl. Guiane 485. 1775. Deciduous shrub or tree 2–20 m tall, flowering when leaves are immature; secondary and tertiary venation on upper surface of leaves impressed; fruits purple-black at maturity, ca. 1 cm diameter. Lowland semideciduous to evergreen forests, savannas, 50–300 m; northern Bolívar (between El Tigre and La Soledad, near Guaniamo, Río Nichare), Amazonas (near Puerto Ayacucho). Anzoátegui, Monagas, Sucre; Trinidad, Guyana, Suriname, French Guiana, coast of Brazil south to Bahia, Amazonian Bolivia. ◆Fig. 13. The pulp of the fruit is described as aromatic and sweet-tasting and is eaten by monkeys.

Campomanesia grandiflora (Aubl.) Sagot, Ann. Sci. Nat. Bot. ser. 6, 20: 182. 1885. —Psidium grandiflorum Aubl., Hist. Pl. Guiane 483, pl. 190. 1775. —Canjilón guayabero. Tree ca. 8–10 m tall; leaf blades membranous, the tertiary venation finely reticulate, perpendicular to arching secondary veins, lower surface with tufts of trichomes present in secondary vein axils; fruits yellow at maturity, ca. 2 cm diameter. Forests, along streams, in savannas, 50–100 m; Bolívar (Túriba), cultivated in Amazonas (San Fernando de Atabapo). Guyana, Suriname, French Guiana, Brazil (Amazonas, Bahia, Pará, Roraima). The fruits of Campomanesia grandiflora are juicy and edible.

Eugenia 17

Fig. 13. Campomanesia aromatica

6. EUGENIA L., Sp. Pl. 470. 1753. [Subtribe Eugeniinae]. Phyllocalyx O. Berg, Linnaea 27: 306. 1856. Stenocalyx O. Berg, Linnaea 27: 309. 1856. by Bruce K. Holst Shrubs or trees. Inflorescence an axillary bracteate shoot, usually racemose or greatly reduced and appearing glomerate or fasciculate. Flowers 4-merous, hypanthium not prolonged beyond ovary. Petals white or rarely pinkish or bluish; calyx lobes free or rarely partly to completely connate, the inner 2 usually longer than outer 2, occasionally subfoliaceous, persistent or rarely deciduous. Stamens numerous; filaments incurved in bud; anthers dorsifixed, opening by longitidinal slits. Fruit globose, oblong, or ellipsoid, usually crowned by the persistent calyx; flesh scant to abundant. Seeds 1–few; embryo homogenous, the radicle broadly adnate to the wholly connate cotyledons. Tropics and subtropics worldwide; ca. 500 species, ca. 80 in Venezuela, 46 of these in the flora area. Eugenia is by far the largest genus of Myrtaceae in the New World. It is narrowly characterized by having 4-merous flowers with free calyx lobes and an embryo with fused cotyledons. There is a tendency, however, in some Eugenia species complexes toward prolongation of the hypanthium and connation of the lobes, which blurs the distinction with related genera such as Calycorectes and Plinia. Eugenia stipitata McVaugh is cultivated in southern Amazonas state for its fruits, but does not appear to naturalize. It is distinguished from all other Eugenia

18

M YRTACEAE

species in the flora area by the pedicels being borne midway on the pedicel, instead of at the base of the hypanthium. Eugenia sp. B is not included in the key as it is only known from fruiting collections. See species entry (page 28) for additional information. Key to the Species of Eugenia 1. 1. 2(1). 2. 3(2). 3.

4(2). 4.

5(4).

5.

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

7. 8(6).

8.

9(1).

Inflorescence racemose, the main axis > 1 cm long .................................. 2 Inflorescence sessile to short-racemose, often appearing fasciculate or glomerate, the axis < 1 cm long ............................................................ 9 Inflorescence and hypanthium glabrous or with sparse, minute, appressed, light coppery trichomes ........................................................... 3 Inflorescence and hypanthium conspicuously pubescent ........................ 4 Upper surface of leaf midvein concave or sulcate; lower sterile portion of inflorescence axis 1/2 or less the length of the fertile portion ...... E. florida Upper surface of leaf midvein convex or elevated and slightly wrinkly upper surface; lower sterile portion of inflorescence axis equaling or longer than the fertile portion ........................................ E. longiracemosa Lower surface of leaf blades glabrous, the secondary and marginal veins usually evident ...................................................................................... 5 Lower surface of leaf blades appressed-pubescent, usually densely so on young leaves, with the secondary and marginal veins usually inconspicuous .................................................................................................. 6 Upper surface of leaf blades usually impressed-punctate and the secondary veins slightly engraved (use 10× magnification); inflorescence and flowers mostly glabrous or with minute light coppery appressed trichomes ....................................................................................... E. florida Upper surface of leaf blades not impressed-punctate nor with engraved secondary veins; inflorescence and hypanthium densely coppery sericeous .................................................................................... E. trinervia Bracteoles rounded or broadly obtuse at apex, glabrous or at least distally so; fruits longitudinally ribbed-tuberculate ................................. 7 Bracteoles acute to sharply acuminate, pubescent over entire surface; fruits smooth .......................................................................................... 8 Leaf blades spatulate-obovate, emarginate at apex, strongly revolute with the reflexed margins nearly touching; bracteoles ca. 1 mm long ............................................................................................... E. emarginata Leaf blades elliptic, obtuse to short-acuminate at apex, the margins plane or slightly revolute; bracteoles ca. 2 mm long ....................... E. pubescens Petioles 2–5 mm long; leaf blades strongly impressed-punctate on upper surface, usually > 3 times as long as wide, the lower surface glabrescent to permanently and densely gray to silvery sericeous, base acute to cuneate; flower buds 3–5 mm wide .......................................... E. biflora Petioles 10–14 mm long; leaf blades not or only slightly impressed-punctate on upper surface, 2–2.5 times as long as wide, the lower surface glabrescent, base rounded to obtuse; flower buds 6–8 mm wide ............................................................................................E. pachystachya Bracteoles linear or filiform and often equaling the flower bud in length, deciduous at or before anthesis and leaving a small, nearly circular

Eugenia 19

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

11.

12(11).

12.

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

15.

16(15). 16. 17(9).

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

scar at the base of the hypanthium .................................................... 10 Bracteoles ovate to broadly ovate, persistent past anthesis and often into fruit ...................................................................................................... 17 Plants thinly pubescent to mostly glabrous; fruits red and pendent at maturity (purple-black in E. ligustrina) .................................................. 11 Plants evidently and often abundantly pubescent, at least on the inflorescence and often also on the leaves and stems .................................... 14 Calyx lobes completely glabrous; bracts at base of pedicels foliaceous (> 6 mm long); fruits purple-black at maturity; plants evergreen ................................................................................................. E. ligustrina Calyx lobes ciliate and often lightly pubescent; bracts at base of pedicels nonfoliaceous (< 3 mm long); fruits red at maturity; plants deciduous to semideciduous .................................................................................. 12 Leaves predominantly widest above the middle (mostly obovate), the midvein on the upper surface convex or elevated in a narrow line ...................................................................................................... E. patrisii Leaves predominantly widest at or below the middle (elliptic, oblong, ovate), the midvein concave or deeply sulcate on the upper surface, marginal vein mostly parallel to margin or only shallowly arching between the secondary veins .................................................................. 13 Calyx lobes oblong, thinly pubescent without, glabrous within; leaf blades with varying-sized gland dots ....................................... E. cribrata Calyx lobes rounded, ciliate but otherwise glabrous; leaf blades with uniform-sized gland dots ......................................................... E. roseiflora Pubescence bright coppery; leaf blades 5–7 cm long .............. E. callichroma Pubescence whitish or grayish or pale coppery; leaf blades mostly > 7 cm long ....................................................................................................... 15 Petioles slender, ca. 1 mm thick; midvein of leaf blades sulcate on upper surface, the secondary venation inconspicuous; calyx lobes subfoliaceous; fruits several-seeded, yellow to orange at maturity, conspicuously glandular ............................................................ E. macrocalyx Petioles stout, 2–4 mm thick; midvein of leaf blades plane to convex on upper surface; the secondary venation evident; calyx lobes not subfoliaceous; fruits 1-seeded, purplish at maturity, not glandular ....... 16 Calyx lobes 1.5–3 mm long, sparsely tomentulose to glabrous; fruits glabrescent; larger leaf blades < 20 cm long ........................ E. gomesiana Calyx lobes 4.5–8 mm long, permanently villous; fruits persistently tomentose; larger leaf blades > 20 cm long .................................... E. magna Inflorescence, or at least the hypanthium, densely pubescent (in one species, E. coffeifolia, the bracteoles obscuring the hypanthium must be removed to see the pubescence) .......................................................... 18 Inflorescence and flowers glabrous, and often the foliage as well, or, at most thinly pubescent with the trichomes not obscuring surfaces ............ 30 Midvein on upper surface of leaves impressed, usually V- or U-shaped in cross section ......................................................................................... 19 Midvein on upper surface of leaves plane or elevated in some fashion (convex, biconvex, or in a narrow elevated line or ridge) .................. 21 Leaf blades rounded to broadly obtuse at each end ............. E. cachoeirensis Leaf blades gradually to abruptly narrowed at one or both ends .......... 20

20

M YRTACEAE

20(19). Leaf blades 5–10(–13) cm long, secondary venation usually slightly impressed on upper surface; ovary and fruits not ribbed; flower buds ca. 1 mm long ................................................................................. E. coffeifolia 20. Leaf blades 15–20 cm long; secondary venation slightly elevated on upper surface; ovary and fruits longitudinally ribbed ...................... E. cuaoensis 21(18). Midvein on upper surface elevated in a narrow line or ridge; calyx lobes partially to completely connate; anthers linear, drying grayish ....... 22 21. Midvein on upper surface convex or biconvex; calyx lobes free; anthers not as above .......................................................................................... 24 22(21). Calyx completely closed in bud; secondary venation of leaf blade impressed on upper surface, prominently elevated on the lower ................. E. yatuae 22. Calyx open in bud; secondary venation of leaf blade plane on upper surface, usually inconspicuous, elevated or not on lower surface .......... 23 23(22). Ovary and fruit longitudinally ribbed; leaf blades thick-coriaceous, the venation inconspicuous on both surfaces; fruit 3–3.5 cm long ............................................................................................. E. diplocampta 23. Ovary and fruit smooth; leaf blades coriaceous, the venation usually conspicuous at least on lower surface; fruits 3–5.5 cm long ................ E. feijoi 24(21). Leaf blades 20–45 cm long, bearing large gland dots on the lower surface that are easily visible without magnification, petioles thick, corkyexfoliating ............................................................................ E. tumulescens 24. Leaf blades mostly < 20 cm long, the gland dots on the lower surface visible only with magnification; petioles smooth or wrinkled, but not as above .................................................................................................... 25 25(24). Flowers large, ca. 5 cm diameter when open ............................ E. moritziana 25. Flowers small to medium-sized, < 3 cm diameter when open ............... 26 26(25). Indument of matted, flexous trichomes; fruits red to orange or yellow at maturity ............................................................................................... 27 26. Indument of appressed, mostly straight trichomes; fruits dark purple to black at maturity ................................................................................. 28 27(26). Midvein in dried leaves definitely convex above, smooth or somewhat wrinkled; bracts, bracteoles, pedicels, and calyx lobes at least at base, cottony-tomentose ............................................................... E. ferreiraeana 27. Midvein flat or concave above, often finely wrinkled; lower surface of leaves and hypanthium floccose-tomentose, the bracteoles and calyx lobes glabrous or pubescent at base only ................................... E. omissa 28(26). Calyx lobes pubescent within with stout reddish trichomes; leaves thickcoriaceous ........................................................................... E. kaieteurensis 28. Calyx lobes glabrous within; leaves membranaceous to thinly coriaceous .............................................................................................................. 29 29(28). Marginal veins 2, the innermost (4–)5–10 mm from the margin and arching between the secondaries, particularly in the upper 1/3 of the blade, midvein prominently convex especially near the petiole; leaf blades 7– 11 cm wide and typically > 15 cm long ........................................ E. baileyi 29. Marginal vein 1, 1–3 mm from the margin and not arching between the secondaries, midvein plane or biconvex; leaf blades 3–5 cm wide and typically < 15 cm long ..................................................... E. chrysophyllum 30(17). Leaf blades drying yellow-green; midvein convex on upper surface; inflorescences mostly borne at leafless nodes, flowers pedicellate; fruits

Eugenia 21

depressed-globose ................................................................... E. flavescens 30. Plants not as above .................................................................................. 31 31(30). Larger calyx lobes 3.5–7 mm long (deciduous at anthesis in E. umbonata) .............................................................................................................. 32 31. Larger calyx lobes ≤ 3 mm long, persistent ............................................ 35 32(31). Calyx lobes deciduous at anthesis; leaves strongly arcuate-conduplicate ................................................................................................. E. umbonata 32. Calyx lobes persistent into fruit; leaves plane or arcuate-conduplicate only at apex .......................................................................................... 33 33(32). Midvein on upper surface of leaves impressed, usually V- or U-shaped in cross section; principal marginal vein 1–3 mm from the margin; fruits red at maturity ............................................................... E. pseudopsidium 33. Midvein on upper surface of leaves convex, biconvex, or plane; principal marginal vein 4–9 mm from the margin; fruits dark purple at maturity or unknown .......................................................................................... 34 34(33). Calyx lobes free, bracteoles connate; petals often with bluish tinge .......................................................................................... E. anastomosans 34. Calyx lobes connate to well above their base, bracteoles free; petals white ............................................................................................ E. marlierioides 35(31). Leaf blades broadly obtuse to rounded at apex ...................................... 36 35. Leaf blades acute to acuminate at apex .................................................. 41 36(35). Leaf blades obovate, coriaceous, upper surface lustrous; inflorescences reduced to one or typically 2 flowers borne in the leaf axils; fruit maturing from green to yellow to bright red at maturity; pubescence of young vegetative growth pale reddish .................................. E. punicifolia 36. Plants not as above .................................................................................. 37 37(36). Leaf blades drying yellowish green on upper surface, slightly arcuateconduplicate; pedicels short, stout, ca. 2 mm long .................. E. citrifolia 37. Leaf blades not drying as above, plane; pedicels > 2 mm long, usually slender .................................................................................................. 38 38(37). Pedicels glabrous; midvein on upper surface of leaf blades biconvex to plane ................................................................................................. E. sp. A 38. Pedicels pubescent; midvein on upper surface of leaf blades impressed, V- or U-shaped in cross section ........................................................... 39 39(38). Pedicels with appressed trichomes; leaf margins revolute ....... E. tepuiensis 39. Pedicels with erect trichomes; leaf blades plane .................................... 40 40(39). Leaf blades plane, membranaceous ..................................... E. amblyosepala 40. Leaf blades slightly to prominently arcuate-conduplicate, coriaceous ........................................................................................... E. tapacumensis 41(35). Midvein on upper surface of leaf blades impressed, U-shaped in cross section ................................................................................. E. lambertiana 41. Midvein on upper surface of leaf blades plane, biconvex, or convex ..... 42 42(41). Leaf blades drying light green on upper surface, lower surface olive brown; flowers nearly sessile ................................................ E. dittocrepis 42. Leaf blades not drying as above; flowers distinctly pedicellate ............. 43 43(42). Leaf blades broadly ovate to broadly oblong, 5–8 cm wide, < 1.8 times longer than wide, thick-coriaceous and usually lustrous on upper surface ................................................................................ E. conduplicata 43. Leaf blades ovate to lanceolate or narrowly elliptic, 1.5–5 cm wide,

22

M YRTACEAE

2 times or more longer than wide; membranaceous to coriaceous, lustrous or not on upper surface .............................................................. 44 44(43). Leaf blades chartaceous, midvein on upper surface persistently puberulent as is the inflorescence ..................................................... E. monticola 44. Leaf blades membranaceous to coriaceous, midvein on upper surface glabrous, the inflorescence glabrous to minutely puberulent ................ 45 45(44). Inflorescence axis evident, 3–10 mm long, somewhat square in cross section; pedicels mostly > 1 cm long; leaf blades usually > 7 cm long and gradually acuminate, venation on lower surface of leaves usually evident .......................................................................................... E. egensis 45. Inflorescence axis not evident or to 2 mm long; pedicels mostly < 1 cm long; leaf blades usually < 6 cm long and abruptly and prominently acuminate, venation on lower surface of leaves usually obscured .................................................................................................... E. protenta Eugenia amblyosepala McVaugh, Mem. New York Bot. Gard. 18: 162. 1969. Shrub 1–2 m tall or tree 5–10 m; leaves thin, obtuse to rounded at apex. Periodically flooded forests along rivers, ca. 100 m; Bolívar and Amazonas along the middle Río Orinoco and tributaries, from lower Río Caura to Puerto Ayacucho. Anzoátegui, Apure. ◆Fig. 14. Eugenia amblyosepala is closely related to and possibly conspecific with E. tapacumensis McVaugh. Eugenia anastomosans DC., Prodr. 3: 269. 1828. Eugenia ptariensis Steyerm., Fieldiana, Bot. 28: 1013. 1957. Slender, glabrous tree to 13(–19) m tall; leaf blades elliptic to obovate, obtuse or rounded at apex to shortly and bluntly acuminate; flowers tend to be borne on old wood, calyx lobes relatively large, petals white, turning light blue with age; fruit oblong. Lowland to upland forests in understory, edges of savannas, tepui escarpments, 100–1300 m; scattered in Delta Amacuro, Bolívar, and Amazonas. Sucre; Guyana, Suriname, French Guiana, Amazonian Ecuador, Brazil, and Bolivia. ◆Fig. 15. The fruit of Eugenia anastomosans is reported to be edible though astringent. Eugenia baileyi Britton, Bull. Torrey Bot. Club 48: 334. 1922, “bailleyi.” Eugenia peregrina McVaugh, Mem. New York Bot. Gard. 18(2): 201. 1969. Tree 4–7 m tall; leaves chartaceous, lower surface coppery to eventually silvery-seri-

ceous; midvein prominently convex on upper surface. Lowland evergreen forests to riparian forests on tepui summits and slopes, 200–1700 m; Delta Amacuro (Serranía de Imataca), Bolívar (Auyán-tepui, Río Acanán, Río Botanamo, Río Pao, Serranía de Imataca, near Upata), Amazonas (Río Coro Coro west of Cerro Yutajé, lower Río Ventuari). Trinidad, Suriname. ◆Fig. 16. See comments under Eugenia chrysophyllum. Eugenia biflora (L.) DC., Prodr. 3: 276. 1828. —Myrtus biflora L., Syst. Nat. ed. 10. 1056. 1759. —Arichai (Arekuna). Eugenia caurensis Steyerm., Fieldiana, Bot. 28: 1010. 1957. Shrub 0.3–3 m tall or tree to 5 m; lower surface of leaves and inflorescence gray to silvery-sericeous; leaves variable in shape, coriaceous, narrowly lanceolate to narrowly elliptic, less commonly ovate, short-mucronate, impressed-punctate on upper surface. Seasonally flooded forested, sandy, or rocky river banks, rocky savannas, shrub islands in savannas, 200–600 m; Amazonas (scattered), more common in Bolívar in all major river basins (except the Cuyuní). Apure, Barinas, Carabobo, Guárico, Lara, Mérida, Miranda, Portuguesa, Táchira, Trujillo, Zulia; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 23. Eugenia cachoeirensis O. Berg in Mart., Fl. Bras. 14(1): 265. 1857. Eugenia lingua O. Berg in Mart., Fl. Bras. 14(1): 266. 1857.

Eugenia

Small tree 3–8 m tall; leaves lustrous on upper surface, coriaceous, drying dark brown; inflorescences sessile, dense, hypanthium coppery brown sericeous; fruits depressed-globose. Periodically flooded blackwater rivers or lagoons, ca. 100 m; Amazonas (Río Casiquiare, Río Negro, Río Yatua). Adjacent Colombia and Brazil (Amazonas: Rio Negro). ◆Fig. 25. Eugenia callichroma McVaugh, Mem. New York Bot. Gard. 18(2): 169. 1969. Shrub 2–5 m tall; pubescence bright copper-colored; bracteoles deciduous; calyx lobes hooded. Habitat unknown; 100–300 m; Bolívar (Cerro La Puerta along middle Río Orinoco). Endemic. Eugenia chrysophyllum Poir. in Lam., Encyc. suppl. 3: 129. 1813. Eugenia llewelynii Steyerm., Fieldiana, Bot. 28: 1011. 1957. Shrub or slender tree 3–7 m tall; young vegetative growth with dense ferruginous trichomes; leaves variable in size, but typically 8–13 × 3–6 cm, upper surface of leaves glabrous, lower surface of leaves sericeous; calyx lobes in unequal pairs; fruit grayish when dry. Moist or wet areas along gallery forests, periodically flooded forests, savanna edges, 50–300(–500) m; Bolívar (Río Acanan, Río Parguaza, Río Pargueni), western Amazonas (Puerto Ayacucho to San Carlos de Río Negro and Río Baría). Apure, Mérida(?); Panama, Colombia, Suriname, Peru, Brazil (Roraima). ◆Fig. 18. Eugenia chrysophyllum is perhaps the oldest name in a complex of species centered on the Guayana Shield that includes at least E. baileyi, E. heterochroma, and E. kaiteurensis. The complex, characterized by lustrous sericeous pubescence on the lower surface of the leaf blades, plane to convex midvein, short-racemose, many-flowered racemes, and ellipsoid fruits, is in need of further study to determine species limits. Characters used in the key to the species and the synonyms listed under the above species should be considered as tentative. Eugenia citrifolia Poir in Lam., Encycl. suppl. 3: 129. 1813. Shrub to 2 m tall; leaves slightly arcuateconduplicate, drying yellowish green on upper surface, midvein on uper surface convex;

23

flowers nearly sessile at leafless nodes. On granitic outcrops, ca. 100 m; Bolívar (Cerro San Borja). Suriname, French Guiana. Eugenia citrifolia is an apparantly rare and poorly understood species. Eugenia coffeifolia DC., Prodr. 3: 272. 1828. Eugenia melinonis Sagot, Ann. Sci. Nat. Bot. sér. 6, 20: 194. 1885. Tree 3–20 m tall; leaf blades membranaceous, secondary venation usually slightly impressed on upper surface; flowers minute, the hypanthium completely hidden by the bracteoles. Semideciduous forests, 200–500 m; Delta Amacuro (Serranía de Imataca), Bolívar (Serranía de Imataca). Carabobo, Yaracuy; Lesser Antilles, Guyana, Suriname, French Guiana, Brazil (Amapá, Maranhão, Pará). ◆Fig. 21. A population from the lower Río Caura in Bolívar (Knab-Vispo & G. Rodriguez 334, SEL; Knab-Vispo & A. Rodriguez 707, SEL) and another from lower Río Ventuari in Amazonas (Liesner 18757, MO, SEL) with larger flowers that have the hypanthium partially visible and the secondary leaf venation convex on the upper surface seems to be related to E. coffeifolia, but may represent a new species. Eugenia conduplicata B. Holst, Selbyana 23: 140. 2002. Treelet; leaves thick-coriacous, relatively broad, conduplicate toward apex, upper surface lustrous; inflorescence few-flowered, pedicels slender, 1–3 cm long; fruit reddish. Riparian floodplain, ca. 200 m; Amazonas (Caño Yagua). Brazil (Amazonas: Rio Urubu). Eugenia cribrata McVaugh, Mem. New York Bot. Gard. 18(2): 174. 1969. —Pendanga, Ya’ra worokopinyo (Panare). Shrub 2–4 m tall or tree to 10 m; bark smooth; leaves deciduous or semideciduous; fruit bright red and juicy when mature. Rocky river banks, periodically flooded forests or savannas, rarely on dry forested slopes, 50–200(–400) m; Delta Amacuro (vicinity Los Castillos), Bolívar (along Río Orinoco and lower portions of its main tributaries, Río Cuyuní basin). Anzoátegui, Apure, Aragua, Barinas, Guárico, Lara, Monagas, Portuguesa, Zulia; Colombia (Macarena), possibly Guyana. ◆Fig. 27.

24

M YRTACEAE

The fruit of Eugenia cribrata is said to have a good taste and is eaten raw or in preserves. Eugenia cuaoensis McVaugh, Mem. New York Bot. Gard. 18(2): 174. 1969. Tree to 6 m tall; upper surface of leaves lustrous; sepals persistent; ovary and fruit densely and minutely puberulent, longitudinally ridged. Evergreen lowland forests; 100–200 m; Amazonas (Río Cataniapo, Río Cuao). Miranda. Eugenia diplocampta Diels, Verh. Bot. Vereins Prov. Brandenburg 48: 191. 1907. Small tree 3 m tall; leaves coriaceous, secondary venation inconspicuous; fruit longitudinally ridged. Lowland evergreen sclerophyllous forests, 100–200 m; Amazonas (near San Carlos de Río Negro). Amazonian Brazil. Eugenia dittocrepis O. Berg in Mart., Fl. Bras. 14(1): 292. 1857. Eugenia congestissima Diels, Verh. Bot. Vereins Prov. Brandenburg 48: 190. 1907. Small tree; leaf blades medium to large, 9–16 cm long, abruptly acuminate, coriaceous, midvein convex; flowers pale lilac. Flooded forests along rivers, ca. 100 m; Amazonas (Río Yatua). Peru, Brazil (Amazonas, Pará), Bolivia. ◆Fig. 24. Eugenia egensis DC., Prodr. 3: 281. 1828. Shrub or small tree 2–5(–8) m tall; upper surface of leaf blades frequently drying darker than lower surface; raceme axis square in cross section; pedicels slender. Riparian forests, river banks, rarely in shrubby savannas; 100–500(–800) m; Bolívar (basins of Río Caura, Río Cuyuní, and Río Paragua), Amazonas (widespread). Mérida(?); Costa Rica, Panama, eastern Colombia, Guyana, Suriname, Peru, Amazonian Brazil, Bolivia, Paraguay. ◆Fig. 26. Several populations with differing leaf shapes seem referable to Eugenia egensis, but may also represent new taxa (e.g., Huber 8477, NY; Gran Sabana, with cordate leaves). Eugenia emarginata (H.B.K.) DC., Prodr. 3: 280. 1828. —Myrtus emarginata H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 142. 1823.

Shrub 1–2 m tall; leaves coriaceous, lower surface sericeous; fruits ridged, grayish. Rocky areas along rivers, granitic outcrops, ca. 100 m; Amazonas (near Puerto Ayacucho). Endemic. ◆Fig. 28. Eugenia emarginata may be only a variety of E. pubescens, known also from the middle Río Orinoco. It is curious that this taxon has not been recollected in more than 40 years, even though the area that it is known from has been relatively well sampled during that time. Eugenia feijoi O. Berg in Mart., Fl. Bras. 14(1): 283. 1857. Shrub or tree to 18 m tall; leaf blades coriaceous, often drying blackish, midvein in a narrow line on upper surface; inflorescence few-flowered; calyx lobes partially connate; anthers grayish. Evergreen lowland forests, along rivers, 50–300 m; Delta Amacuro (scattered), northern Bolivar, Amazonas (widespread). Colombia, Ecuador, Peru, Brazil (Amazonas, Pará, Rondônia). ◆Fig. 30. Eugenia ferreiraeana O. Berg in Mart., Fl. Bras. 14(1): 285. 1828. Eugenia ramiflora var. montana Amshoff, Bull. Torrey Bot. Club 75: 535. 1948. Shrub or slender tree 2–6 m tall; young leaves and stems red-brown-floccose; leaves variable in shape, forest and laja forms ovate to broadly elliptic, riparian forms narrowly elliptic; fruits depressed-globose, yellow or orangish, covered with gray-brown matted trichomes. Along rivers, slope forests, rock outcrops, 100–500 m; Bolívar (near Cerro Camarón, Cerro Ichún, Río Caura), Amazonas (central to southern regions), central to southern Amazonas. Guyana, Suriname, Brazil (Amazonas, Pará). Eugenia flavescens DC., Prodr. 3: 272. 1828. Eugenia flavescens var. guianensis Sagot, Ann. Sci. Nat. Bot. sér 6, 20: 192. 1885. Glabrous shrub or treelet to 3 m tall; leaves, flowers, and fruits drying yellowgreen; midvein convex, new leaves very thin, eventually coriaceous; fruit depressed-globose. Lowland semideciduous forests, 100– 300 m; northern Bolívar. Miranda, Sucre, Táchira, Zulia; Guyana, Suriname, French Guiana, Brazil, Bolivia.

Eugenia 25

Eugenia florida DC., Prodr. 3: 283. 1828. —Ipiya (Yeral), Piitirijrida (Curripaco). Shrub or tree to 8(–12) m tall; leaves variable in size and thickness depending on habitat, upper surface usually impressed-punctate and with the secondary veins delicately engraved; inflorescence typically glabrous, but can be sparsely to densely sericeous, racemose or occasionally a panicle of racemes; mature fruit purple-black. Found in a wide array of natural and disturbed habitats, but usually in lowland to upland moist evergreen forests or along rivers, 50–1000 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Apure, Barinas, Falcón, Guárico, Lara; Costa Rica, Colombia, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 22. Eugenia florida is treated here in the broad sense, but several rather distinct populations occur in the flora area. A close study of this wide-ranging species complex is needed. Eugenia gomesiana O. Berg in Mart., Fl. Bras. 14(1): 254. 1857. Eugenia prosoneura O. Berg, Linnaea 31: 255. 1862. Shrub or tree 3–5 m tall; leaf blades mostly < 20 cm long, usually glabrescent, occasionally permanently pilose; inflorescences tomentulose; sepals erect or incurved in fruit, the apices often glabrous; fruits ellipsoid, glabrescent. Periodically flooded forests along black-water rivers, 100–200 m; southwestern Amazonas. Amazonian Peru, Brazil, and Bolivia. Eugenia kaieteurensis Amshoff, Bull. Torrey Bot. Club 75: 536. 1948. Shrub or more commonly a tree 8–12 m tall; pubescence ferruginous when young, turning grayish with age; leaves thick-coriaceous, obovate; inflorescence rachis to 5 mm long; fruits orange when mature, oblong-ellipsoid. Humid montane forests, 1200–1600 m; Bolívar (vicinity La Escalera-Cerro Venamo, between Ptari-tepui and Sororopántepui). Guyana (Kaieteur Plateau). Eugenia lambertiana DC., Prodr. 3: 270. 1828. Lesser Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru,

Amazonian Brazil, Bolivia; 2 varieties, 1 in Venezuela. Eugenia lambertiana var. hispidula McVaugh, is known from Suriname and Pará, Brazil. E. lambertiana var. lambertiana Eugenia schomburgkii O. Berg, J. Bot. (Hooker) 2: 321. 1840. Shrub or week-stemmed treelet 2–4 m tall or tree 5–15(–20) m, mostly glabrous; leaves variable in size, typically ovate to lanceolate, acuminate; flowers fragrant; fruits globose, purple-black at maturity. Occurring in a wide variety of habitats: tall primary forests, open or closed riparian forests, seasonally flooded forests, forest-savanna borders, shrubby thickets, 100–500 m; Delta Amacuro (Serranía de Imataca), northern Bolívar, Amazonas (widespread under 200 m). Other distribution as in species. ◆Fig. 29. Eugenia ligustrina (Sw.) Willd., Sp. 962. 1800. —Myrtus ligustrina Prodr. 78. 1788. West Indies, Venezuela, Guyana, name, Brazil, Bolivia; 2 varieties, Venezuela.

Pl. 2: Sw., Suri1 in

E. ligustrina var. ligustrina Glabrous shrub or more frequently a small tree, 3–7 m tall, bark smooth, thin, peeling in plates; leaves coriaceous, upper surface smooth and the venation obscure; inflorescence with several overlapping bracts basally and the pedicels subtended by a subfoliaceous bract. Understory of deciduous or semideciduous forests, rocky slopes, 100– 400 m; north-central and northeastern Bolívar. Falcón, Lara; West Indies, Guyana, Brazil (Minas Gerais, Rio de Janeiro, Santa Catarina), Bolivia. ◆Fig. 17. Eugenia longiracemosa Kiaersk., Enum. Myrt. Bras. 149. 1893. Tree to ca. 5 m tall, densely branched; leaves membranaceous, ovate to elliptic or obovate, the upper surface impressed-punctate; petioles 2–3 mm long. Along streams, ca. 100 m; Amazonas (Caño Ucata in middle Orinoco). Brazil (Amazonas: Rio Negro basin). Eugenia macrocalyx (Rusby) McVaugh, Fieldiana, Bot. 29: 212. 1956.

26

M YRTACEAE

—Calycorectes macrocalyx Rusby, Mem. New York Bot. Gard. 7: 313. 1927. Eugenia wentii Amshoff, Recueil Trav. Bot. Néerl. 39: 158. 1942. Tree to 10 m tall; leaves long and slender petiolate, the blades with inconspicuous venation, drying pale greenish, mostly glabrous; inflorescence and young shoots densely sericeous; fruits large, severalseeded. Upland evergreen forests, 900–1000 m; Bolivar (southern Gran Sabana). Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Amazonas, Pará), Bolivia. Eugenia magna B. Holst, Selbyana 23: 142. 2002. Tree 3–15 m tall; leaf blades mostly > 20 cm long, permanently velutinous, petioles stout; inflorescences densely villous; calyx lobes cupped. Upland forests, 700–900 m; Bolívar (southern Gran Sabana). Guyana, Brazil (Amazonas: Serra Aracá). Eugenia marlierioides Rusby, Bull. New York Bot. Gard. 8: 108. 1912. Shrub to 2 m or clambering over rocks; leaves 12–19 cm long, the margin thickened, midvein convex; inflorescence abbreviated, few-flowered; pedicles relatively long and slender; calyx lobes connate, thin, forming a globe much larger than the ovary; fruits probably large. Lowland evergreen forests, 100–200 m; Amazonas (near San Carlos de Río Negro). Amazonian Bolivia. Fruits are unknown from the sole Venezuelan collection, but immature fruits from a Bolivian collection (Daly et al. 6595, SEL) are described as 5 cm diameter and depressedglobose. Eugenia monticola (Sw.) DC., Prodr. 3: 275. 1828. —Myrtus monticola Sw., Prodr. 78. 1788. ?Eugenia monticola var. racemosa Amshoff in Pulle, Fl. Suriname 3(2): 121. 1951. ?Eugenia monticola var. latifolia Krug & Urb., Bot. Jahrb. Syst. 19: 636. 1895. Tree 3–10 m tall; leaf blades thin, the basal veins strongly ascending. Evergreen or semideciduous forests, 100–500 m; northcentral and northeastern Bolívar. Mexico, Central America, West Indies, Colombia,

Guyana, Suriname, Ecuador. ◆Fig. 19. Additional work on the amount of variability of Eugenia monticola needs to be done before the varieties can be recognized. Eugenia moritziana H. Karst., Auswahl Gew. Venez. 18, t. 6. 1848. Tree to 7 m tall; leaves coriaceous, upper surface midvein biconvex to convex, lower surface finely and obscurely sericeous; inflorescence appressed-velutinous; flowers large. Lowland evergreen forests, ca. 300 m; Bolívar (Guaniamo). Northern Venezuela (exactly locality uncertain), Colombia. Eugenia omissa McVaugh, Mem. New York Bot. Gard. 18(2): 197. 1969. Tree 4–10 m tall; indument of leaves, stems, and inflorescence pale orangish, turning gray with age; bracteoles connate at base; flowers fragrant, usually borne at leafless nodes. Evergreen forests, 100–1000 m; Bolívar (upper Río Caura), Amazonas (Caño Caname, Cerro Huachamacari, Río Asisa, Río Casiquiare, Río Guayapo). Suriname, French Guiana, Peru, Brazil (Acre, Amapá, Amazonas, Pará, Roraima), Bolivia. Eugenia omissa is closely related to E. ramiflora Desv. of the Guianas and Brazil (Pará, Rondônia), and it is possible that the two may represent the same species. Eugenia pachystachya McVaugh, Mem. New York Bot. Gard. 18(2): 197. 1969. Shrub or tree 1–7 m tall; leaf blades coriaceous, racemes occasionally bearing leaves; inflorescence densely gray- or gray-brown-tomentose; fruits round, smooth. Periodically flooded riparian forests, laja vegetation, ca. 50–100 m; Bolívar and Amazonas along Río Orinoco and lower portions of tributaries from Puerto Ordaz to confluence with Río Ventuari. Llanos (Anzoátegui, Apure). ◆Fig. 32. Eugenia patrisii Vahl, Eclog. Amer. 2: 35. 1798. —Stenocalyx patrisii (Vahl) O. Berg, Linnaea 29: 247. 1858. —Guayabo dantero, Majako shodö (Yekuana), Pendanga, Pendango. Shrub or tree 1–10(–18) m tall; leaf blades typically obovate, texture and size variable, basal veins strongly ascending and broadly

Eugenia

arching between the secondaries; flowers fragrant; mature fruit bright red, pendulous. Evergreen, riparian, periodically flooded, and secondary forests, forest-savanna borders, disturbed areas, 100–600 m; Delta Amacuro (Serranía de Imataca), Bolívar (scattered, but rare in Gran Sabana), Amazonas (widespread). Amazonian Colombia, Guyana, Suriname, French Guiana, Amazonian Peru, Brazil, and Bolivia. ◆Fig. 33. Eugenia protenta McVaugh, Mem. New York Bot. Gard. 18(2): 204. 1969. —Eugenia protracta O. Berg, Linnaea 31: 254. 1862, non Steud. 1843. Shrub or small tree to 5 m tall; leaves usually abruptly acuminate; fruits purple at maturity. Scrub forests, lowland evergreen forests, 100–1400 m; Bolívar (Altiplanicie de Nuria, Lago Guri, Río Paragua), Amazonas (Cerro Aracamuni, Cerro Duida, near San Carlos de Río Negro). Guyana (Kanuku Mountains), Brazil (Amazonas). ◆Fig. 39. Eugenia pseudopsidium Jacq., Enum. Syst. Pl. 23. 1760. Shrub or tree to 24 m tall; leaves variable in texture; inflorescences few–many-flowered, pedicels slender; fruit red at maturity. Semideciduous to evergreen lowland to montane forests, shrub savannas, 100–1400 m; Delta Amacuro (Río Acure, Serranía de Imataca), Bolivar (Cerro Venamo, Serranía de Imataca). Anzoátegui, Barinas, Distrito Federal, Mérida, Monagas; Puerto Rico, Guadeloupe, Martinique, Guyana, Suriname, French Guiana, Brazil (Acre, Amapá, Amazonas). Eugenia pubescens (H.B.K.) DC., Prodr. 3: 282. 1828. —Myrtus pubescens H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 143. 1823. —Paduari-yek (Arekuna). Eugenia ottonis O. Berg, Linnaea 27: 298. 1856. Shrub or tree to 9 m tall, abundantly grayto light-brown-pubescent; leaf blades coriaceous, the venation inconspicuous; bracteoles mostly glabrous, contrasting with the pubescent hypanthium; fruits longitudinally ridged-tuberculate, reddish when mature. Rocky outcrops along rivers, gallery forests,

27

premontane forests, 50–400 m; Bolívar (near Ciudad Bolívar, Río Caroní, Río Caura, Río Nichare, Río Paragua), Amazonas (middle Río Orinoco). Endemic. ◆Fig. 34. Eugenia punicifolia (H.B.K.) DC., Prodr. 3: 267. 1828. —Myrtus punicifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 149. 1823. Eugenia kunthiana DC., Prodr. 3: 264. 1828. Eugenia subalterna Benth., J. Bot. (Hooker) 2: 320. 1840. Eugenia benthami O. Berg, Linnaea 27: 164. 1856. Subshrub, shrub, or small tree, 0.3–4 m tall; leaves typically obovate, variable in size and indument; flowers 2 per node; fruits red at maturity. Savannas, shrub islands, gallery forests, granitic outrcops, lowland to montane evergreen forests, disturbed areas, 50–1700 m; widespread in Bolivar and Amazonas. Apure, Aragua, Carabobo, Falcón, Guárico, Lara, Mérida, Monagas, Sucre, Zulia; West Indies, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Africa. ◆Fig. 38. The red fruits of Eugenia punicifolia are edible, and the crushed plant is effective as an astringent for cuts. This commonly collected species is perhaps the only species of Myrtaceae that is naturally distributed in the Neotropics as well as Africa. Eugenia roseiflora McVaugh, Mem. New York Bot. Gard. 18(2): 210. 1969. Deciduous or semideciduous shrub or tree 2–13(–24) m, rarely scandent; flowers fragrant, pedicels to 4 cm long, slender; fruits bright red at maturity, pendent. Flooded forests, commonly along black-water rivers, rarely in semideciduous forests, 100–300 m; northwestern Bolívar, Amazonas (Caño Ucata southeast of Síquita, Río Atabapo, Río Atacavi, Río Baría, Río Guasacavi, Río Guayapo, Río Negro, middle to upper Río Orinoco, Río Pasimoni, Río Sipapo, Río Temi). Adjacent Colombia (Vichada), Peru(?), Brazil (upper Rio Negro), Bolivia(?). ◆Fig. 36. Eugenia tapacumensis O. Berg, Linnaea 27: 222. 1856. Eugenia ochra O. Berg, Linnaea 27: 216. 1856.

28

M YRTACEAE

?Eugenia roriamana O. Berg, Linnaea 27: 219. 1856. Shrub or tree to 10(–20) m tall; leaves often lustrous on upper surface; fruit purpleblack at maturity. Lowland evergreen or semideciduous forests, 100–300 m; Delta Amacuro (Serranía de Imataca), northern Bolivar. Apure; Panama, West Indies, Guyana, Suriname, Peru, Brazil (Amapa, Maranhao, Pará, Piauhi). ◆Fig. 31. Eugenia amblyosepala McVaugh is difficult to distinguish from E. tapacumensis. Eugenia tepuiensis Steyerm., Fieldiana, Bot. 28: 1015. 1957. Tree ca. 12 m tall; leaf blades coriaceous, with revolute margins. Tepui slope forests, 1500–1600 m; Bolívar (vicinities of Ptaritepui and Sororopán-tepui). Endemic. Eugenia trinervia Vahl, Eclog. Amer. 2: 36. 1798. Shrub or tree to 5 m tall; leaves thin, the principal marginal vein conspicuous and 5– 10 mm from the margin, a second lighter marginal vein present; inflorescence and particularly the hypanthium golden-sericeous; fruit to 2 cm diameter. Evergreen submontane or montane forests, 400–1400 Bolívar (near El Paují, Río Supamo, slopes of Macizo del Chimantá). Miranda; Lesser Antilles, French Guiana. Eugenia tumulescens McVaugh, Fieldiana, Bot. 29: 221. 1956. Understory tree 4–8 m tall; leaves with large, evident gland dots, the petioles corkyexfoliating; fruit yellow or orange-red at maturity. Evergreen lowland forests, along rivers, 100–200 m; Amazonas (Río Mawarinuma, near San Carlos de Río Negro). Brazil (Amazonas: Rio Negro basin). Eugenia umbonata McVaugh, Mem. New York Bot. Gard. 18(2): 215. 1969. Shrub or tree 2–8 m tall; leaf blades arcuate-conduplicate, ovate, 5–7 cm long; flowers borne at leafless nodes, relatively large, 6–8 mm wide in bud, calyx lobes deciduous at anthesis, an unusual feature in the genus; fruits globose, dark purple at maturity. Shrub islands around granitic outcrops, 100– 200(–800) m; Bolívar (Los Pijiguaos), Amazonas (near Puerto Ayacucho, Raudal de Atu-

res). Endemic. ◆Fig. 35. Eugenia yatuae (McVaugh) B. Holst, Selbyana 23: 144. 2002. —Calycorectes yatuae McVaugh, Mem. New York Bot. Gard. 18: 226. 1969. Tree 3–15 m tall; inflorescence, flowers, and young branches coppery-sericeous; leaf blades drying dark brown and lustrous on upper surface, secondary venation impressed on upper surface, markedly raised on lower surface; inflorescence borne at leafless nodes, calyx closed in bud; fruit orange-red at maturity. Periodically flooded evergreen lowland forests, 100–200 m; Amazonas (Río Mawarinuma, Río Yatua). Brazil (Amazonas: Rio Cauaburí). ◆Fig. 37. The fruit of Eugenia yatuae is reported to be edible and spicy. Eugenia sp. A Tree to 15 m tall; leaf blades coriaceous, lustrous on upper surface, broadly obtuse to rounded at apex, midvein shallowly impressed to shallowly convex on upper surface; inflorescence glabrous, pedicels slender. Lowland, evergreen forests, seasonally flooded forests, 50– 100 m; Amazonas (scattered). Possibly also Apure state and Amazonian Brazil and Bolivia. This taxon is well represented in herbaria, but sufficiently close to several other species (Eugenia lambertiana DC., E. egensis DC., E. pseudopsidium Jacq.) that a more thorough study is warranted before describing it as new. Representative collections are Williams 15123 (F, US) and Liesner 9057 (MO, NY). Eugenia sp. B. —Uraurek-boni-yek (Arekuna). Eugenia producta auct. non DC. 1828: sensu McVaugh, New York Bot. Gard. 18: 203. 1969. Tree 7–8 m tall; fruit ellipsoid. Forested slopes, 1100–1400 m; Bolívar (near Ptaritepui, Quebrada O-paru-má near Kavanayén). Endemic. ◆Fig. 20. The type of Eugenia producta has globose fruit, while the two collections of this taxon in Venezuela, Steyermark 60440 (F, NY, VEN) and 60609 (F), both have ellipsoid fruits. The leaves are identical with those of E. pseudopsidium, but that also has round fruits. Flowering collections should help solve this problem.

Eugenia 29

Fig. 14. Eugenia amblyosepala

Fig. 15. Eugenia anastamosans

30

M YRTACEAE

Fig. 16. Eugenia baileyi

Fig. 17. Eugenia ligustrina var. ligustrina

Eugenia 31

Fig. 18. Eugenia chrysophyllum

Fig. 19. Eugenia monticola

Fig. 20. Eugenia sp. B

32

M YRTACEAE

Fig. 21. Eugenia coffeifolia

Fig. 22. Eugenia florida

Fig. 23. Eugenia biflora

Eugenia 33

Fig. 24. Eugenia dittocrepis

Fig. 25. Eugenia cachoeirensis

34

M YRTACEAE

Fig. 26. Eugenia egensis

Fig. 27. Eugenia cribrata

Eugenia 35

Fig. 28. Eugenia emarginata

Fig. 29. Eugenia lambertiana var. lambertiana

36

M YRTACEAE

Fig. 30. Eugenia feijoi

Fig. 31. Eugenia tapacumensis

Eugenia 37

Fig. 32. Eugenia pachystachya

Fig. 33. Eugenia patrisii

38

M YRTACEAE

Fig. 34. Eugenia pubescens

Fig. 36. Eugenia roseiflora

Fig. 35. Eugenia umbonata

Eugenia 39

Fig. 37. Eugenia yatuae

40

M YRTACEAE

Fig. 38. Eugenia punicifolia

Fig. 39. Eugenia protenta

7. MARLIEREA Cambess. in A. St.-Hil., Fl. Bras. Merid. 2: 373 (folio ed. 269). pl. 156. 1829 [1830]. [Subtribe Myrciinae]. Krugia Urb., Ber. Deutsch. Bot. Ges. 11: 375. 1893. by Bruce K. Holst Shrubs or trees; branchlets terete, complanate, or shallowly winged, when winged, the wings of the stem terminate directly below the petioles and are sometimes continuous with them; plants glabrous to densely pubescent, the trichomes simple or conspicuously dibrachiate. Leaves typically petiolate; midvein sulcate to noticeably convex; marginal vein distinct. Inflorescences lateral to subterminal, paniculate, often paired in the axils, usually borne among the leaves, bracteate, the bracts early deciduous. Flowers 4- or 5-merous; hypanthium prolonged beyond the summit of the ovary; calyx completely closed in bud and apiculate or the lobes free, in either case tearing irregularly and longitudinally prior to anthesis, the lobes usually deciduous; petals present and conspicuous to much reduced. Stamens many, adnate to the hypanthium. Style elongate. Fruits globose, crowned by the persistent hypanthium. Seeds 1; embryo myrcioid. Panama, West Indies, South America east of the Andes (most diverse in the Guayana and Brazilian Shields); ca. 100 species, ca. 35 in Venezuela, 31 of these in the flora area. Marlierea is a poorly defined genus that is becoming increasingly difficult to maintain. It is distinguished entirely by a single character: the method of opening of the calyx (closed in bud or with small lobes and tearing longitudinally into the hypanthium). As more collections are studied, however, that is proving to be a variable character, at times variable even on the same individual.

Marlierea 41

At one extreme, some Marlierea species resemble Calyptranthes. In these cases, the inflorescence has paired panicles, and the calyx is closed in bud and roughly calyptrate (the edges of the hypanthium with 1–3 smaller lobes, but the major lobe containing the apiculum intact). Some examples of this are Marlierea schomburgkiana, M. cana, M. ligustrina, and M. suborbicularis. At the other extreme, the calyx is open, if only slightly so, and tears longitudinally into 4 or 5 lobes (e.g., M. rugosior, M. umbraticola) and these appear to be allied with some groups of Myrcia. Key to the Species of Marlierea 1. 1. 2(1).

2. 3(2). 3. 4(3). 4. 5(4). 5. 6(3). 6. 7(6). 7. 8(2). 8. 9(8). 9. 10(9). 10. 11(10).

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

Flower buds glabrous, the inflorescence glabrous or thinly pubescent ...... 2 Flower buds densely pubescent, often also the entire inflorescence ..... 15 Branchlets narrowly winged or sharply angled, the wing or angle in the same plane as the petiole and often appearing decurrent from the petiole ........................................................................................................... 3 Branchlets terete to compressed ............................................................... 8 Leaf blades to 7 cm long, the apex usually obtuse to rounded; petioles 1–3 mm long ........................................................................................... 4 Leaf blades > 7 cm long, the apex usually acute to acuminate, petioles > 2.5 mm long ........................................................................................ 6 Leaf blades subcordate to broadly rounded at base ............. M. lituatinervia Leaf blades narrowed to the base .............................................................. 5 Plants glabrous; leaf blades < 3 times longer than wide ....... M. karuaiensis Plants thinly pubescent; leaf blades > 3 times longer than wide .............. .................................................................................................. M. maguirei Petioles 5–6 mm long ............................................................ M. guildingiana Petioles 2.5–4 mm long .............................................................................. 7 Inflorescences to 9-flowered on each main branch, the axes slender to filiform; inflorescence and young shoots thinly pubescent ..... M. bipennis Inflorescences ca. 50-flowered on each main branch, the axes stout; plants glabrous ...................................................................... M. ensiformis Midvein of leaf blades deeply sulcate; inflorescence moderately graypubescent ............................................................................ M. umbraticola Midvein of leaf blade convex, biconvex, or plane; inflorescence mostly glabrous ....................................................................................................... 9 Leaf blades sharply acuminate to caudate at apex ................................ 10 Leaf blades acute, obtuse, or rounded at apex, or if acuminate, then bluntly and shortly so .......................................................................... 12 Calyx with a small opening in bud .......................................... M. scytophylla Calyx completely closed in bud ............................................................... 11 Midvein of leaf blade prominently convex; leaf blades mostly obovate, apex caudate-acuminate, lamina not impressed-punctate .................... ..................................................................................... M. schomburgkiana Midvein of leaf blade shallowly sulcate; leaf blades mostly ovate to elliptic, apex acuminate, lamina impressed-punctate ............ M. ventuarensis Leaves petiolate, gradually narrowed to base ........................................ 13 Leaves subsessile, broadly rounded to subcordate at base .................... 14 Flowers 1 or 3 per inflorescence .................................................... M. uniflora Flowers 30–50 per inflorescence ................................................ M. ligustrina

42

M YRTACEAE

14(12). Flower buds not wrinkled in drying; calyx usually not calyptrate, but splitting irregularly longitudinally, the 3 or 4 lobe-like segments reflexed and persistent ............................................................... M. montana 14. Flower buds fleshy, coarsely wrinkled in drying; calyx calyptrate, the calyptra deciduous, the hypanthium margin becoming irregularly 5– 8-lobulate ........................................................................ M. suborbicularis 15(1). Midvein of leaf blades convex, biconvex, or plane on upper surface ...... 16 15. Midvein of leaf blades sulcate, often deeply so, or at least V-shaped in cross section ......................................................................................... 22 16(15). Indument of coppery, ferruginous, brownish, or reddish trichomes ..... 17 16. Indument of whitish, grayish, yellowish, or straw-colored trichomes ...... 20 17(16). Petioles 4 mm wide (including trichomes); trichomes of stems and petioles spreading ............................................................................ M. insignis 17. Petioles 1–2 mm wide; trichomes of stems and petioles appressed ....... 18 18(17). Leaf blades 4–7 cm long, apex long-caudate................................. M. caudata 18. Leaf blades (10)15–30 cm long, apex acuminate .................................... 19 19(18). Leaf blades elliptic to obovate, the apex reflexed, conduplicate; petioles 1–2 cm long ............................................................................ M. caesariata 19. Leaf blades oblong to elliptic, the apex not reflexed (plane) nor conduplicate; petioles < 1 cm long ................................................... M. convexivenia 20(16). Leaf blades pendent, lower surface lustrous, densely gray-sericeous, 1– 1.5 cm wide .................................................................................. M. pudica 20. Leaf blades not pendent, the lower surface glabrous or thinly sericeous, 4–14 cm wide ....................................................................................... 21 21(20). Leaf blades with revolute margins, upper surface lustrous; fruits not glaucous ........................................................................................... M. cana 21. Leaf blades with planar margins, upper surface dull; fruits glaucous ......................................................................................................... M. sp. A 22(15). Indument of whitish, grayish, yellowish, or straw-colored trichomes; calyx closed in bud .................................................................................. 23 22. Indument of coppery, ferruginous, brownish, or reddish trichomes; calyx open in bud, with 4 or 5 small lobes ................................................... 26 23(22). Leaf blades subcordate to cordate at base .............................................. 24 23. Leaf blades rounded to cuneate at base, clearly petiolate, the secondary veins plane or slightly elevated on upper surface; fruit rounded, muricate ............................................................................................... 25 24(23). Leaf blades broadly obtuse to rounded at apex, 7–10 cm long, the secondary veins of leaf blades slightly impressed on upper surface and conspicuously elevated on lower surface, lower surface glabrous to sparsely pilosulose; flower buds densely sericeous on hypanthium, calyx glabrous ....................................................................... M. subcordata 24. Leaf blades acute to acuminate at apex, > 15 cm long, the secondary veins of leaf blades plane on upper surface, inconspicous on lower surface, lower surface densely yellowish-sericous; flower buds sericeous on hypanthium and calyx ................................................................. M. subulata 25(23). Fruits 0.7–1 cm wide; leaf blades rounded to obtuse or acute, 5.4–9 cm long ......................................................................................... M. mcvaughii 25. Fruits 1.7–1.9 cm wide; leaf blades acuminate, 9–18 cm long .... M. spruceana 26(22). Leaves mostly glabrous at maturity ....................................................... 27 26. Leaves persistently and densely pubescent on lower surface, at least

Marlierea 43

along the veins ..................................................................................... 28 Petioles wrinkled, corky ................................................................... M. cuprea Petioles smooth ............................................................................... M. summa Leaf blades obtuse to rounded at apex .................................................... 29 Leaf blades acuminate to caudate ........................................................... 30 Leaf blades short-petiolate, acute to obtuse at base; inflorescences > 5flowered .................................................................................... M. buxifolia 29. Leaf blades sessile, subcordate at base; inflorescences 3-flowered .................................................................................................. M. foveolata 30(28). Leaf blades rugose-bullate, trichomes on lower surface predominating on the veins; petioles 1.2–2 cm long ............................................. M. rugosior 30. Leaf blades smooth; trichomes on lower surface evenly distributed on veins and lamina; petioles < 1 cm long ............................................... 31 31(30). Leaf blades not impressed-punctate on upper surface; trichomes pale brown ...................................................................................... M. ferruginea 31. Leaf blades impressed-punctate on upper surface; trichomes dark reddish brown .................................................................................... M summa 27(26). 27. 28(26). 28. 29(28).

Marlierea bipennis (O. Berg) McVaugh, Mem. New York Bot. Gard. 10(1): 79. 1958. —Myrciaria bipennis O. Berg, Linnaea 31: 259. 1862. —Myrcia bipennis (O. Berg) McVaugh, Fieldiana, Bot. 29: 189. 1956. Shrub or small tree 2.5–4 m tall; leaves ovate to elliptic, petioles 2–4 mm long; inflorescence few-flowered, mostly glabrous. Moist evergreen forests along rivers, ca. 100 m; Amazonas (Caño Yagua, Río Casiquiare, Río Negro). Brazil (Amazonas). ◆Fig. 52. Marlierea buxifolia Amshoff, Bull. Torrey Bot. Club 75: 529. 1948. Shrub or small tree 1–2(–6) m tall; leaf blades coriaceous, revolute, the lower surface dark brown or ferruginous-pubescent; inflorescence densely villous, 3–5-flowered; fruits purple. Montane, moist dwarf forests, shrub islands, 1200–1400 m; Bolívar (Cerro Venamo, Sierra de Lema). Guyana. ◆Fig. 45. Marlierea caesariata McVaugh, Mem. New York Bot. Gard. 10(1): 81. 1958. Tree 12 m tall; inflorescence and young growth ferruginous-velutinous; leaf blades abruptly acuminate, midvein flat or convex on upper surface; inflorescences many flowered; flowers nearly closed in bud, with 4 or 5 lobes. Lower montane Clusia forests, 600– 700 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 42. Marlierea cana McVaugh, Mem. New York Bot. Gard. 10(1): 82. 1958.

Spindly or scandent shrub 1–3 m tall; leaf blades mostly elliptic, margins revolute, apex obtuse to rounded; flowers and inflorescence branches appressed white- or yellowish-pubescent on hypanthium; fruit purple. Moist scrub forests, rocky slopes and ridges, 1100– 2000 m; Amazonas (Cerro Duida, Cerro Parú, Cerro Yutajé). Endemic. ◆Fig. 53. Marlierea caudata McVaugh, Fieldiana, Bot. 29: 176. 1956. Tree 5–6 m tall; pubescence coppery; leaves abruptly long-caudate, the midvein convex and higher order venation inconspicuous; inflorescence few-flowered. Nonflooded evergreen lowland forests, ca. 100 m; Amazonas (Caño Caname, Río Guainía). Amazonian Colombia, Peru, and Brazil. Marlierea convexivenia B. Holst, Selbyana 23: 144. 2002. Tree to 5 m tall; young vegetative parts and inflorescence densely coppery-sericeous; leaf blades mostly oblong to elliptic, sharply acuminate, midvein convex on upper surface, secondary venation inconspicuous. Lowland evergreen forests on white sand, periodically flooded riparian forests, shrub savannas, 100–200 m; Amazonas (Caño Duapo in Río Siapa basin, Caño San Miguel, Río Atabapo, Río Pasimoni). Endemic. Marlierea cuprea Amshoff, Bull. Torrey Bot. Club 75: 530. 1948. Shrub or tree to 5 m tall; leaf blades coriaceous, secondary venation inconspicuous,

44

M YRTACEAE

midvein sulcate, petioles wrinkled and corky; inflorescence coppery-pubescent. Along black-water rivers or upland evergreen forests, 100–500 m; Bolívar (La Escalera), Amazonas (Caño San Miguel, Río Emoni). Guyana. The Venezuelan Amazonian collections of Marlierea cuprea (Aymard 9050, SEL; Stergios et al. 8269, MO, SEL) are very much out of altitudinal and geographical range from the type locality populations, but the material seems indistinguishable. Marlierea ensiformis McVaugh, Mem. New York Bot. Gard. 18(2): 64. 1969. Tree 10 m tall, glabrous; leaf blades elliptic, impressed-punctate on upper surface, midvein sulcate; inflorescences of paired panicles. Lower montane forests, 700–800 m; presumably Amazonas (Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). Marlierea ferruginea (Poir.) McVaugh, Mem. New York Bot. Gard. 10(1): 83. 1958. —Eugenia ferruginea Poir. in Lam., Encycl. suppl. 3: 124. 1813. —Myrcia ferruginea (Poir.) DC., Prodr. 3: 245. 1828. —Krugia ferruginea (Poir.) Urb. in Krug & Urb., Bot. Jahrb. Syst. 19: 604. 1895. —Pawaruyek (Arekuna). Marlierea acuminata O. Berg, Linnaea 27: 13. 1855. Shrub 2–4 m tall, or usually a tree 4–12 (–20) m; lower surface of leaf blades densely pale brown-tomentulose; fruit purple-black. Lower montane to montane forests, gallery forests, 500–1200 m; Bolívar (Gran Sabana), Amazonas (Cerro Aratitiyope, Cerro Sipapo, Los Pijiguaos, Piedra Arauicaua). Trinidad, Lesser Antilles, Guyana, Suriname, French Guiana, Brazil (Amazonas). The inflorescence bracts on the collections of Marlierea ferruginea from Venezuela are caducous, whereas they are persistent on specimens from other areas. Marlierea foveolata B. Holst, Selbyana 23: 145. 2002. Shrub 1–2 m tall; leaves stiff, lower surface of young blades densely ferruginous-tomentose. Open rocky areas, 2000–2200 m; Bolívar-Amazonas border (Sierra de Maigualida). Endemic. Marlierea guildingiana (Griseb.) Krug & Urb., Bot. Jahrb. Syst. 19: 591. 1895.

—Psidium guildingianum Griseb., Fl. Brit. W. I. 242. 1860. Tree 4–12(–24) m tall; branchlets narrowly winged; fruits reddish or orangish turning black. Evergreen lowland forests, 100–300 m; Delta Amacuro (Serranía de Imataca), Bolívar (Altiplanicie de Nuria, Río Nichare, Serranía de Imataca). St. Vincent, Trinidad, Guyana. ◆Fig. 40. Marlierea guildingiana is closely related to, and scarcely distinct from, M. bipennis. Marlierea insignis McVaugh, Fieldiana, Bot. 29: 176. 1956. Tree to 7 m tall with dense crown; bark rough, fissured; young vegetative growth and inflorescence densly ferruginous-velutinous. Seasonally flooded river banks, ca. 100 m; Amazonas (Río Guainía). Amazonian Colombia and Peru. The single collection of Marlierea insignis known from Venezuela (Williams 14302, F, US) has the leaves relatively narrower than those of the type collection from Colombia. A collection from Bolívar state, lower Río Caura (Stergios & Elcoro 12182, PORT, SEL), may represent an additional population of Marlierea insignis for Venezuela, or perhaps a distinct, related taxon, but flowers are needed for final placement. Marlierea karuaiensis (Steyerm.) McVaugh, Mem. New York Bot. Gard. 10(1): 85. 1958. —Aulomyrcia karuaiensis Steyerm., Fieldiana, Bot. 28: 1005. 1957. Shrub or small tree to 8 m tall, mostly glabrous; leaves variable in width, secondary veins inconspicuous; inflorescences few-flowered; fruits purple-black at maturity. Lowland to upland scrub forests, rock crevices, along rocky streams, gallery forests, 300– 1700 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Gran Sabana, Macizo del Chimantá [Apacará-tepui], Ptari-tepui), Amazonas (Cerro Avispa, Cerro Vinilla, Sierra de la Neblina). Guyana. Marlierea ligustrina McVaugh, Mem. New York Bot. Gard. 10(1): 86. 1958. Small tree to 10 m tall; leaves coriaceous, short-petiolate, midvein convex, secondary venation inconspicuous, apex obtuse; inflorescences many-flowered; fruit purple-black. Upland scrub forests, 1000–1200 m; Amazonas (Cerro Huachamacari, Cerro Yapa-

Marlierea 45

cana). Endemic. ◆Fig. 48. Marlierea ligustrina is closely related to, and perhaps not distinct from, M. montana Aubl. Marlierea lituatinervia (O. Berg) McVaugh, Mem. New York Bot. Gard. 18(2): 65. 1969. —Myrciaria lituatinervia O. Berg, Linnaea 27: 322. 1856. Habit unknown; branchlets markedly keeled; leaf blades to 7 cm long, subcordate at base, rounded at apex. Among rocks, near brooks, ca. 400 m; Bolívar (near Roraimatepui), or possibly Guyana. The only known collection of this species (Schomburgk 558/877b, K) was apparently made along the Venezuela/Guyana border. Marlierea maguirei McVaugh, Mem. New York Bot. Gard. 10(1): 87. 1958. Shrub or small tree 0.5–3 m tall; branchlets narrowly winged; fruit changing from green to orange to dark purple, slightly spicy. Along rocky streams, low forested slopes, 200–1500 m; Amazonas (Cerro Coro Coro, Cerro Yutajé). Endemic. ◆Fig. 55. The leaf width and shape in Marlierea maguirei is extremely variable; collections from along rivers have nearly linear leaves while those from drier, forested areas are elliptic or oblong. Marlierea mcvaughii B. Holst, Selbyana 23: 147. 2002. Shrub or tree 3–12 m tall; indument of lower leaf surface, inflorescence, and flowers whitish or dull yellowish; secondary leaf venation inconspicuous; flowers densely sericeous, fruits muricate. Periodically flooded white-sand shrubby savannas, ca. 100 m; Amazonas (Río Negro basin). Río Negro basin of Colombia (Meta) and Brazil (Amazonas). Marlierea montana (Aubl.) Amshoff, Recueil Trav. Bot. Néerl. 39: 147. 1942. —Eugenia montana Aubl., Hist. Pl. Guiane 495, t. 195. 1775. Calyptranthes obtusa Benth., J. Bot. (Hooker) 2: 319. 1840. —Marlierea obtusa (Benth.) O. Berg, Linnaea 27: 15. 1855. Few-stemmed, mostly glabrous shrub or spindly tree to 5(–8) m tall; leaf blades stiffcoriaceous, ovate, rounded to cordate at base, nearly sessile, obtuse to rounded at apex; inflorescences subterminal, many-flowered,

glabrous. Upland scrub forests, gallery forests, mixed lowland evergreen forests, (50–) 500–100 m; Delta Amacuro (Caño Simoina), Bolívar (Cerro Guaiquinima, Cerro Guanacoco, Gran Sabana, middle Río Caroní, Serranía Senkopirén). Guyana, Suriname, French Guiana, Brazil (Amazonas: Serra Araca). ◆Fig. 46. The single collection of Marlierea montana from Delta Amacuro at 50 m elevation (Steyermark et al. 114398, VEN) has leaves thinner than those from other areas. Other populations in Venezuela are somewhat variable as to the size of the leaves. An additional collection from 1100 m on Cerro Parú, Amazonas state (Huber 4280, MICH, NY, VEN), may represent a variety of M. montana, or perhaps a distinct species. It has generally smaller leaves that are imbricate along the stem, giving it a distinct aspect. Marlierea pudica McVaugh, Brittonia 33: 35. 1981. Shrub 1–3 m tall; branches elongated, arching or pendulous; leaf blades drooping, margins revolute, the lower surface densely gray-sericeous, lustrous; inflorescence fewflowered. Sandy, rocky, open areas on tepui summit, 900–1000 m; Bolívar (Cerro Guaiquinima). Endemic. ◆Fig. 41. Marlierea rugosior McVaugh, Mem. New York Bot. Gard. 18(2): 67. 1969. Slender tree 2–10 m tall; leaves large, rugose-bullate, indument of lower surface and young stems dark brown; flower buds 4–5 mm long, calyx open in bud, 5-lobed. Evergreen forests on tepui summit, 1700–1800 m; Bolívar (Auyán-tepui). Endemic. ◆Fig. 47. This appears to be a species of Myrcia (Gomidesia), but is known only from the type, which is past flower, and one additional sterile collection. Marlierea schomburgkiana O. Berg, Linnaea 29: 209. 1858. —Dau dau. Tree 4–15(–25) m tall, rarely a shrub 1–2 m; leaf blades thin, abruptly caudate-acuminate, midvein convex on upper surface, foliar glands relatively few and sparse; inflorescence glabrous; fruit dark blue at maturity, the walls thick. Understory of premontane to montane, deciduous to evergreen forests, riverbanks, 200–1000 m; Delta Amacuro (Serranía de Imataca), Bolívar (Gran Sabana, Río Cuyuní basin, Serranía de

46

M YRTACEAE

Imataca), Amazonas (base of Sierra de la Neblina). Guyana, Suriname, Ecuador, Peru. ◆Fig. 44. The wood of Marlierea schomburgkiana is sought for its hardness, and the fruit has been described as sweet. Marlierea scytophylla Diels, Verh. Bot. Vereins Prov. Brandenburg 48: 187. 1907. Small tree; leaf blades coriaceous, abruptly acuminate, midvein convex on upper surface, petioles wrinkled; fruits glaucous. Flooded riparian forests, ca. 100 m; Amazonas (Caño Cuweje south of San Carlos de Río Negro). Brazil (Amazonas). Marlierea spruceana O. Berg in Mart., Fl. Bras. 14(1): 34. 1857. Marlierea uaupensis O. Berg in Mart., Fl. Bras. 14(1): 516. 1857. Shrub to 3 m or tree 3–15 m tall; inflorescence densely appressed-pubescent with yellowish trichomes; calyx closed in bud, not apiculate; fruits muricate, yellow when mature. Seasonally flooded riparian forests, forested river banks, scrub forests, 100–200 m; Amazonas (near Cerro Yapacana, basins of Río Casiquiare and Río Negro, Río Guayapo, Río Sipapo, Santa Barbara del Orinoco). Amazonian Colombia, Peru, and Brazil. ◆Fig. 43. Marlierea subcordata B. Holst, Selbyana 23: 150. 2002. Tree 8–10 m tall; leaves lustrous, midvein on upper surface slightly concave; inflorescence, flower buds, and lower surface of leaf blades sericeous. Lowland evergreen forests along peridiodically flooded river banks, ca. 100 m; Amazonas (Río Baría, Río Pasimoni). Endemic. Marlierea suborbicularis McVaugh, Mem. New York Bot. Gard. 10(1): 88. 1958. Shrub 1–2.5 m tall; leaves coriacous, rounded at apex, nearly sessile; flowers white or cream-colored, the buds wrinkled; fruit purple-black at maturity. Lowland moist savannas, white-sand savannas, forest understory, 100–200 m; Amazonas (Caño Cotua, Cerro Yapacana base, Río Autana basin, Río Guainía, junction of Río Orinoco and Río Ventuari). Endemic. ◆Fig. 50.

Apparently variable in leaf shape, Marlierea suborbicularis is closely related to Marlierea montana and M. ligustrina. Marlierea subulata McVaugh, Fieldiana, Bot. 29: 177. 1956. Shrub or small tree 2–3 m tall; lateral and marginal veins impressed on upper leaf surface; calyx closed in bud, apiculate; fruit oblate. Along streams, nonflooded forests, ca. 100(–700) m; Amazonas (near San Carlos de Río Negro, Sierra de la Neblina). Amazonian Peru, Brazil (Amazonas, Rondônia), and Bolivia. Marlierea summa McVaugh, Mem. New York Bot. Gard. 10(1): 89. 1958. Shrub or tree to 12 m tall; most vegetative and reproductive parts of plant abundantly ferruginous-pubescent; leaf blades impressed-punctate on upper surface, midvein sulcate, apex acuminate to caudate. Usually in upland or highland tepui forests, 1400– 2300 m; Bolívar (Auyán-tepui, Cerro Jaua, Ilú-tepui, Macizo del Chimantá), Amazonas (Cerro Aracamuni, Cerro Marahuaka, Cerro Sipapo, Sierra de la Neblina). Guyana, Brazil (Amazonas: Serra Aracá). ◆Fig. 49. Marlierea summa is a widespread and variable species of higher elevations in the Guayana Shield. Three varieties have been recognized (var. calva McVaugh, var. superior McVaugh, and var. summa). However, there seem to be several additional fairly distinct populations that cannot be accomodated with the current infraspecific classification scheme. Some of these may merit recognition at the species level, e.g., a collection from the Macizo del Chimantá (Pipoly et al. 7211, NY, SEL) has the flowers twice the size of typical M. summa, and another, known from two localities from low elevation areas around the Río Atacavi (Piñate & Mondolfi 1007, MICH) and Río Negro on the Colombian side (Schultes & López 9399, US), differ by having exceptionally long petioles, lustrous leaf blades, and in lacking the impressed punctations of the typical higher elevation collections. Another closely related species, M. vicina McVaugh, occurs in Guyana. Marlierea umbraticola (H.B.K.) O. Berg, Linnaea 27: 17. 1855. —Myrtus umbraticola H.B.K., Nov. Gen. Sp. (folio ed.) 7: 258. 1825.

Marlierea 47

Shrub or tree to 8 m tall; leaves coriacous, the petiole conspicuously flaky-corky; inflorescence gray-puberulent, bracts persistent, flowers opening sequentially. Seasonally flooded banks of largely black-water rivers, lowland evergreen forests, 100–200 m; western Bolívar, Amazonas (widespread). Amazonian Colombia, Peru, Brazil, and Bolivia. ◆Fig. 51. Marlierea umbraticola has affinities to several species of Myrcia (e.g., M. decorticans DC., M. nigrescens DC., M. rupta Kawasaki & B. Holst), and may eventually be transferred there. Marlierea uniflora McVaugh, Mem. New York Bot. Gard. 18(2): 69. 1969. Shrub or small tree 1–3 m tall; leaf blades coriaceous, mostly obovate, rounded at apex, venation inconspicuous, impressed-punctate on upper surface; inflorescences 1- or 3-flowered, glabrous. Flooded forests along blackwater rivers, savannas, 100–200 m; southwest Amazonas (Río Atabapo, Río Atacavi, Río Casiquiare, Río Guainía, Río Pasimoni, Río Temi). Brazil (Amazonas: upper Rio Negro). ◆Fig. 54.

Shrub or treelet to 10 m tall; leaf blades coriaceous, acuminate, impressed-punctate and with shallowly sulcate midvein on upper surface, venation inconspicuous; inflorescence many-flowered, the peduncles and pedicels branching at 90° angles. Shrubby savannas or scrub, 200–400 m; Amazonas (Cerro Moriche, Río Manapiare basin). Endemic. Marlierea sp. A Tree 4–12 m tall; petioles rimose or not; fruit glaucous. Seasonally flooded riparian forests, ca. 100 m; Amazonas (Río Casiquiare, Río Guainía, Río Negro). Endemic. While this is likely an undescribed species, all eight known collections of it are galled, making it difficult to place. It is possibly a species of Myrcia.

Marlierea ventuarensis B. Holst, Selbyana 23: 152. 2002.

Fig. 40. Marlierea guildingiana

48

M YRTACEAE

Fig. 41. Marlierea pudica

Fig. 42. Marlierea caesariata

Marlierea 49

Fig. 43. Marlierea spruceana

Fig. 44. Marlierea schomburgkiana

Fig. 45. Marlierea buxifolia

50

M YRTACEAE

Fig. 46. Marlierea montana

Fig. 47. Marlierea rugosior

Marlierea 51

Fig. 48. Marlierea ligustrina

Fig. 49. Marlierea summa

Fig. 50. Marlierea suborbicularis

52

M YRTACEAE

Fig. 51. Marlierea umbraticola

Fig. 52. Marlierea bipennis

Marlierea 53

Fig. 53. Marlierea cana

Fig. 54. Marlierea uniflora

Fig. 55. Marlierea maguirei

54

M YRTACEAE

8. MYRCIA DC. ex Guill. in Bory, Dict. Class. Hist. Nat. 11: 401. 1827. [Subtribe Myrciinae]. Aulomyrcia O. Berg, Linnaea 27: 35. 1855. Gomidesia O. Berg, Linnaea 27: 6. 1855. by Bruce K. Holst Shrubs or trees; stems usually terete or compressed. Leaves variable in size, shape, and indument. Inflorescence an axillary or subterminal panicle, rarely reduced to one flower, bracts present, small, persistent or deciduous. Flowers 4- or 5merous. Calyx lobes imbricate, equal to markedly unequal, rarely subfoliaceous, hypanthium equal to the summit of the ovary (section Myrcia and what is sometimes recognized as the distinct genus Gomidesia) to prolonged above it (sections Armeriela, Aulomyrcia). Stamens numerous; filaments incurved in bud; anthers ovate, dorsifixed, opening by longitudinal slits. Ovary 2- or 3(4)-locular; ovules 2 per locule. Fruit round or ellipsoid (section Myrcia). Seeds 1 or 2(3); seed coat cartilaginous, shiny; embryo myrcioid. New World tropics; ca. 300 species, ca. 60 in Venezuela, 54 of these in the flora area. Myrcia is considered in the broad sense here, though it contains such diverse elements that it may need realignment in the future. Myrcia is one of the most difficult genera in regard to species delimitation, particularly in section Myrcia. The section itself is well distinguished by the lack of hypanthial prolongation and by the spear shape of the vegetative buds (the spear shape is the result of adherent developing leaves, with the petioles resembling the handle of the spear, and the small blades the tip), but it is difficult to draw the line between many described names. That difficulty is reflected in the latter section of the key presented below and users will undoubtedly encounter difficulty in placing a specimen with any certainty. A more extensive key to these and other northern South American species can be found in R. McVaugh, Mem. New York Bot. Gard. 18(2): 75–149. 1969. Other groups of Myrcia (excluding gomidesioid plants—see following), have cone-shaped vegetative buds. Gomidesia may deserve generic recognition, but it so closely resembles Myrcia section Myrcia with the characteristic spear-shaped buds, reticulate leaf venation, and lack of a prolonged hypanthium, that it is included here. If Gomidesia were to be recognized, species from the flora area possibly included there are M. bonnetiasylvestris, M. clausa, M. fenzliana, and M. sipapensis. Key to the Species of Myrcia 1. 1. 2(1). 2. 3(2). 3. 4(3).

Midvein on upper surface of leaves plane or elevated in some fashion (convex, biconvex, or in a narrow elevated line) .................................. 2 Midvein on upper surface of leaves impressed, usually V- or U-shaped in cross section ......................................................................................... 16 Flowers copiously pubescent (minutely so in M. pistrinalis), the indument often persistent into fruit ..................................................... 3 Flowers glabrous, or at most with a few trichomes basally ..................... 9 Leaf blades acuminate to caudate at apex................................................ 4 Leaf blades rounded to obtuse at apex...................................................... 6 Lower surface of leaf blades yellow-tan appressed-pubescent; leaf blades

Myrcia 55

4.

5(4).

5. 6(3). 6. 7(6). 7.

8(6).

8.

9(2). 9. 10(9). 10. 11(10).

11.

12(10). 12. 13(9). 13. 14(13). 14. 15(13). 15. 16(1). 16.

obovate to broadly elliptic, caudate-acuminate at apex; flowers 4-merous, hypanthium yellow-tan pubescent ........................... M. compta Leaves mostly glabrous on both surfaces; leaf blades elliptic to oblong, the apex acuminate; flowers 5-merous; hypanthium white-sericeous or puberulent .............................................................................................. 5 Hypanthium puberulent; inflorescence bracts deciduous at or before anthesis; flowers stout-pedicellate, pedicels 5–10 mm long; ovary 3-locular ................................................................................................ M. pistrinalis Hypanthium white-sericeous; inflorescence bracts mostly persistent into fruit; flowers sessile; ovary 2-locular ....................................... M. pyrifolia Flowers 4-merous ....................................................................................... 7 Flowers 5-merous ....................................................................................... 8 Inflorescences and young stems coppery-sericeous; inflorescences broadly paniculate .................................................................................. M. nubicola Inflorescences and young stems abundantly and softly tomentose with mostly whitish or brownish yellow trichomes; inflorescences racemose or with occasional branches basally ...................................... M. ptariensis Secondary and tertiary venation obscure on upper surface of leaves; leaves and young branches pale-strigose with appressed trichomes ................................................................................................... M. salticola Secondary and tertiary venation impressed on upper surface of leaves, evident; leaves and young branches soft-villous with yellowish trichomes .................................................................................... M. tomentosa Flowers 4-merous, calyx lobes equal or equal in 2 pairs ........................ 10 Flowers 5-merous, calyx lobes usually unequal ..................................... 13 Leaf blades obtuse to rounded or emarginate at apex ........................... 11 Leaf blades acute to acuminate at apex .................................................. 12 Leaves short-petiolate to nearly sessile, oblong, cordate at base, 6–12 cm long; secondary veins evident, widely separated with the tertiary veins forming a conspicuous reticulation; calyx lobes densely pubescent on inner surfaces ................................................................. M. ehrenbergiana Leaves petiolate, mostly obovate, cuneate at base, 2–4 cm long, secondary veins fine, scarcely evident, numerous, parallel, inconspicous; calyx lobes glabrous on inner surfaces ........................................... M. rotundata Inflorescence glabrous; calyx lobes of equal pairs, glabrous within .................................................................................................... M. grandis Inflorescence reddish pubescent; calyx lobes of unequal pairs; sparsely appressed-pubescent within ................................................. M. quitarensis Calyx lobes glabrous within; ovary 2-locular ......................................... 14 Calyx lobes appressed-pubescent within; ovary 3-locular ..................... 15 Leaves acuminate at apex, 2.5–3.5 times longer than wide ......... M. liesneri Leaves rounded to emarginate at apex, < 1.5 times longer than wide ............................................................................................. M. revolutifolia Flower buds 3–3.5 cm long ........................................................... M. citrifolia Flower buds 1–2 mm long ......................................................... M. guianensis Hypanthium prolonged beyond the summit of the ovary, present as short cylindrical crown in fruit; terminal vegetative bud conical ............... 17 Hypanthium not prolonged beyond the summit of the ovary; summit of

56

M YRTACEAE

17(16). 17. 18(17). 18. 19(18). 19. 20(18). 20. 21(20).

21.

22(20).

22.

23(22).

23. 24(23). 24. 25(17). 25 26(25). 26. 27(26). 27. 28(27). 28.

ovary persistently pubescent to hirsute; terminal vegetative bud shaped like a short-handled spear (the shape the result of adherent developing leaves, with the petioles resembling the handle of the spear, and the small blades the tip) .................................................... 35 Hypanthium glabrous without, or at most with a few sparse trichomes .............................................................................................................. 18 Hypanthium pubescent without, usually densely so ............................. 25 Petioles corky-rimose; secondary and tertiary venation inconspicuous, especially on upper surface ..................................................................... 19 Petioles smooth; secondary and tertiary venation evident on both surfaces ...................................................................................................... 20 Fruits thick-walled, 10–20(–30) mm diameter; leaves usually abruptly acuminate at apex ............................................................... M. decorticans Fruits thin-walled, 4–7 mm diameter; leaves usually obtuse to rounded at apex ............................................................................................. M. sp. B Inflorescence bracts persistent past anthesis and usually into fruit .... 21 Inflorescence bracts deciduous at or before anthesis ............................. 22 Leaf blades rounded to scarcely cordate at base, often appearing polished and lustrous with the tertiary venation inconspicuous on upper surface, gradually long-acuminate at apex ....................................... M. lucida Leaf blades obtuse to cuneate at base, not appearing polished, the tertiary venation forming an evident reticulum, abruptly short-acuminate at apex ........................................................................ M. inaequiloba Flower buds 1–1.5 mm long; bark on year-old twigs reddish, flaky, peeling in small patches; inflorescence branches usually thinly puberulent ............................................................................................... M. amazonica Flower buds 2–4 mm long; bark on year-old twigs peeling in thin strips or not at all; inflorescence branches glabrous or with sparse, thick, dibrachiate trichomes .......................................................................... 23 Leaf blades round to broadly elliptic to broadly ovate, rounded at base and rounded to shallowly emarginate at apex; inflorescences sparsely branched, appearing racemose ..................................................... M. sp. D Leaf blades ovate, elliptic, or obovate, narrowed at either base or apex; inflorescences distinctly branched ...................................................... 24 Petals in bud forming a conspicuous globe above the calyx; leaf blades membranaceous, the apex slightly conduplicate and recurved ...... M. multiflora Petals not forming a conspicuous globe above the calyx; leaf blades coriaceous, the apex plane ............................................................ M. platyclada Leaves rounded, obtuse, acute, or rarely shortly and bluntly acuminate at apex ...................................................................................................... 26 Leaves prominently acuminate to caudate ............................................. 30 Leaf blades 1.4–1.8 cm long ............................................................ M. gentryi Leaf blades > 3 cm long ........................................................................... 27 Leaf blades > 12 cm long; petioles ca. 1 cm long ............................... M. sp. A Leaf blades 3.5–8 cm long ........................................................................ 28 Leaf blades strongly revolute, stems and inflorescence branches tomentose ........................................................................... M. kylistophylla Leaf blades plane or the margins scarcely revolute; stems and inflores-

Myrcia 57

cence branches sericeous or appressed-pubescent ............................. 29 29(28). Flowers 4-merous; leaf blades broadly obtuse to rounded, the upper surface somewhat pebbly and impressed-punctate, the venation inconspicuous; petioles corky-rimose .................................. M. albido-tomentosa 29. Flowers 5-merous; leaf blades shortly and bluntly acuminate at apex, upper surface with evident reticulate venation, not impressed-punctate; petioles smooth ................................................................... M. exploratoris 30(25). Lower surface of young leaf blades densely sericeous, usually permanently so ............................................................................................... 31 30. Lower surface of all leaf blades glabrous or sparsely pubescent ........... 33 31(30). Calyx lobes 3–5 mm long, densely appressed-pubescent within; inflorescences > 3 cm long ................................................................ M. calycampa 31. Calyx lobes ca. < 1.5 mm long, glabrous within; inflorescences 0.5–2 cm long ....................................................................................................... 32 32(31). Inflorescence axis 5–12 mm long; pubescence of fruit somewhat loose, not closely appressed ......................................................................... M. induta 32. Inflorescence axis to 3 mm long; pubescence of fruit fine, appressed ............................................................................................... M. sessiliflora 33(30). Inflorescence persistently bracteate, the bracts foliaceous; young stems and inflorescence branches hirsute with long erect yellow trichomes; leaf blades gradually acuminate at apex ............................... M. bracteata 33. Inflorescence bracts deciduous at or before anthesis, the bracts not foliaceous; young stems and inflorescence branches with appressed trichomes; leaf blades caudate to caudate-acuminate at apex .............. 34 34(33). Leaf blades with numerous parallel fine secondary veins and a marginal vein mostly parallel with margin, these plane and scarcely evident on upper surface; calyx lobes glabrous within ....................... M. bolivarensis 34. Leaf blades with 6–8 evident, secondary veins and a looping marginal vein, these usually slightly impressed on upper surface and evident on lower surface; calyx lobes pubescent within ....................... M. minutiflora 35(16). Indument dark reddish brown to chestnut brown ................................. 36 35. Indument pale yellow, pale yellow-brown, coppery, tawny, or silvery .... 39 36(35). Leaf blades nearly rounded to broadly ovate, the apex abruptly and shortly acuminate, the acumen to 5 mm long, secondary leaf venation impressed on upper surface ........................................................... M. sp. C 36. Leaf blades obovate, elliptic, or ovate, the apex long-caudate-acuminate (acumen > 10 mm long) or rounded to acute, secondary leaf venation plane to slightly elevated on upper surface ........................................ 37 37(36). Leaf blades long-caudate-acuminate, 10–17 cm long ................ M sipapensis 37. Leaf blades rounded to obtuse or rarely acute at apex, 3–9 cm long .... 38 38(37). Leaves glabrous to glabrate or only sparsely pubescent ............................ ................................................................................... M. bonnetiasylvestris 38. Leaves persistently and densely tomentose on lower surface ........ M. crispa 39(35). Calyx lobes imbricate and erect or incurved in fruit, the calyx forming a crown .................................................................................................... 40 39. Calyx lobes usually spreading in fruit, not imbricate ............................ 42 40(39). Leaf blades sparsely pubescent to glabrous on lower surface, the margins plane; secondary veins slightly elevated on upper surface ......... M. aliena

58

M YRTACEAE

40.

41(40). 41. 42(39). 42. 43(42). 43. 44(42). 44. 45(44).

45. 46(45). 46. 47(45). 47. 48(47). 48. 49(44).

49. 50(49).

50.

51(49). 51. 52(51).

52.

Leaf blades abundantly pubescent on lower surface, especially so along the veins, the margins revolute; secondary veins slightly impressed on upper surface ....................................................................................... 41 Leaf blades caudate-acuminate, not splitting longitudinally at the apex when pressed flat ......................................................................... M. clausa Leaf blades obtuse to acuminate, often splitting longitudinally at the apex when pressed flat ............................................................ M. fenzliana Upper surface of leaves impressed-punctate (use hand lens) ................ 43 Upper surface of leaves not impressed-punctate .................................... 44 Leaf blades mostly > 5 cm long, obtuse to rounded at base; petioles 4–7 mm long ................................................................................. M. paivae Leaf blades mostly < 5 cm long, cuneate at base; petioles to 3 mm long .................................................................................................. M. sylvatica Inflorescence, young stems, and leaves densely pubescent ................... 45 Inflorescence, young stems, and leaves sparsely pubescent .................. 49 Lower surface of leaf blades finely and minutely pebbled (use hand lens) or densely golden sericeous and the surface not evident; sinus between calyx lobes rounded ............................................................................. 46 Plants not as above .................................................................................. 47 Fruits 6–9 mm diameter; petioles < 1 cm long; lower surface of leaf blades dull, minutely pebbled ................................................................ M. deflexa Fruits 15–25 mm diameter; petioles 2–2.5 cm long; lower surface of leaf blades densely golden sericeous, the pebbling inconspicuous ..... M. intonsa Calyx lobes glabrous on inner surface; inflorescence hirsutulous ........................................................................................ M. sororopanensis Calyx lobes densely pubescent on inner surface; inflorescence appressedpubescent ............................................................................................. 48 Leaf blades 2–3 cm wide ................................................................. M. servata Leaf blades 4–8 cm wide ....................................................... M. magnoliifolia Leaf blades broadly ovate to obovate, rounded to obtuse at apex, mostly < 7 cm long (see also the key to the varieties of M. subsessilis, a variable species in regard to petiole length and leaf size) ....................... 50 Leaf blades ovate-lanceolate to elliptic, acuminate at apex, mostly > 7 cm long ....................................................................................................... 51 Leaf blades rounded to emarginate at apex, somewhat conduplicate and often splitting lengthwise when pressed flat; calyx lobes pubescent within ...................................................................................... M. clusiifolia Leaf blades broadly obtuse to subacuminate at apex, not conduplicate and splitting lengthwise when pressed flat; calxy lobes glabrous within ................................................................................................. M. tepuiensis Leaves subsessile, the blades subcordate or broadly rounded at base, petioles 1–3 mm long ........................................................................... 52 Leaves evidently petiolate, the blades usually acute to cuneate at base, petioles > 3 mm long ............................................................................ 53 Inflorescence axes narrow and terete, the peduncle 1–1.5 mm wide below the lowest inflorescence branches; leaf blades broadly rounded to scarcely subcordate at base .................................................... M. splendens Inflorescence axes broad and compressed, the peduncle 2–2.5 mm wide

Myrcia 59

below the lowest inflorescence branches; leaf blades usually evidently subcordate at base ................................................................. M. subsessilis 53(51). Larger calyx lobes 2–2.5 mm long, pubescent within; flower buds 3–4 mm long ........................................................................................ M. dichasialis 53. Larger calyx lobes 1–1.5 mm long, glabrous within; flower buds 2–2.5 mm long ................................................................................................ M. fallax Myrcia albido-tomentosa (Amshoff) McVaugh, Mem. New York Bot. Gard. 18(2): 79. 1969. —Aulomyrcia albidotomentosa Amshoff, Bull. Torrey Bot. Club 75: 532. 1948. —Chipo-cuy-yek (Arekuna). Aulomyrcia amshoffiana Steyerm., Fieldiana, Bot. 28: 1003. 1957. Shrub or small tree 1–4 m (taller in Guyana and Suriname); leaves coriaceous, the venation inconspicuous, petioles corky-rimose; trichomes of inflorescence relatively long and slender, abundantly whitish-sericeous. Sandy soils in gallery forests, wooded slopes, scrub forests, and rock outcrops, 600– 1400(–2600) m; Bolívar (Cerro Guaiquinima, Gran Sabana), Amazonas (Cerro Coro Coro, Cerro Duida, Cerro Huachamacari, Cerro Sipapo). Guyana, Suriname, Brazil (Amazonas: Serra Aracá). The fruit of Myrcia albido-tomentosa is juicy and edible and has a spicy taste. Myrcia aliena McVaugh, Field Mus. Nat. Hist., Bot. Ser. 13(4.2): 627. 1958. Aulomyrcia chilensis O. Berg, Linnaea 27: 38. 1855, non Myrcia chilensis O. Berg, 1855. Tall tree; leaves mostly glabrous, drying reddish brown; inflorescence many-flowered. Lowland evergreen forests, 100–300 m; Amazonas (Caño Yureba in Río Ventuari basin, Río Cunucunuma). Amazonian Ecuador, Peru, Brazil, and Bolivia. Myrcia amazonica DC., Prodr. 3: 250. 1828. —Curtido, Curtidor, Uraroene (Arekuna). ?Myrcia leptoclada DC., Prodr. 3: 244. 1828. Aulomyrcia hostmanniana O. Berg, Linnaea 27: 42. 1855. Aulomyrcia spruceana O. Berg in Mart., Fl. Bras. 14(1): 76. 1857. Shrub or small tree 2–15(–20) m tall; year-old twigs reddish, with thin bark peel-

ing in small patches; leaves and inflorescence often drying dark reddish brown; one calyx lobe often held erect in bud. In forests, along rivers, in rocky areas, 100–1000 m; widespread in Bolívar and Amazonas. Guyana, Suriname, northern and central Brazil. ◆Fig. 56. The only difference between Myrcia amazonica and M. leptoclada is the presence (M. amazonica) or lack of pubescence (M. leptoclada). Myrcia bolivarensis (Steyerm.) McVaugh, Mem. New York Bot. Gard. 18(2): 81. 1969. —Aulomyrcia bolivarensis Steyerm., Fieldiana, Bot. 28: 1004. 1957. Shrub or small tree 1–4(–8) m tall; young stems and inflorescences appressed-pubescent; leaf blades membranaceous, caudateacuminate; inflorescences few-flowered. Crevices, rocky places, and forested slopes on tepui summits and slopes, (800–)1400–2100 m; Bolívar (Cerro Guaiquinima, Cerro Venamo, Ptari-tepui), Amazonas (widespread on tepuis and granitic domes). Guyana, Peru(?). ◆Fig. 57. Myrcia bonnetiasylvestris (Steyerm.) Steyerm., Ann. Missouri Bot. Gard. 71: 330. 1984. —Gomidesia bonnetiasylvestris Steyerm., Fieldiana, Bot. 28: 1016. 1957. Variable habit shrub 0.3–3 m, rarely a small tree 6–8 m; flowers fragrant, petals white to brown or pinkish. In Bonnetia and Clusia thickets, exposed rocky escarpments, crevices and small gallery forests on tepui summits, 1800–2600 m; Bolívar (Auyán-tepui, Macizo del Chimantá). Endemic. ◆Fig. 58. Two specimens have the same general appearance of Myrcia bonnetiasylvestris, but have larger leaves Myrcia bracteata (Rich.) DC., Prodr. 3: 245. 1828. —Eugenia bracteata Rich., Actes Soc. Hist. Nat. Paris 1: 110. 1792.

60

M YRTACEAE

—Chipoki (Arekuna), Curame, Guayabillo, Kanau, Taribara-yek. Myrcia hirtellaefolia Gleason, Bull. Torrey Bot. Club 58: 411. 1931. Shrub or treelet 1–8 m tall; leaf blades membranous, gradually acuminate at apex; inflorescence yellowish-hirsute, bracts conspicuous, foliaceous; fruit turning from red to black. Commonly occurring in moist, primary and secondary forests, 100–1300 m; widespread in Bolívar and Amazonas in areas with sand. Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil, Bolivia. ◆Fig. 63. Myrcia calycampa Amshoff, Recueil Trav. Bot. Néerl. 39: 153. 1942. —Calycampe latifolia O. Berg, Linnaea 27: 130. 1856, non Myrcia latifolia O. Berg 1855. —Myrtus latifolia (O. Berg) Badillo, Pittier et al, Cat. Fl. Venez. 2: 196. 1947. Calycampe angustifolia O. Berg, Linnaea 27: 131. 1856, non Myrcia angustifolia (O. Berg) Niedenzu 1853. Shrub or small tree to 5(–14) m; upper surface of leaves evidently reticulateveined, the lower usually obscured by the trichomes; flowers cream-colored; inflorescences densely light-brown-pubescent; calyx lobes large, rounded apically. Evergreen lowland forests, ca. 200 m; eastern Bolívar (Río Cuyuní). Guyana, Suriname, Brazil (Pará). Collections of Myrcia calycampa from Suriname and Rio Branco, Brazil, have the petioles much longer and the fruits permanently pubescent. Myrcia citrifolia (Aubl.) Urb., Repert. Spec. Nov. Regni Veg. Beih. 16: 150. 1920. —Myrtus citrifolia Aubl., Hist. Pl. Guiane index p. 20. 1775. —Aulomyrcia citrifolia (Aubl.) Amshoff, Bull. Torrey Bot. Club 75: 531. 1948. Eugenia acetosans Poir. in Lam., Encycl. suppl. 3: 125. 1813. ?Aulomyrcia triflora O. Berg, Linnaea 27: 79. 1855. Shrub or small tree 1–8 m tall; inflorescence cymosely branched, lateral flowers of a cluster often long-pedicellate. Scarce in savannas at middle elevations (600–800 m), in shrub or dwarf forests at higher elevations (1500–1850 m); Bolívar (Auyán-tepui, Cerro

Bolívar, Roraima-tepui, Uaipán-tepui). West Indies, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 61. Collections from above 1000 m in the Guayana Shield differ from normal Myrcia citrifolia in that they have the leaves broadly rounded to subcordate at the base, and dry reddish brown. They would represent Aulomyrcia triflora (Myrcia triflora) if that is a distinct species. Myrcia clausa McVaugh, Mem. New York Bot. Gard. 18(2): 128. 1969. Tree ca. 8 m tall; young stems, leaves, and inflorescence densely appressed-pubescent; leaf blades elliptic, caudate-acuminate; calyx lobes infolded; fruit fleshy, dull orange-yellow. Moist montane forests, 900–1000 m; Bolívar (Cerro Venamo). Endemic. Myrcia clausa is possibly a synonym of Myrcia fenzliana O. Berg. Myrcia clusiifolia (H.B.K.) DC., Prodr. 3: 255. 1828. —Myrtus clusiaefolia H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 138. 1823. Shrub or small tree 1–4 m; flowers white or whitish pink, fragrant; fruit turning shiny purple-black. White-sand savannas, shrub islands in and around savannas, white-sand scrub (bana), gallery forests, 100–200 m; common in southern Amazonas (Río Orinoco and Río Negro basins). Adjacent Colombia, Brazil (Amazonas, to 1200 m on Serra Aracá; Pará). ◆Fig. 62. Myrcia compta McVaugh, Mem. New York Bot. Gard. 18(2): 82. 1969. Shrub or small tree to 5 m tall; indument appressed-yellow-tan; leaf blades rigidly coriaceous, caudate-acuminate; inflorescence pendulous. Savannas and along streams on tepui summits and slopes, 1600–2000 m; Amazonas (Cerro Cuao, Cerro Sipapo). Endemic. Myrcia crispa McVaugh, Mem. New York Bot. Gard. 18(2): 130. 1969. Shrub or small tree 1.5–5 m tall, young stems, lower surface of leaves, stems, and inflorescence with dense reddish brown tomentum turning olive green to grayish with age, upper surface lustrous; leaf blades rigidly coriaceous, margins revolute. Dwarf or scrub forests, gallery forests, along escarpments on

Myrcia 61

tepui summits and slopes, 1300–2100 m; Bolívar (Cerro Duida, Cerro Guanacoco, Cerro Jaua, Cerro Sarisariñama), Amazonas (Cerro Huachamacari, Cerro Parú, Sierra de la Neblina). Endemic. ◆Fig. 65. Myrcia decorticans DC., Prodr. 3: 252. 1828. Shrub or tree to 10 m tall; leaves typically thick-coriaceous, the venation obscure, petioles corky and flaky. Lowland to upland evergreen or semideciduous forests, 200–700 m; Delta Amacuro (Serranía de Imataca), central and northeastern Bolívar. Sucre; Guyana, French Guiana, Brazil. ◆Fig. 68. Myrcia decorticans is difficult to distinguish from Marlierea umbraticola and Myrcia sp. B other than by its large, thickwalled fruits. Myrcia deflexa (Poir.) DC., Prodr. 3: 244. 1828. —Eugenia deflexa Poir. in Lam., Encycl. suppl. 3: 124. 1813. —Myrtus deflexa (Poir.) H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 142. 1823. Myrcia humboltiana DC., Prodr. 3: 256. 1828. Shrub or tree to 10 m tall, variable in amount and color of pubescence; lower surface of leaves minutely pebbled. Lowland to upland evergreen forests, gallery forests, along rivers, 100–1100 m; Bolívar (Gran Sabana), Amazonas (widespread). Lara, Miranda, Táchira, Yaracuy; West Indies, Guyana, Suriname, French Guiana, Amazonian Peru and Brazil (Amazonas, Pará, Rondônia, Roraima). ◆Fig. 67. Myrcia dichasialis McVaugh, Fieldiana, Bot. 29: 190. 1956. Shrub or tree to 6 m tall; leaves ovate; inflorescences with somewhat dichasial branching; flowers relatively large. Lowland savannas, evergreen forests, gallery forests, along riverbanks, 100–200(–900) m; Bolívar (Gran Sabana, Río Parguaza), Amazonas (widespread). Amazonian Colombia, Peru, and Brazil. ◆Fig. 60. Myrcia ehrenbergiana (O. Berg) McVaugh, Mem. New York Bot. Gard. 18(2): 85. 1969. —Myrciaria ehrenbergiana O. Berg, Linnaea 27: 321. 1856.

Shrub or small tree 3–4 m tall; leaves subsessile, cordate, oblong; tertiary venation evident, reticulate; inflorescences manyflowered, appressed-pubescent; flowers 4merous. Savannas, along gallery forests, 700–1100 m; Bolívar (Gran Sabana near Roraima-tepui, Sierra Pakaraima). Adjacent Guyana and Brazil (Roraima). Myrcia exploratoris McVaugh, Mem. New York Bot. Gard. 18(2): 86. 1969. Shrub or tree to 8 m tall; leaves coriaceous, rounded at apex, upper surface lustrous; inflorescences few- to many-flowered; fruit with spicy odor, purple-black at maturity. Wooded slopes and dense dwarf forests on upper tepui slopes and summits, 1800– 2500 m; Bolívar (Carrao-tepui, Cerro Jaua, Ilú-tepui, Karaurín-tepui, Sierra de Maigualida, Sororopán-tepui), Amazonas (Serranía Uasadi, Sierra de Maigualida). Guyana (Mount Wokomung), Suriname (Tafelberg). ◆Fig. 59. Myrcia fallax (Rich.) DC., Prodr. 3: 244. 1828. —Eugenia fallax Rich., Actes Soc. Hist. Nat. Paris 1: 110. 1792. Myrcia kegeliana O. Berg, Linnaea 27: 99. 1855. Myrcia spruceana O. Berg in Mart., Fl. Bras. 14(1): 165. 1857. Myrcia aguitensis Gleason, Bull. Torrey Bot. Club 58: 409. 1931. Shrub or tree to 12 m tall; leaves typically ovate-elliptic, acuminate at apex. Occurring in a wide range of habitats, from disturbed to primary areas, from savannas to forests, and from lowland to upland areas, 50–1700 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Apure, Aragua, Distrito Federal, Falcón, Lara, Mérida, Táchira, Yaracuy, Zulia; Lesser Antilles, Panama, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Rondonia, Pará, Acre, Amazonas, Roraima). ◆Fig. 70. The type of Myrcia ayresiana O. Berg, Ayres 872, collected from the banks of the Río Orinoco, was apparently destroyed in Berlin. It was compared by O. Berg to Myrcia kegeliana and M. berberis DC., both of which are synonyms of M. fallax. Myrcia compressa Gleason (holotype Tate 834, NY) is another poorly known species that may be a synonym of M. fallax.

62

M YRTACEAE

Myrcia fenzliana O. Berg in Mart., Fl. Bras. 14(1): 196. 1857. Gomidesia lindeniana O. Berg, Linnaea 29: 208. 1858. Shrub or treelet to 2–7 m; leaves often strongly revolute, softly pubescent; fruits yellow-orange turning dark purple at maturity, pilose, crowned by the erect or incurved calyx lobes. Upland thickets in savannas and gallery forests, 1100–1300 m; Bolívar (Gran Sabana). Mérida, Táchira; West Indies, Colombia (Santander), Guyana, Brazil (Distrito Federal, Minas Gerais), Bolivia. Myrcia gentryi B. Holst, Biollania 10: 4. 1994. Shrub to 2 m tall; young growth and flowers white-sericeous; upper surface of leaf blade impressed-punctate, the midvein impressed and higher-order venation obscure; inflorescence 1- or 3-flowered. Gallery forests and open areas on tepui summit, ca. 1700 m; Amazonas (Cerro Huachamacari). Endemic. ◆Fig. 81. Myrcia grandis McVaugh, Mem. New York Bot. Gard. 18(2): 114. 1969. Shrub or small tree 1–8 m, twigs ash gray; leaves ovate, abruptly acuminate, the secondary venation inconspicuous; mature fruit purple-black. Along seasonally flooded blackwater rivers, in white-sand savannas, around clearings along rivers and in forests, 100–200 m; western Amazonas. Guyana, Amazonian Colombia and Brazil. ◆Fig. 73. Myrcia guianensis (Aubl.) DC., Prodr. 3: 245. 1828. —Eugenia guianensis Aubl., Hist. Pl. Guiane 506, t. 201. 1775. Panama, Colombia, Venezuela, TrinidadTobago, Guyana, Suriname, Peru, Amazonian Brazil, Bolivia; 2 varieties, both in Venezuela, 1 of these in the flora area. Myrcia guianensis is one of the most commonly collected, widespread, and variable species of Myrcia in northern South America. The typical variety occurs widely in the Venezuelan Guayana. Variety cuneata (O. Berg) McVaugh is limited to cloud forests of the coastal mountain range of Venezuela. M. guianensis var. guianensis. —Farasu (Yekwana), Jujuli, (Guahibo). Myrcia obtusa Schauer, Linnaea 21. 272.

1848. —Aulomyrcia obtusa (Schaeur) O. Berg, Linnaea 27: 64. 1855. Aulomyrcia roraimensis O. Berg, Linnaea 27: 68. 1855. Myrcia roraimae Oliv. ex Thurn, Timehri 5: 192. 1886. —Aulomyrcia roraimae (Oliv. ex Thurn) Steyerm., Fieldiana, Bot. 28: 1007. 1957. Shrub 1–4 m tall or small tree 5–10 m; leaves ovate to obovate, usually obtuse to acutish at apex; flowers fragrant, white, creamy white, or pinkish. Shrub and tree islands in savannas, on and around granitic outcrops, gallery forests, wooded backwater areas, wooded slopes and roadside scrub, 50– 1300 m; widespread in Delta Amacuro, Bolívar, and Amazonas. North-central Venezuela (Anzoátegui, Aragua, Cojedes, Miranda, Sucre, Táchira, Trujillo); Panama, Colombia, Trinidad-Tobago, Guyana, Suriname, Peru, Amazonian Brazil, Bolivia. ◆Fig. 75. Populations occurring in the Venezuelan Guayana are relatively uniform except for two that occur in southern Amazonas: (1) high-altitude (1700–2000 m) on Sierra de la Neblina, with inflorescence reduced to a few flowers and leaves smaller (e.g., Nee 30788, SEL); (2) large-leaved, mostly glabrous population on the upper Río Negro (e.g., Stergios & Aymard 4114, SEL). Myrcia inaequiloba (DC.) Legrand, Atas do Simposio sobre a Biota Amazonica 149. “1967” [1968, sem 1]; McVaugh, Taxon 17: 378. 27 Aug 1968. —Eugenia inaequiloba DC., Prodr. 3: 282. 1828. —Chunchunillo blanco, Jarasa, Kumachi (Yekwana), Mahchó kaí mën, Maybapio-keu-yek (Arekuna), Tupurú ke tipeñ mën (Piaroa), Yucú. Eugenia ?nitida Benth., J. Bot. (Hooker) 2: 322. 1840. —Myrciaria nitida (Benth.) O. Berg, Linnaea 27: 324. 1856. ?Aulomyrcia paniculata O. Berg, Linnaea 27: 49. 1855. —Aulomyrcia inaequiloba var. paniculata (O. Berg) Amshoff, Recueil Trav. Bot. Néerl. 42: 7. 1950. Shrub 1–5 m or tree 3–10(–15) m tall; leaf blades elliptic and abruptly and shortly acuminate to ovate and acuminate; inflorescence many-flowered, sparsely to densely puberulent, at least some of the bracts persistent into fruit; flowers fragrant. Common along rivers with sandy or rocky shores, on

Myrcia 63

and around granitic outcrops, dwarf forests on sandstone, gallery forests, dry riparian woodlands, rocky shrub savannas, edges of forests, 100–1300 m; widespread in Bolívar and Amazonas (to 500 m). Panama, Colombia (Choco), Guyana, Suriname, French Guiana, Brazil (Roraima). ◆Fig. 66. Almost all of the Venezuelan collections of Myrcia inaequiloba differ from the type collection from Suriname, which has the petioles short and corky-rimose. The Venezuelan collections have petioles that are long and smooth and would be referrable to Aulomyrcia inaequiloba (Myrcia inaequiloba) var. paniculata if that proves to be distinct. Myrcia induta McVaugh, Mem. New York Bot. Gard. 18(2): 99. 1969. Tree ca. 4 m tall; branchlets and inflorescence densely appressed-hirsute; leaves caudate-acuminate, margins revolute, upper surface impressed-punctate; inflorescence < 2 cm long; flowers 4-merous. Escarpments on tepui summits, ca. 2000 m; Amazonas (Cerro Parú). Endemic. Myrcia intonsa (McVaugh) B. Holst, Selbyana 23: 152. 2002. —Marlierea intonsa McVaugh, Mem. New York Bot. Gard. 10(1): 85. 1958. Tree 8–12 m tall; petioles stout; leaf blades ovate to elliptic, densely golden sericeous on lower surface; fruits round, densely pubescent; flowers and fruits large for the genus. Bases of tepui bluffs, gallery forests, 1100–1700 m; Bolívar (near Kavanayén, Macizo del Chimantá [Chimantá-tepui]). Brazil (Amazonas: Serra Aracá). Myrcia kylistophylla B. Holst, Selbyana 23: 154. 2002. Shrub 1–3 m tall; young stems whitish-tomentose; leaves strongly revolute, the lower surface greenish white, the upper surface impressed-punctate; inflorescences few-flowered. Tepui summits on rocky outcrops or along streams, 1900–2100 m; Bolívar (Cerro Jaua, Sierra de Maigualida). Endemic. Myrcia liesneri B. Holst, Selbyana 23: 154. 2002. Tree to 4 m tall; leaves stiff-coriaceous, short-petiolate, subcordate, the midvein prominently convex to biconvex on upper

surface; inflorescence few-flowered; flowers obscurely but densely puberulent without; fruits slightly oblate. Lowland evergreen moist forests, 100–400 m; Amazonas (near La Esmeralda, base of Sierra de la Neblina). Endemic. Myrcia lucida McVaugh, Mem. New York Bot. Gard. 18(2): 100. 1969. —Myrcia laevis O. Berg, Linnaea 31: 252. 1862, non M. laevis G. Don 1832.—Curame (Baniva), Dumatsi, Guayabito, Iwapichuna. Myrcia lucida var. attenuata McVaugh, Mem. New York Bot. Gard. 18(2): 100. 1969. Shrub 1–2 m or tree 3–10 m tall; leaves usually lustrous and appearing polished on upper surface; flowers fragrant. Sandy or rocky areas along rivers, seasonally flooded forests, shrub islands in savannas, 100–200 m; Bolívar (lower Río Parguaza), Amazonas (Rio Cataniapo, Río Negro basin). Apure; Amazonian Colombia and Brazil (Amazonas, Mato Grosso), Bolivia. ◆Fig. 76. Myrcia magnoliifolia DC., Prodr 3: 248. 1828, “magnoliaefolia.” Shrub or tree to 7 m, abundantly pilose; leaf blades large for the genus, to 20 cm long. Lowland to upland evergreen forests or disturbed areas, along river banks, 100–1200 m; Bolívar (Gran Sabana), Amazonas (widespread). Guyana, Suriname, Amazonian Brazil and Bolivia. ◆Fig. 72. Myrcia minutiflora Sagot, Ann. Sci. Nat. Bot. sér. 6, 20: 185. 1885. Shrub or tree 1–10 m tall; leaves prominently caudate-acuminate, the marginal vein arching between and connecting the secondary veins; inflorescences few-flowered. Understory plant of forests, tepui summits, 200–1800 m; Bolívar (Cerro Jaua, Río Antavari, upper Río Caura, Rio Paragua, upper Río Suapure), Amazonas (Río Mavaca). Guyana, Suriname, French Guiana, Amazonian Peru, Brazil (Amapa, Rondônia, Roraima, Amazonas, Acre, Marahnao, Pará). ◆Fig. 74. Myrcia multiflora (Lam.) DC., Prodr. 3: 244. 1828. —Eugenia multiflora Lam., Encycl. 3: 302. 1789.

64

M YRTACEAE

Aulomyrcia vinacea Steyerm., Fieldiana, Bot. 28: 1008. 1957. Shrub or small tree 3–4 m tall; globe of the corolla in bud much enlarged over the calyx; leaf blades membranaceous, the lower surface venation often with a reddish tinge. Savannas, rocky slopes, gallery forests, 100– 500 m; northern Bolívar, Amazonas (Puerto Ayacucho). Falcón, Sucre; widespread elsewhere in South America east of the Andes. ◆Fig. 71. Myrcia nubicola McVaugh, Mem. New York Bot. Gard. 18(2): 116. 1969. Shrub or small tree 1–8(–15) m tall; leaves stiff-coriaceous, mostly obovate, the secondary venation inconspicuous; flowers fragrant, inflorescence coppery-sericeous. Along streams and escarpments, dwarf or scrub forests on tepui summits, forested areas on tepui slopes, 1000–2300 m; Bolívar (Auyántepui, Cerro Guaiquinima, Cerro Jaua, Macizo del Chimantá, Sierra Pakaraima), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú, Sierra Parima, Sierra de la Neblina). Guyana (Mount Ayanganna). Myrcia paivae O. Berg in Mart., Fl. Bras. 14(1): 179. 1857. Shrub or tree to 12 m tall; leaf blades usually elliptic, prominently acuminate, impressed-punctate on upper surface; inflorescences largely axillary. Moist evergreen lowland to upland forests, 100–1200 m; widespread in Bolívar and Amazonas. Anzoátegui, Apure, Barinas, Miranda, Táchira, Trujillo; Panama, Colombia, Guyana, Amazonian Peru, Brazil, and Bolivia. Myrcia pistrinalis McVaugh, Mem. New York Bot. Gard. 18(2): 117. 1969. Small trees to ca. 8 m tall; leaf blades membranous; inflorescence few-flowered, sparsely branched; hypanthium covered with minute, erect, dense trichomes; pedicels 4–10 mm long, stout in fruit. Lower montane forests, 800–1100 m; Bolívar (southern Gran Sabana), Amazonas (Sierra Parima). Suriname. Myrcia pistrinalis is apparently uncommon, though it is fairly nondescript and is perhaps overlooked in herbaria.

Myrcia platyclada DC., Prodr. 3: 244. 1828. —Aulomyrcia platyclada (DC.) Amshoff, Bull. Torrey Bot. Club 75: 531. 1948. Aulomyrcia dumosa O. Berg, Linnaea 30: 656. 1861. —Myrcia dumosa (O. Berg) Krug & Urb., Bot. Jahrb. Syst. 19: 580. 1895. Aulomyrcia edulis O. Berg, Linnaea 30: 657. 1861. —Myrcia edulis (O. Berg) Krug & Urb., Bot. Jahrb. Syst. 19: 581. 1895. Aulomyrcia platyclada var. kaieteurensis Amshoff, Bull. Torrey Bot. Club 75: 532. 1948. Shrub 1–4 m or tree 10–15 m tall; leaves coriaceous, venation evident or obscured in thick-leaved forms, often broadly rounded to obtuse at apex; flower buds turbinate. Lowland to upland forests, shrub thickets in savannas, 400–1800 m; Bolívar (Auyán-tepui, Gran Sabana, Serranía de Imataca). Lesser Antilles, Guyana, Suriname, French Guiana, Brazil (Pará). ◆Fig. 79. A collection from Cerro Parú (Cowan & Wurdack 31122, MICH, NY, US) resembles Myrcia platyclada vegetatively, but has 2 inflorescence branches per node, a feature more characteristic of Calyptranthes. Glabrous forms of Marlierea summa are difficult to differentiate from Myrcia platyclada. Myrcia ptariensis (Steyerm.) McVaugh, Mem. New York Bot. Gard. 18(2): 103. 1969. —Aulomyrcia ptariensis Steyerm., Fieldiana, Bot. 28: 1006. 1957. Shrub 1–5 m tall; flowers white to pinkish, fragrant, abundantly and softly white- to light coppery-tomentose; leaf blade margins revolute, the secondary venation inconspicuous. Shrub islands, gallery forests, Bonnetia roraimae forests, open rocky slopes, tepui summits and slopes, 1700–2300 m elevation; Bolívar (Auyán-tepui, Kamarkawarai-tepui, Macizo del Chimantá, Ptari-tepui). Endemic. ◆Fig. 78. Myrcia pyrifolia (Desv. ex Ham.) Nied. in Engl. & Prantl, Nat. Pflanzenf. 3(7): 76. 1893. —Eugenia pyrifolia Desv. ex Ham., Prodr. Pl. Ind. Occid. 44. 1825. —Chipo-yek (Arekuna), Farasu (Yekwana), Ha-dá-sa, Jadasa. Shrub or tree 2–5(–8) m tall; leaves drying dark brownish, midvein convex on upper sur-

Myrcia 65

face; inflorescences many flowered, the flowers fragrant, hypanthium white-sericeous. Sandy, rocky, open or forested areas along rivers or lagoons, 100–500 m; Bolívar (Río Canaracuni, Río Caroní, Río Carrao, Río Carún, Río Caura, Río Paragua), Amazonas (Río Casiquiare, Río Cunucunuma, Río Guayapo, junction of Río Orinoco and Río Ventuari, Río Sipapo). Colombia, Guyana, Suriname, French Guiana, Amazonian Peru, Brazil (Amazonas, Pará). Collections of Myrcia pyrifolia from the Río Cunucunuma region of Amazonas have longer petioles and narrower leaf bases than populations from other localities. Myrcia quitarensis (Benth.) Sagot, Ann. Sci. Nat. Bot. sér. 6, 20: 184. 1885. —Eugenia quitarensis Benth., J. Bot. (Hooker) 2: 322. 1840. —Myrciaria quitarensis (Benth.) O. Berg, Linnaea 27: 323. 1854. Rare tree of uncertain height, bark smooth; leaf blades membranous, midvein plane to slightly convex, the upper surface impressed-punctate; inflorescence sparsely reddish-pubescent, branching at 90° angles. Along highway, most likely in evergreen forests, ca. 200 m; Bolívar (Río Cuyuní basin). Guyana (Río Quitaro). Myrcia revolutifolia McVaugh, Mem. New York Bot. Gard. 18(2): 121. 1969. Myrcia planifolia McVaugh, Mem. New York Gard. 18(2): 118. 1969. Dense to wiry shrub 0.5–4 m tall or tree 5–10 m; bark smooth with reddish or coppery patches; leaf blades coriaceous, rounded to retuse at apex, the secondary veins fine, parallel, numerous; branches of inflorescence frequently reddish. Occurring in a wide range of habitats from lowlands to tepui summits, but always associated with sandstone, shrub thickets in white-sand savannas, open or dense dwarf forests, along small streams, 100–2000 m; Bolívar (Cerro Sarisariñama), Amazonas (widespread). Eastern Colombia, Brazil (Amazonas: Serra Aracá). ◆Fig. 69. Myrcia rotundata (Amshoff) McVaugh, Mem. New York Bot. Gard. 18(2): 122. 1969. —Aulomyrcia rotundata Amshoff, Recueil Trav. Bot. Néerl. 42: 8. 1950.

Spindly to bushy shrub 0.5–6 m tall or small tree 3–8 m; young leaves, branches of inflorescence, and calyx lobes reddish green to maroon. Dwarf Bonnetia roraimae forests, moist dwarf forests, on flat rocky areas with sparse shrubs, and exposed sandstone bluffs, tepui summits and slopes, 500–2200 m. Southern Venezuela, Guyana; 2 varieties, both in the flora area. Key to the Varieties of M. rotundata 1. Leaf blades bluntly rounded to bluntly acuminate; inflorescence 3-flowered ........................................... var. atrans 1. Leaf blades broadly rounded to emarginate at the apex; inflorescences to 40flowered ....................... var. rotundata M. rotundata var. atrans McVaugh, Mem. New York Bot. Gard. 18(2): 122. 1969. Spindly or bushy shrub 0.5–3 m tall. Areas with exposed sandstone and low, open vegetation, 1700–2200 m; Bolívar (Auyántepui). Endemic. M. rotundata var. rotundata Aulomyrcia parvifolia Steyerm., Fieldiana, Bot. 28: 1006. 1957. Shrub or tree 1–8 m tall. Bonnetia roraimae and other dwarf, moist forests, slopes and summits of tepuis, 500–2100; Bolívar (Cerro Venamo, Murisipán-tepui, Ptari-tepui, Uei-tepui), Amazonas (Cerro Sipapo). Guyana. ◆Fig. 77. Myrcia salticola (Steyerm.) McVaugh, Mem. New York Bot. Gard. 18: 123. 1969. —Aulomyrcia salticola Steyerm., Fieldiana, Bot. 28: 1007. 1957. —Ai-yek (Arekuna). Shrub ca. 4.5 m tall; leaves strongly bicolorous, midvein convex on upper surface; calyx lobes pubescent within. Montane forests, ca. 1600 m, Bolívar (Sororopán-tepui). Endemic. Myrcia servata McVaugh, Mem. New York Bot. Gard. 18(2): 139. 1969 Myrcia schomburgkiana O. Berg, Linnaea 27: 110. 1855, nom. illeg. Tree 8–15 m tall; leaf blades relatively narrow; calyx lobes pubescent on both surfaces. Cloud forests, moist evergreen forests,

66

M YRTACEAE

(100–)500–1300 m; Delta Amacuro (Santa Catalina), Bolívar (Altiplanicie de Nuria, Cerro Venamo), Amazonas (Sierra Parima). Distrito Federal; Guyana. ◆Fig. 64. Myrcia servata is a poorly understood species in the M. fallax alliance. Myrcia sessiliflora McVaugh, Mem. New York Bot. Gard. 18(2): 105. 1969. Shrub 1–3 m tall; young stems, lower surface of leaves, and fruits gray- or gray-greenpubescent, upper surface impressed-punctate, the venation obscure; leaf blades abruptly acuminate-caudate; inflorescence sessile, or nearly so. Along streams or on moist escarpments, 1400–1800 m; Bolívar (Cerro Guanay), Amazonas (Cerro Sipapo, Sierra Uasadi). Endemic. Myrcia sessiliflora is possibly conspecific with M. induta McVaugh, but too few collections are available to be certain. Myrcia sipapensis McVaugh, Mem. New York Bot. Gard. 18: 140. 1969. Tree to 15 m tall, young stems, leaves on lower surface, and inflorescence covered with dark brown pubescence; upper surface of leaves glabrous, drying dark brown; petioles stout; inflorescences many-flowered. Slope forests or rocky areas on tepuis, 1700–2000 m; Amazonas (Cerro Sipapo, Sierra de la Neblina). Endemic. Myrcia sororopanensis Steyerm., Fieldiana, Bot. 28: 1019. 1957. Shrub or tree to 8 m tall; vegetative growth and inflorescence densely hirsutulous. Montane evergreen forests, dwarf tepui forests, along rivers, 1200–2300 m; Bolívar (widespread on tepui summits and slopes), Amazonas (Sierra Parima). Guyana. ◆Fig. 80. Myrcia splendens (Sw.) DC., Prodr. 3: 244. 1828. —Myrtus splendens Sw., Prodr. 79. 1788. Shrub or tree to 10 m tall; leaves shortpetiolate, mostly ovate; inflorescences pubescent. Lowland to highland evergreen forests, along rivers, rocky savannas, 100–2100 m; Delta Amacuro (Río Acure), Bolívar (Gran Sabana, Ptari-tepui), Amazonas (widespread). Anzoátegui, Aragua, Distrito Federal, Sucre, Táchira, Zulia; Central America,

West Indies, Colombia, Tobago, Suriname, Peru, Brazil (Amapa, Amazonas, Mato Grosso), Bolivia. ◆Fig. 82. Myrcia subsessilis O. Berg, Linnaea 31: 251. 1862. Colombia, Venezuela, French Guiana, Brazil (Amapá, Pará), Bolivia; 2 varieties, both in the flora area. Key to the Varieties of M. subsessilis 1. Leaf blades elliptic, gradually narrowed to the base; calyx lobes glabrous within ............................................ var. ovalis 1. Leaf blades ovate, subcordate; calyx lobes pubescent within ....... var. subsessilis M.

subsessilis var. ovalis (O. Berg) McVaugh, Mem. New York Bot. Gard. 18(2): 144. 1969. —Myrcia subsessilis ß ovalis O. Berg, Linnaea 31: 252. 1862. Shrub or tree to 8 m tall. Lowland to upland evergreen forests, Clusia scrub forests, granitic outcrops, 100–1100 m; southern Amazonas. Endemic. M. subsessilis var. subsessilis Myrcia subsessilis var. subcordata O. Berg, Linnaea 31: 251. 1862. Shrub or tree to 6 m. Lowland to upland evergreen forests, seasonally flooded forests, shrub islands in savannas, gallery forests, secondary forests, granitic outcrops, 100– 300(–900) m; Bolívar (northwestern portions, Cerro Jaua), Amazonas (widespread). Apure; Colombia, French Guiana, Brazil (Amazonas, Pará), Bolivia. ◆Fig. 84. Myrcia sylvatica (G. Mey.) DC., Prodr. 3: 244. 1828. —Myrtus sylvatica G. Mey., Prim. Fl. Esseq. 191. 1818. Shrub or tree to 5 m tall; leaf blades stiffcoriaceous, the upper surface impressedpunctate, midvein prominent and deeply impressed, the secondary venation inconspicuous, margins revolute. Lowland to highland, evergreen to subsclerophyllous forests, gallery forests, scrubland, shrub savannas, along creeks, 100–2200 m; widespread in Bolívar and Amazonas. Anzoátegui, Sucre, Táchira; Panama, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Acre, Amapa, Amazonas, Pará, Roraima).

Myrcia 67

Myrcia sylvatica is one of the most widespread and commonly collected species of Myrtaceae in South America. It is often found in nutrient-poor, sandy soils. Myrcia tepuiensis Steyerm., Fieldiana, Bot. 28: 1019. 1957. Shrub 1.5–5 m tall; leaf blades small, 3–7 cm long, conspicuously reticulate, apex broadly obtuse to subacuminate, petioles short, 2–4(–6) mm long. Upper slopes and summits of tepuis, 1500–2600 m; Bolívar (Ilú-tepui, Ptari-tepui, Sororopán-tepui), Amazonas (Cerro Sipapo, Cerro Yutajé). Endemic. ◆Fig. 85. Myrcia tomentosa (Aubl.) DC., Prodr. 3: 245. 1828. —Eugenia tomentosa Aubl., Hist. Pl. Guiane 504, t. 200. 1775. —Aulomyrcia tomentosa (Aubl.) Amshoff, Recueil Trav. Bot. Néerl. 39: 153. 1942. Shrub or small tree 1.5–10 m tall; young twigs and inflorescences densely villous; leaves softly pilose, obovate, the secondary veins slightly impressed on upper surface and evident on lower surface; sepals unequal. Thickets, riparian forests, ecotone between forests and savannas, 100–900 m; Bolívar (Gran Sabana, Lago Guri, Río Aro, lower Río Paragua). Aragua, Distrito Federal, Falcón, Lara, Miranda, Monagas, Sucre; Panama, West Indies, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 83. Myrcia tomentosa is a variable species with many leaf sizes and textures and a variable amount of pubescence. When a thorough study of the species complex to which M. tomentosa belongs is completed, many more names, especially from Brazil, will be added to the list of synonomy. Myrcia sp. A Tree ca. 8 m tall; stems stout; leaves broadly elliptic, drying dark brown on upper surface, olive green on lower surface, the midvein impressed on upper surface; inflorescence many-flowered, sparsely yellowishsericeous; fruit unknown. Montane forest, ca. 1300 m; Amazonas (Sierra de la Neblina). Endemic. This taxon is known from a single collection that is between flower and fruit (Nee 31142, ASU, NY). It is distinct because of its

bicolorous leaves and many-flowered inflorescence, but insufficient material is available for a full description. Myrcia sp. B Shrub; leaves stiff-coriaceous, mostly obovate and rounded to obtuse at apex, the petioles corky-rimose; inflorescence bracts persistent. Scrub forests on white-sand, 100– 200 m; southern Amazonas. Northern Amazonian Brazil. This is apparently a common shrub in white-sand areas, but belongs to a complex of species in need of revision. Closely related species are Myrcia decorticans and Marlierea umbraticola. Myrcia sp. C Slender tree to 8 m tall; stem and inflorescence axes ferruginous- or brownishvelutinous; leaves rounded to broadly ovate, the secondary venation impressed on upper surface. Evergreen tepui-summit forests, 1700–1800 m; Bolívar (Auyán-tepui). Endemic. This distinct taxon is known from two collections, one sterile (Holst 3824, MO, VEN) and one in fruit (Steyermark 93609, MICH, NY, US, VEN); flowers are needed for a complete description. Myrcia sp. D Shrub to 2 m, few-stemmed; leaves stiffcoriaceous, round, to broadly elliptic to broadly ovate, rounded at base and rounded to shallowly emarginate at apex, secondary veins numerous, midvein shallowly sulcate; inflorescences subterminal; inflorescences sparsely flowered. Tepui scrub forests, 1200–1300 m; Bolívar (Cerro Guaiquinima). Endemic. As Rogers McVaugh pointed out on an annotation label in 1979 (Steyermark et al. 117425, MICH), this collection likely represents an undescribed species. However, the few flowers remaining on the inflorescence of the only known specimen are well past anthesis. This taxon would key out in the neighborhood of Myrcia multiflora or M. platyclada, but differs from them by having round to broadly oblong leaf blades (versus ovate, elliptic, or obovate).

68

M YRTACEAE

Fig. 56. Myrcia amazonica

Fig. 57. Myrcia bolivarensis

Myrcia 69

Fig. 58. Myrcia bonnetiasylvestris

Fig. 60. Myrcia dichasialis

Fig. 59. Myrcia exploratoris

70

M YRTACEAE

Fig. 61. Myrcia citrifolia Fig. 62. Myrcia clusiifolia

Fig. 63. Myrcia bracteata

Myrcia 71

Fig. 64. Myrcia servata

Fig. 65. Myrcia crispa

Fig. 66. Myrcia inaequiloba

72

M YRTACEAE

Fig. 67. Myrcia deflexa

Fig. 68. Myrcia decorticans

Myrcia 73

Fig. 69. Myrcia revolutifolia

Fig. 70. Myrcia fallax

74

M YRTACEAE

Fig. 71. Myrcia multiflora

Fig. 72. Myrcia magnoliifolia

Fig. 73. Myrcia grandis

Myrcia 75

Fig. 74. Myrcia minutiflora

Fig. 76. Myrcia lucida

Fig. 75. Myrcia guianensis var. guianensis

76

M YRTACEAE

Fig. 77. Myrcia rotundata var. rotundata

Fig. 78. Myrcia ptariensis

Fig. 79. Myrcia platyclada

Myrcia 77

Fig. 80. Myrcia sororopanensis

Fig. 81. Myrcia gentryi

78

M YRTACEAE

Fig. 82. Myrcia splendens

Fig. 84. Myrcia subsessilis var. subsessilis

Fig. 83. Myrcia tomentosa

Fig. 85. Myrcia tepuiensis

Myrcianthes 79

9. MYRCIANTHES O. Berg, Linnaea 27: 315. 1856. [Subtribe Eugeniinae]. Anamomis Griseb., Fl. Brit. W. I. 240. 1860. by Francesa Grifo Shrubs or trees to 40 m, larger trees with buttressed trunks; bark exfoliating, often reddish in color. Leaves cartilaginous or coriaceous, opposite or ternate; petioles often channeled; blade margins sometimes revolute; midvein often impressed on upper surface, marginal vein evident. Inflorescence a dichasium, simple or much branched and sometimes resembling a panicle, 1–31-flowered, the terminal flowers sessile and the lateral flowers pedicellate; peduncles 0.4–9 cm long, often flattened distally or somewhat compressed; bracts and bracteoles mostly linear or lanceolate, sometimes subulate or narrowly triangular, scarious, mostly early deciduous. Hypanthium not prolonged beyond ovary summit, mostly smooth, sometimes 4-angled, broadly obconic or campanulate. Flowers 4- or 5-merous; calyx lobes distinct, imbricate, in equal or unequal pairs, persistent; petals imbricate, reflexed at anthesis, white or greenish white or rarely white aging pink; disk quadrangular or round. Stamens 50–300; anthers longitudinally dehiscent. Ovary 2-locular (3-locular in M. prodigiosa); ovules (5–)10–15(–30); style 5–12 mm long; stigma occasionally capitate, never peltate. Fruit spherical, to 6 cm diameter, yellow, orange, red, or deep purple-black; some sweet with abundant flesh, others insipid. Seeds 1–4, reniform, to 4 cm long; testa light brown, normally paper-like and peeling easily or leathery (M. prodigiosa); cotyledons 2, plano-convex, distinct except where joined apically, large, fleshy; radicle usually about as long as the cotyledons; plumule evident in mature seeds, often sericeous, much shorter than radicle. U.S.A. (Florida), Mexico, Central America, West Indies, South America south to Argentina (most diverse in the Andes); ca. 50 species, 15 in Venezuela, 3 of these in the flora area. Key to the Species of Myrcianthes 1. 1. 2(1). 2.

Ovaries densely gray-pubescent; leaves and inflorescences mostly 2–5 cm long; petioles and peduncles thin ............................................. M. fragrans Ovaries glabrous or thinly pubescent; leaves and inflorescences mostly 7–12 cm long; petioles and peduncles stout ......................................... 2 Flowers ca. 7 mm wide at anthesis; fruits ca. 4 cm diameter; ovary trilocular ................................................................................ M. prodigiosa Flowers ca. 6 mm wide at anthesis; fruits 1–2 cm diameter; ovary bilocular ................................................................................ M. rhopaloides

Myrcianthes fragrans (Sw.) McVaugh, Fieldiana, Bot. 29(8): 485. 1963. —Myrtus fragrans Sw., Prodr. 79. 1788. —Arrayán, Guayabillo. Eugenia steyermarkii Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 360. 1940. Eugenia rondonensis Steyerm., Fieldiana, Bot. 28: 1013. 1957. Eugenia turumiquirensis Steyerm., Fieldiana, Bot. 28: 1015. 1957.

Shrub or tree to 15 m tall; leaves coriaceous, usually densely black-glandular-punctate; mature fruits orange. Tall, lowland to montane evergreen forests, 200–2200 m; Bolívar (near El Dorado, Macizo del Chimantá [Agparamán-tepui], Roraima-tepui). Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Sucre, Trujillo, Zulia; U.S.A. (southern Florida), Bahamas, Turks and Caicos, Antilles, Mexico,

80

M YRTACEAE

Fig. 86. Myrcianthes prodigiosa

Central America, Colombia, Ecuador, Peru. ◆Fig. 87. Myrcianthes prodigiosa McVaugh, Fieldiana, Bot. 29(8): 492. 1963. Treelet or tree to 15 m tall; leaves thickleathery, secondary venation obscure; mature fruits unknown (immature fruits known to be 4+ cm diameter). Evergreen lowland to montane forests, (100–)1000–1600 m; Amazonas (Cerro Duida, Cerro Parú, Río Sipapo). Guyana, Suriname, Ecuador. ◆Fig. 86.

Fig. 87. Myrcianthes fragrans

Myrcianthes rhopaloides (H.B.K.) McVaugh, Publ. Field Mus. Nat. Hist., Bot. 13(4): 771. 1958. —Myrtus rhopaloides H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 137. 1823. Tree to 8 m tall; leaves coriaceous, secondary venation prominent; mature fruits purple-black. Evergreen montane forests, 1800–2000 m; Bolívar (Macizo del Chimantá [Apacará-tepui]). Mérida; Costa Rica, Panama, Colombia, Ecuador, Peru, Bolivia.

Myrciaria 81

10. MYRCIARIA O. Berg, Linnaea 27: 320. 1856. [Subtribe Eugeniinae]. by Bruce K. Holst Shrubs or small trees, bark guava-like. Leaf-blade midvein convex or flat on upper surface, primary veins numerous, mostly parallel. Inflorescence a much-reduced bracteate shoot, appearing glomerate, axillary in the leaves or cauliflorous; bracteoles enclosing the ovary, often connate. Flowers 4-merous, sessile to shortpedicellate; hypanthium elongated above the ovary, circumscissile at point of attachment with ovary. Petals small; calyx open in bud; petals and stamens borne on hypanthium. Stamens many; filaments incurved in bud; anthers ovate. Ovary bilocular; ovules usually 2 per locule, ascending. Fruit globose, the hypanthium scar evident as a ring. Seeds 1–3; embryo eugenioid, undivided. Central America, West Indies, South America mostly east of the Andes; ca. 20 species, 7 in Venezuela, 6 of these in the flora area. In the abscence of flowers, the leaf venation described above can often be used to help identify the genus. In fruit, the genus can usually be recognized by the abscence of any calyx lobes or hypanthium, and when dry, the fruits have a bumpy appearance due to the large surface glands. Many of the species have juicy fruits that are widely eaten fresh or used in preserves and beverages. Myrciaria dubia is especially noteworthy for the high ascorbic acid content of its fruits. Key to the Species of Myrciaria 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(3). 5.

Leaves < 2 cm long, obtuse to rounded at apex .............................. M. tenella Leaves > 3 cm long, acute to acuminate at apex ...................................... 2 Inflorescence axis 4–5 mm long; bracts 3 mm long, exceeding the summit of the ovary of the accompanying flower ........................... M. puberulenta Inflorescence axis 0–2 mm long; bracts < 1 mm long, not exceeding the summit of the ovary of the accompanying flower................................. 3 Leaves shallowly cordate at base .............................................................. 4 Leaves acute to rounded at base ............................................................... 5 Leaf apex obtuse or rounded; petiole glabrous ............................. M. cordata Leaf apex long-acuminate; petiole puberulent ........................ M. vismeifolia Flower buds 3 mm wide; leaves acute to acuminate; mature fruits (when dry) > 1.5 cm diameter .................................................................. M. dubia Flower buds 1–1.5 mm wide; leaves abruptly long-acuminate; mature fruits (when dry) < 1 cm diameter ....................................... M. floribunda

Myrciaria cordata O. Berg, Linnaea 27: 337. 1856. Probably a shrub or tree. Along rivers, ca. 600 m; Bolívar (near Roraima-tepui). Brazil (Roraima, 600 m). Known only from two collections, Schomburgk 608 (F, G, K, MICH, W) from Venezuela and Ule 8437 (K) from Brazil. Neither collection gives information as to the habit or habitat. It is expected to occur along rivers, as do most members of the genus.

Myrciaria dubia (H.B.K.) McVaugh, Fieldiana, Bot. 29(8): 501. 1963. —Psidium dubium H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 152. 1823. —Agracia ácido, Limoncillo. Myrciaria paraensis O. Berg in Mart., Fl. Bras. 14(1): 364. 1857. Myrciaria caurensis Steyerm., Fieldiana, Bot. 28: 1020. 1957. Shrub or small tree 1–6 m tall; bark exfoliating, mottled; leaves slightly bicolored;

82

M YRTACEAE

flowers fragrant; fruits red to dark purple at maturity. Frequent to locally dominant on sandy or rocky, seasonally flooded river banks, islands, and forests, 50–200(–500) m; Bolívar (lower Río Caura, Río Ichún, Río Orinoco, Río Paragua, Río Parhueña), Amazonas (basins of Río Casiquiare, Río Negro, Río Orinoco, and Río Ventuari). Anzoátegui, Apure; Amazonian Colombia, Guyana, Ecuador, Amazonian Peru, Brazil (Amazonas, Mato Grosso, Pará, Roraima, Rondônia), Bolivia. ◆Fig. 88. The acidic fruits of Myrciaria dubia are edible and are eaten raw or with salt and condiments, or made into juice or ice cream. They are high in ascorbic acid and widely used in the Amazon Basin where they are known by the name “camu-camu.” Myrciaria floribunda (West ex Willd.) O. Berg, Linnaea 27: 330. 1856. —Eugenia floribunda West ex Willd., Sp. Pl. 2: 960.

1800. —Guayabillo, Guayabillo blanco, Guayabo montañero, Turek (Pemón). Eugenia protracta Steud., Flora 26: 762. 1843. —Myrciaria protracta (Steud.) O. Berg, Linnaea 27: 330. 1856. Myrciaria uliginosa O. Berg, Linnaea 27: 329. 1856. Myrciaria verticillata O. Berg, Linnaea 27: 332. 1856. Shrub 2–3 m tall or tree 4–10(–18) m; young branches and petioles puberulous; leaves typically abruptly long-acuminate to caudate at apex; fruit orange to red when ripe. Forested or rocky river banks, moist forests, around granitic outcrops, 50–500(–900) m; Delta Amacuro (Río Grande), Bolívar (widespread), Amazonas (widespread). Apure, Falcón, Nueva Esparta, Táchira, Zulia; southern Mexico to Panama, West Indies, Colombia, Trinidad, Tobago, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 89.

Fig. 88. Myrciaria dubia

Myrciaria 83

Fig. 89. Myrciaria floribunda

Fig. 90. Myrciaria vismeifolia

84

M YRTACEAE

The common name, “turek” (horsefly), is given to this plant in Pemón because of the Indian belief that the flies are born from the plant. A gall with two concave halves, of unknown cause, commonly develops on the branches throughout its range. Myrciaria puberulenta B. Holst, Selbyana 23: 156. 2002. Tree to 15 m tall; leaves oblong, 15 cm long, membranous; inflorescence puberulent; fruit 2.5–3.5 cm diameter, green with winered tints when mature. Moist, tall evergreen forests, 100–800 m; Amazonas (Río Cataniapo, Río Cuao). Endemic. Myrciaria puberulenta is apparently rare since only three collections have been made from the heavily collected and easily accessible area just south of Puerto Ayacucho. It is perhaps related to M. vexator McVaugh, which is known from northern Venezuela and is cultivated in Costa Rica and Panama.

Myrciaria tenella (DC.) O. Berg in Mart., Fl. Bras. 14(1): 368. 1857. —Eugenia tenella DC., Prodr. 3: 272. 1828. Compact shrub or spindly tree 0.5–4 m tall; flowers white or pinkish; fruit purpleblack at maturity. River banks, 300–500 m; Bolívar (Caño Cucurital on Río Ventuari, Río Carrao basin). Brazil (Maranhão and Pará, south to Río Grande do Sul), Paraguay, Argentina, Uruguay. Myrciaria vismeifolia (Benth.) O. Berg, Linnaea 27: 336. 1856. —Eugenia vismeaefolia Benth., J. Bot. (Hooker) 2: 320. 1840. Small tree; twigs and petioles puberulent; leaf blades oblong, coriaceous, shallowly cordate at base; flowers borne on branches or trunk; fruits ca. 1 cm diameter (when dry), purple, acidic. Edge of rivers, ca. 100 m; Bolívar (Río Cuyuní, Río Venamo). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Acre, Amazonas). ◆Fig. 90.

11. MYRTEOLA O. Berg, Linnaea 27: 393. 1856. [Subtribe Myrtinae]. by Leslie Landrum Subshrubs or shrubs to ca. 4 m tall; trichomes simple, unicellular. Leaves opposite, decussate, often markedly 4-ranked, to ca. 11 mm long, coriaceous. Flower borne on a solitary uniflorous peduncle in the axil of a leaf, 4-or 5-merous, sometimes both types on a single plant; bracteoles foliaceous, linear to elliptic, persistent. Stamens 9–75; filament mainly longer than the anther during development

Fig. 91. Myrteola nummularia

Plinia 85

and at maturity. Ovary 2- or 3-locular, the septum between locules sometimes incomplete, the ovules 2–14, biseriate. Fruit a berry 5–8 mm diameter. Seeds 1.2–3 mm long, hard, shiny, smooth, not strongly angular, the seed coat ca. 0.1 mm thick, the inner cavity of the seed C-shaped; embryo C-shaped, the cotyledons nearly as long as the hypocotyl, not folded back against the hypocotyl. Western South America from Colombia and Venezuela to Chile; 3 species, 1 in Venezuela. Myrteola nummularia (Poir.) O. Berg, Linnaea 27: 396. 1856. —Myrtus nummularia Poir. in Lam., Encycl. 4: 407. 1797 [1798]. Creeping subshrub or small shrub to ca. 1 m tall; leaves < 1 cm long; inflorescences 1flowered. Tepui summits, 2500–2700 m;

Amazonas (Cerro Marahuaka). Apure, Mérida, Táchira; Andean Colombia, Ecuador, and Peru, Brazil (Amazonas: Serra da Neblina), Andean Bolivia, to sea level in southern Chile and Argentina; and the Falkland Islands (Islas Malvinas). ◆ Fig. 91.

12. PLINIA L., Sp. Pl. 516. 1753. [Subtribe Eugeniinae]. by Bruce K. Holst Shrubs or trees, often cauliflorous, often appressed-hirsutulous or sericeous. Leaf blades usually with convex midvein on upper surface. Inflorescences of small, nearly sessile, multibracteate shoots, or well developed and paniculate. Flowers 4merous; buds closed or nearly so; hypanthium prolonged beyond the summit of the ovary; calyx lobes free to nearly fully connate, small, tearing irregularly and longitudinally at anthesis, persistent. Stamens borne on hypanthium; ovary 2-locular; ovules 2(–4) per locule. Fruit round or tapering at both ends, occasionally with longitudinal ridges, often orange at maturity. Seeds 1 or 2; embryo eugenioid, with 2, free, plano-convex cotyledons. West Indies, tropical South America east of the Andes and south to Paraguay; ca. 20 species, 6 or 7 in Venezuela, 5 of these in the flora area. Plinia is a seemingly artificial genus containing two disparate elements and is much in need of revision. The element that includes the type species has dense, bracteate, nearly sessile shoots that are usually borne on older branches or trunks, and the other has paniculate inflorescences borne in leaf axils on younger branches. The fruits of many species are reported to be edible. Key to the Species of Plinia 1.

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

Inflorescence racemose or paniculate, the main axis > 1 cm long, evident; plants glabrous or with the inflorescence sparsely puberulous .................................................................................................... P. rivularis Inflorescence axis reduced and hidden by the bracts; plants pubescent, at least conspicuously so on stems, petioles, and flowers ........................ 2 Secondary veins impressed on upper surface of leaf blades .............. P. sp. A Secondary veins plane or slightly elevated on upper surface of leaf blades, not impressed ......................................................................................... 3 Larger leaf blades 5–9 cm long ...................................................... P. pinnata Leaf blades > (10–)13 cm long ................................................................... 4

86

4(3). 4.

M YRTACEAE

Fruits and lower surface of leaf blades pubescent ....................P. involucrata Fruits and leaf blades glabrous to sparsely sericeous ........................ P. sp. B

Plinia involucrata (O. Berg) McVaugh, Mem. New York Bot. Gard. 18(2): 228. 1969. —Myrciaria involucrata O. Berg in Mart., Fl. Bras. 14(1): 375. 1857. Shrub or tree 2–5 m tall; vegetative portions pubescent with soft yellowish green trichomes; leaf margins revolute, apex sharply acuminate; fruit pubescent. Margins of granitic outcrops, ca. 100 m; Bolívar (Río Orinoco near Río Suapure), Amazonas (Puerto Ayacucho). Apure; Brazil (Amazonas). ◆Fig. 92. Plinia pinnata L., Sp. Pl. 516. 1753. —Tu‘ kinke me‘n (Panare). Shrub or tree 3–10 m tall, cauliflorous; leaves sparsely sericeous, coriaceous-semisucculent, the secondary venation inconspicuous; flowers cream-colored, fruits orange or golden yellow at maturity, smooth. Lowland riparian forests, moist evergreen forests, premontane semideciduous forests, 100–400 m; Bolívar (Represa Raúl Leoni, Río Botanamo, Río Caroní, Río Cuyuní, Río

Fig. 92. Plinia involucrata

Paragua basin, Sierra de la Cerbatana), Amazonas (Río Cataniapo, Río Orinoco, Río Ventuari basin). Lesser Antilles, Trinidad, Guyana, Suriname. The fruit of Plinia pinnata is edible and acidulous. Populations from the Lesser Antilles have ridged fruits, while those from South America are smooth; it is possible that more than one taxon is included here. Plinia rivularis (Cambess.) Rotman, Boletín de la Sociedad Argentina de Botánica 24(1–2): 196. 1985. —Eugenia rivularis Cambess. in A. St.-Hil., Fl. Bras. Merid. 2: 337. 1829 [1832]. Myrcia granulata R.O. Williams, Fl. Trinidad 1: 340. 1934. Shrub or tree 3–18 m tall; leaf size, shape and texture variable, typically ovate to elliptic with abruptly acuminate apex, secondary veins numerous, fine; inflorescence racemose

Pseudanamomis 87

or paniculate. Seasonally flooded, evergreen forests along rivers, 100–200 m; Amazonas (Caño Iguapo, La Esmeralda, near Puerto Ayacucho). Barinas, Distrito Federal, Yaracuy; Trinidad, Suriname(?), Brazil, Paraguay, Argentina, Uruguay. Plinia rivularis belongs to a group of species, largely Brazilian, that have well-developed inflorescences which do not match the typical elements of Plinia with reduced bracteate shoots. In flower, members of the Plinia rivularis group resemble tetramerous species of Myrcia (e.g., Myrcia grandis McVaugh), but the embryo is distinctly eugenioid. Plinia sp. A. —Iramatemoko (Yanomami). Tree 15 m tall; leaves to 27 cm long, the midvein broad and convex on upper surface, the secondary veins impressed; fruits dark yellow. Lowland floodplain forests, 100–200 m; Amazonas (Río Jénita in Río Ocamo basin).

The only known collection of this taxon from Venezuela (Angel Fernandez 7301, SEL), appears conspecific with collections from the Peruvian Amazon, though it is not clear which, if any, name is best applied. The Venezuelan specimen is in fruit. Plinia sp. B. —Raek-i (Piaroa). Tree 4–5 m tall; leaves abruptly acuminate, secondary veins inconspicuous on upper surface; fruit yellow to orange. Secondary forest areas, old gardens, 700–800 m; Amazonas (Río Cuao basin). As in Plinia sp. A, this appears related to other Plinia collections known from the Amazon basin, though flowers are not known from the Venezuelan collections (e.g., Zent 785-35b, SEL; Zent 186-13, SEL). Hence, it is difficult to provide a name for this taxon. The fruits are reported as edible for humans, as well as various other forest dwellers.

13. PSEUDANAMOMIS Kausel, Ark. Bot. ser. 2, 3(15): 512. 1955. [Subtribe Eugeniinae]. by Francesca Grifo Shrubs or trees to 15 m tall; bark exfoliating, sometimes reddish in color. Leaves cartilaginous or chartaceous; blade orbicular, obovate, obcordate, or ovate, the apex retuse or emarginate; midvein impressed on upper surface, marginal veins indistinct. Inflorescence a dichasium, often much branched and resembling an umbel, 3–12-flowered, central flowers sessile, lateral flowers usually pedicellate; peduncle to 2 cm long; bracts and bracteoles linear or lanceolate, scarious, early deciduous. Flowers 4- or 5-merous; hypanthium not prolonged beyond ovary summit, tomentose; calyx lobes distinct, imbricate, in equal or unequal pairs, deciduous, variously pubescent; petals imbricate, concave, reflexed at anthesis, white or greenish white; disk quadrangular or round. Stamens many; anthers longitudinally dehiscent. Ovary bilocular; ovules 5 or 6 per locule. Fruit spherical, 2–5 cm diameter, smooth, yellow, orange, or red at maturity, with abundant sweet flesh. Seeds 1–3, reniform, to 4 cm long; testa light brown, papery and peeling easily; cotyledons 2, plano-convex, united laterally for less than 1/5 their length, large, fleshy. Hispaniola, Puerto Rico, Venezuela, Colombia, Trinidad, Guyana; 1 species. Pseudanamomis umbellulifera (H.B.K.) Kausel, Ark. Bot. ser. 2, 3(15): 512. 1955. —Myrtus umbellulifera H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 135. 1823. —Myrcianthes umbellulifera (H.B.K.) Alain, Brittonia 20: 159. 1968. —Paují. Myrtus erythroxyloides H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 149. 1823. Much-branched shrub or tree to 12 m tall; leaves light green; fruits usually orange at

maturity. Lowland to premontane semideciduous forests, 50–500 m; northern Bolívar. Anzoátegui, Aragua, Dependencias Federales, Distrito Federal, Falcón, Guárico, Lara, Miranda, Nueva Esparta, Portuguesa, Sucre, Zulia; other distribution as in genus. ◆Fig. 93. It is not clear what influence cultivation has had on the range of this species, which is grown for its sweet, edible fruits.

88

M YRTACEAE

Fig. 93. Pseudanamomis umbellulifera

14. PSIDIUM L., Sp. Pl. 470. 1753. [Subtribe Myrtinae]. by Leslie Landrum Shrubs or trees; trichomes unicellular, simple. Inflorescences uniflorous or a 3(rarely more)-flowered dichasium. Flowers 5-merous, whitish; calyx lobes free or fused beyond the summit of the ovary in a hypanthial tube, sometimes forming a calyptra. Stamens ca. 100 to over 500. Ovary 2–5-locular; ovules few to numerous, biseriate or multiseriate; placenta bilamelate and often protruding in a peltate structure, the ovules then attached to the edge of the lamelae. Fruit a greenish, yellowish, or reddish berry. Seeds few to numerous, the seed coat bony, dense, ca. 9–30 cells thick at narrowest point, covered with a thin layer of pulpy tissue when wet (or a glaze or crusty tissue when dry). Mexico and the Caribbean to Argentina; ca. 70 species, ca. 20 in Venezuela, 10 of these in the flora area. Species of Psidium are often cultivated for their edible fruits, particularly Psidium guajava (guava) and Psidium cattleianum (strawberry guava). The soft fruits, which have a limited shelf life, are processed into ice creams, beverages, and preserves.

Psidium 89

Key to the Species of Psidium 1.

Twigs with 4 wings or 4 sharp angles, ± quadrangular in cross section ................................................................................................................ 2 1. Twigs without 4 wings, ± terete in cross section ...................................... 4 2(1). Leaves with ca. 12–20 pairs of lateral veins; twigs and leaves covered with grayish, appressed trichomes ............................................ P. guajava 2. Leaves normally with < 12 pairs of lateral veins; twigs and leaves not covered with grayish, appressed trichomes ............................................... 3 3(2). Small shrubs of savannas; leaves usually obovate; calyx generally not closed in bud; seed rounded, not angled nor C-shaped ........... P. cinereum 3. Large trees or shrubs, usually of riparian habitats; leaves elliptic; calyx closed or nearly so in bud; seeds generally angular, C-shaped ............................................................................................ P. acutangulum 4(1). Plants in flowering condition .................................................................... 5 4. Plants with mature fruits ........................................................................ 14 5(4). Calyx open, not enclosing the globe of the closed corolla ......................... 6 5. Calyx closed, enclosing the corolla, or with a small open pore at the apex only ......................................................................................................... 8 6(5). Tree of riparian habitats; leaves lanceolate, 5–15(–19) cm long; petiole 4–14 mm long ....................................................................... P. densicomum 6. Shrubs (rarely small trees) of savannas; leaves variously shaped, usually < 7 cm long; petiole 0–2 mm long .......................................................... 7 7(6). Plants yellowish or grayish tomentose on young growth ..... P. laruotteanum 7. Plants glabrous to thinly pubescent ............................................... P. salutare 8(5). Twigs glabrous ........................................................................................... 9 8. Twigs moderately to densely pubescent .................................................. 11 9(8). Leaves usually 2.4–7 cm long; anthers 0.3–0.5 mm long; style ca. 5 mm long; calyx opening by means of a calyptra, or tearing irregularly, or tears producing 5 lobes ........................................................ P. sartorianum 9. Leaves 5–12 cm long; anthers ca. 1 mm long; style 8–20 mm long; calyx tearing irregularly at anthesis ............................................................ 10 10(9). Petiole usually < 4 mm long; leaves elliptic, ovate, or lanceolate, the lateral veins prominent or faint; style 11–20 mm long; petals ca. 1.5 mm long; riparian habitats ....................................................... P. acutangulum 10. Petiole usually > 4 mm long; leaves elliptic, elliptic-oblong, or obovate, the lateral veins prominent; style 8–10 mm long; petals 7–11 mm long; various habitats ....................................................................... P. guineense 11(8). Leaves elliptic, oblanceolate, or oval, usually widest above the middle, the margin usually crenulate; calyx glabrous within ........... P. maribense 11. Leaves variously shaped, widest at or below the middle, the margin usually not crenulate; calyx pubescent within (except sometimes glabrous in P. sartorianum) ................................................................................ 12 12(11). Midvein ± flat on upper surface of leaf, not impressed except slightly near base of leaf; lateral veins not prominent; calyx opening by means of a calyptra, or tearing irregularly, or tears producing 5 lobes ..... P. sartorianum 12. Midvein impressed on upper surface of leaf; lateral veins prominent; calyx tearing irregularly at anthesis ...................................................... 13

90

M YRTACEAE

13(12). Blades submembranous to subcoriaceous; petiole 1–3 mm long; base often subcordate ...................................................................... P. striatulum 13. Blades stiffly coriaceous; petiole 4–12 mm long; base not normally subcordate ................................................................................ P. guineense 14(4). Seeds angular, usually C-shaped; plants of riparian habitats ............... 15 14. Seeds smooth, rounded; plants of various habitats, often upland ......... 19 15(14). Twigs glabrous ......................................................................................... 16 15. Twigs pubescent ....................................................................................... 17 16(15). Petiole (0–)1–6 mm long; peduncle rarely triflorous; calyx tearing irregularly, 5 lobes usually not evident ...................................... P. acutangulum 16. Petiole 4–14 mm long; peduncle often triflorous; calyx torn between the lobes, 5 lobes usually evident .............................................. P. densicomum 17(15). Leaves usually widest above the middle, the margin usually crenulate; remnants of calyx glabrous within ........................................ P. maribense 17. Leaves usually widest at the middle or below, the margin entire or obscurely sinuate-crenulate; remnants of the calyx glabrous or pubescent within ................................................................................................... 18 18(17). Petiole 1–3 mm long; leaf usually 2.2–7 cm long, elliptic to oblonglanceolate ................................................................................ P. striatulum 18. Petiole 4–14 mm long; leaf usually 5–15 cm long, lanceolate to ovate .............................................................................................. P. densicomum 19(14). Seeds ca. 3 mm long, ca. 100 per fruit; leaves 4–11.5 cm long, the lateral veins prominent; petiole 4–12 mm long .................................. P. guineense 19. Seeds 4–6 mm long, 1–13(–20) per fruit; leaves mainly 2–7 cm long, the lateral veins not prominent; petiole 0–3 mm long .............................. 20 20(19). Trees of forests; calyx closed, not persisting on the fruit, or if persisting, then as a dish-like calyptra, or as 5 more or less irregular lobes, the sinuses between the lobes produced by tearing .................. P. sartorianum 20. Shrubs or small trees of savannas; calyx open, 5-lobed, the lobes usually evident on the fruit, little tearing occurring between the lobes ........ 21 21(20). Leaves and twigs densely yellowish or grayish tomentose ..... P. laruotteanum 21. Leaves and twigs glabrous to only moderately pubescent ............ P. salutare Psidium acutangulum DC., Prodr. 3: 233. 1828. —Guayabo de caño danero. Psidium persoonii McVaugh, Mem. New York Bot. Gard. 18: 255. 1969. Shrub or tree 6–10 m tall; flowers long-pedunculate. Along rivers and in gallery forests and savannas, 50–500 m; Delta Amacuro (Caño Jota-Sabuca near Caño Mariusa), northern and central Bolívar, Amazonas (widespread). Anzoátegui, Apure, Barinas; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Amapá; Mato Grosso, Pará, Rondonia), Bolivia. ◆Fig. 95. Psidium acutangulum is a widespread and variable species. In our area a segregate species P. persoonii has been recognized, but here a more conservative approach has been taken because many intermediates exist. Ad-

ditional studies of the complex would be useful. The fruits are edible. In their ideal states these entities can be distinguished by the following sets of characters. In Psidium acutangulum, the twigs are strongly winged, the lateral veins are usually impressed, the calyx is closed and apiculate, and the young twigs and petioles are sparsely hairy. In P. persoonii, the twigs are scarcely winged, the lateral veins are not impressed, the calyx is closed or has a pore at the apex, and the young twigs and petioles are glabrous. Psidium cinereum DC., Prodr. 3: 234. 1828. Psidium australe Cambess. in A. St.-Hil. et al., Fl. Bras. Merid. 2: 283. 1829 [1832].

Psidium 91

Psidium submetrale McVaugh, Mem. New York Bot. Gard. 18(2): 261. 1969. Shrub or subshrub to ca. 0.7 m tall, new stems sprouting from a woody base; leaf blades obovate, coriaceous, sub-tripliveined. Savannas subject to fire, 50–500 m; northern Bolívar. Guyana, Brazil (Goias, Maranhão, Minas Gerais, Paraná, Roraima), Bolivia, Paraguay, Argentina. Psidium cinereum is a widespread and variable species of the savannas, ranging from Venezuela to southern Brazil. Psidium densicomum DC., Prodr. 3: 235. 1828. —Guayabo. Psidium ovatifolium O. Berg in Mart., Fl. Bras. 14(1): 385. 1857. Shrub or tree to 10 m tall; leaf blades typically ovate and long and evenly tapered to a sharp point. Lowland to premontane gallery forests, along rivers, around lakes, 50–400 m; Bolívar (Hato La Vergareña, Río Aro, Río Paragua), Amazonas (ca. 20 km south of confluence of Río Negro and Brazo Casiquiare, along Río Orinoco at Perro de Agua). Apure, Anzoátegui; Colombia, Guyana, Brazil (Amazonas, Pará), Bolivia. ◆Fig. 98. Typical Psidium densicomum from the western Amazon basin is glabrous. The name P. ovatifolium has been used for populations of P. densicomum in eastern South America that have softly hirsute young growth. With age, most (perhaps all) indument is lost from the young growth, so that the distinction between P. densicomum and P. ovatifolium becomes difficult, if not impossible, to make. Thus, the two taxa are united here. Psidium guajava L., Sp. Pl. 470. 1753. —Guayabo. Shrub or tree to 5 m tall; leaves coriaceous, the secondary veins impressed on the upper surface; fruits yellow-green at maturity with pinkish pulp. Cultivated and in disturbed areas near houses and along roads, also growing at the edges of forests and along streams, 50–1000 m; Bolívar (widespread). Aragua, Falcón, Nueva Esparta, Sucre, Zulia; common from Mexico and the Caribbean to Bolivia, Paraguay, and northeastern Argentina, naturalized on some Pacific islands. Guava is one of the most recognized and important tropical and subtropical fruits. It is high in Vitamin C and is eaten raw or made into beverages, preserves, and ice

creams. Psidium guajava and P. guineense commonly hybridize; for discussion see P. guineense. Guava is a serious noxious weed in Fiji and Hawai’i. Psidium guineense Sw., Prodr. 77. 1788. —Guayabo, Guayabo sabanero. Psidium guyanense Pers., Syn. Pl. 2: 27. 1806 [1807]. Shrub or tree to 4 m tall, usually abundantly pilose with light brown soft trichomes; leaves coriaceous, typically obovate; fruits yellow-green at maturity with cream-colored pulp. Disturbed areas, savannas, gallery forests, 50–800 m; widespread in Bolívar and Amazonas. Apure, Aragua, Distrito Federal, Falcón, Guárico, Mérida, Táchira, Zulia; common from Mexico and the Caribbean to Bolivia, Paraguay, and northeastern Argentina. ◆Fig. 97. Psidium guineense, perhaps the most often collected species in the genus, commonly hybridizes with P. guajava. Intermediates can usually be recognized by having a combination of quadrangular twigs (characteristic of Psidium guajava) and erect, reddish brown trichomes on the lower surface of the leaf (characteristic of Psidium guineense). Psidium laruotteanum Cambess. in A. St.Hil., Fl. Bras. Merid. 2: 282. 1829 [1832]. —Kanomi-dek (Pemón-Taurepan). Psidium glaucescens O. Berg in Mart., Fl. Bras. 14(1): 600. 1859. Psidium quinquedentatum Amshoff, Recueil Trav. Bot. Néerl. 39: 164. 1942. Shrub to 2 m tall, abundantly yellowish to grayish pilose; fruits yellow-orange at maturity. Upland savannas, 800–1000 m; Bolívar (Gran Sabana). Zulia; Costa Rica, Guyana, Brazil (Goias, Minas Gerais, Paraná, São Paulo). The pleasantly flavored fruits are edible and used to treat stomach upset and colic. The bark is boiled to make a tea. Psidium maribense DC., Prodr. 3: 233. 1828. —Guayabito de agua, Guayabo blanco, Guayabo tortugero. Shrub to ca. 4 m tall; leaves crenate. Apparently a common element in shrubby riparian vegetation in sandy soils, 50–100 m; Bolívar (Río Orinoco and tributaries). Anzoátegui, Apure, Guárico; Colombia (Vichada). ◆Fig. 99.

92

M YRTACEAE

The crenate-margined leaves of this species are unusual in the family. The fruits are said to be eaten by turtles. Psidium salutare (H.B.K.) O. Berg, Linnaea 27: 356. 1856. —Myrtus salutaris H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 132. 1823. —Guayabita, Guayabo rajado. Shrub to 1 m tall throughout most of range, the fire-resistant woody base apparently adapted to frequent fires in savannas; becoming a tree to 10 m tall near Puerto Ayacucho; leaves coriaceous. Savannas and gallery forests, 50–400 m; northern Bolívar, western Amazonas. Apure, Barinas, Cojedes, Guárico, Lara, Mérida, Monagas, Portuguesa, Sucre, Zulia; Central America, Cuba, Dominican Republic, Colombia, Guyana, Brazil (Goias, Mato Grosso, Pará, Paraná, São Paulo). ◆Fig. 94. The arboreal forms near Puerto Ayacucho are unusual for this species and merit further study. The fruits are edible and sometimes gathered. Psidium sartorianum (O. Berg) Nied. in Engl. & Prantl, Nat. Pflanzenfam. 3(7): 69. 1893. —Mitranthes sartoriana O. Berg, Linnaea 29: 248. 1858. Shrub or tree to 25 m tall; fruits 1–2 cm diameter, yellowish green at maturity, the flesh creamy yellow. Upland gallery forests, lower montane forests, 500–600 m; Bolívar (Altiplanicie de Nuria). Aragua, Falcón, Lara, Miranda, Monagas, Zulia; Mexico to Panama, Colombia, Ecuador, Brazil (Espirito Santo, Pará, Rio de Janeiro, Rondônia), Bolivia. ◆Fig. 96. The smallish fruits have a pleasant taste and are used to make preserves. Psidium sartorianum is perhaps not distinct from P. galapageium of the Galapagos Islands, in which case the latter name would have priority. Psidium striatulum DC., Prodr. 3: 233. 1828. Psidium aquaticum Benth., J. Bot. (Hooker) 2: 318. 1840.

Fig. 94. Psidium salutare

Psidium parviflorum Benth., J. Bot. (Hooker) 2: 318. 1840. Shrub or small tree to 6 m tall. Riparian habitats subject to flooding, ca. 100 m; Amazonas (Río Orinoco near Isla Ratón and Isla Hormiga). Guyana, Suriname, northern Brazil (Amazonas, Roraima).

Psidium 93

Fig. 95. Psidium acutangulum

Fig. 97. Psidium guineense

Fig. 96. Psidium sartorianum

94

M YRTACEAE

Fig. 98. Psidium densicomum

Fig. 99. Psidium maribense

15. SIPHONEUGENA O. Berg, Linnaea 27. 344. 1856. [Subtribe Eugeniinae]. by Bruce K. Holst Shrubs or small trees; twigs laterally compressed when young. Leaf midvein convex or plane on upper surface, lateral veins numerous, parallel, marginal vein mostly straight, not arching between the lateral veins. Inflorescence axillary, occasionally subterminal, usually occurring at leafless nodes, usually shortly decussateracemose, or rarely the flowers few and appearing fasciculate; bracts and bracteoles nonfoliaceous, usually persisting into fruit. Flowers 4-merous; hypanthium funnelshaped, much prolonged and constricted just above the summit of the ovary, the

Siphoneugena 95

apex strongly reflexed at anthesis, entire hypanthium cleanly circumscissile above the summit of the ovary and falling along with perianth and androecium after anthesis. Calyx open to completely closed in bud, lobes when present small, longitudinal tearing between them, pubescent within; petals imbricate. Stamens borne near summit of hypanthium, incurved in bud; anthers oblong. Ovary bilocular, ovules (2)3–5(–7) per locule; placentation axile or nearly basal. Fruit round, crowned by the circular hypanthium scar. Seeds 1(–4); testa membranaceous; embryo eugenioid, of 2 plano-convex cotyledons; hypocotyl minute. Panama, Puerto Rico, Lesser Antilles, Venezuela, Suriname, Peru, Brazil, Bolivia, Argentina; 8 species, 1 in Venezuela. Siphoneugena dussii (Krug & Urb.) Proença, Edinb. J. Bot. 47: 251. 1990. —Marlierea dussii Krug & Urb., Bot. Jahrb. Syst. 19: 590. 1895. Plinia fruticosa Steyerm., Fieldiana, Bot. 28: 1023. 1957. Siphoneugena densiflora var. tepuiensis Steyerm., Fieldiana, Bot. 28: 1023. 1957. Siphoneugena dussii var. salicifolia McVaugh, Mem. New York Bot. Gard. 18(2): 231. 1969. —Siphoneugena densiflora var. salicifolia (McVaugh) Proença, Edingb. J. Bot. 47: 254. 1990. Siphoneugena densiflora auct. non O. Berg 1857: McVaugh, Mem. New York Bot. Gard. 18(2): 230. 1969. Shrub or small tree 1.5–5(–12) m tall; leaves varying in shape from narrowly lanceolate (when plants growing along rivers) to ovate or broadly elliptic, lustrous, midvein prominently convex; inflorescence appearing glomerate; calyx lobes reddish; fruit with dull taste, purple-black at maturity. Along rocky or forested portions of rivers, moist or

dry exposed sandstone slopes and ledges, forested slopes, tepui summits, (600–)1000– 2200 m; Bolívar (Altiplanicie de Nuria, Cerro Venamo, Gran Sabana, Macizo del Chimantá, Sororopán-tepui), Amazonas (widespread). Coastal Cordillera (Anzoátegui, Aragua, Distrito Federal, Miranda, Sucre, Táchira, Trujillo); Costa Rica, Panama, Puerto Rico, Lesser Antilles, Suriname (Tafelberg), Peru, Brazil (Amazonas, Mato Grosso, Pará). ◆Fig. 100.

Fig. 100. Siphoneugena dussii

96

M YRTACEAE

16. SYZYGIUM Gaertn., Fruct. Sem. Pl. 1: 166, t. 33. 1788. [Subtribe Eugeniinae]. Caryophyllus L., Gen. Pl. ed. 5, 515. 1754. Jambosa DC., Prodr. 3: 286. 1828. by Bruce K. Holst Trees or shrubs, rarely decumbent; twigs usually glabrous. Leaves petiolate, rarely almost sessile, marginal vein usually visible. Inflorescence terminal, axillary, ramal or cauline, usually paniculate, occasionally umbellate or racemose, sometimes few-flowered and cymose; bracts usually absent, rarely present at anthesis. Flowers 4- or 5-merous, rarely 3- or 6-merous, buds clavate to obovoid; calyx lobes usually present, persistent, or absent on calyptrate species; petals free or coherent, occasionally ± calyptrate. Anthers variable, anther sacs usually parallel. Ovary 2(3)-locular; ovules usually radiating from a centrally located axile placenta, occasionally superimposed along a vertical placenta, rarely pendulous from a placenta at the apex of each locule; style short or long. Fruits ± baccate, pericarp usually fleshy. Seeds 1, rarely 2–5; cotyledons distinct from one another, radicle short, hidden between the cotyledons. Paleotropics, introduced to all tropical regions; ca. 500 species, ca. 5 species cultivated in Venezuela, 3 of these cultivated and escaping in the flora area. Collections of Syzygium in the flora area are infrequent since they are widely recognized as being exotic species and frequently passed up by collectors. All of the species in the flora area are cultivated for ornament and their edible fruits. Key to the Species of Syzygium 1.

1.

2(1).

2.

Flowers rose or reddish purple; fruits obpyriform; marginal veins 2, the slighter marginal arising from the base, the principal one arising from the 3rd or 4th lateral vein and arching between them ....... S. malaccense Flowers white; fruits globose or broadly oblong to ellipsoid; marginal vein 1, arising from the base of leaf and running ± straight along margin ................................................................................................................ 2 Flowers small, 3–5 mm across at anthesis; calyx truncate; petals (frequently mistaken for the calyx) calyptrate; inflorescence a branched cyme, many-flowered; fruit oblong or ellipsoid, dark purple to black when mature; leaves elliptic, oblong, to obovate ........................ S. cumini Flowers large, 15 mm wide, or usually much wider at anthesis; petals not calyptrate, free; calyx with 4 evident lobes; inflorescence a short corymb, few-flowered; fruit globose or ovoid, white, pinkish, or yellowish when mature; leaves lanceolate to elliptic-lanceolate .......... S. jambos

Syzygium cumini (L.) Skeels, U.S.D.A. Bur. Pl. Industr. Bull. 248: 25. 1912. —Myrtus cumini L., Sp. Pl. 471. 1753. —Eugenia cumini (L.) Druce, Bot. Exch. Club Soc. Brit. Isles 3: 418. 1914. —Yamú. Tree to 20 m tall, glabrous; twigs white; petioles 1.5–2 cm long; leaf blade with lateral veins numerous and close together; flow-

ers white; filaments < 6 mm long; fruit crowned by persistent hypanthium. Cultivated, possibly escaping, near sea level to 200 m; Delta Amacuro, Bolívar (Río Cuyuní), Amazonas (Gavilán). Widespread elsewhere in Venezuela; native to the Indo-Malaysian region, cultivated pantropically. ◆Fig. 102. Syzygium cumini is widely cultivated in the tropics for ornament and its edible fruit.

Syzygium 97

Fig. 101. Syzygium jambos Fig. 102. Syzygium cumini

Fig. 103. Syzygium malaccense

98

M YRTACEAE

Syzygium jambos (L.) Alston in Trimen, Handb. Fl. Ceylon (Suppl.) 6: 115. 1931. —Eugenia jambos L., Sp. Pl. 470. 1753. —Manzana rosa, Pomarosa. Jambosa vulgaris DC., Prodr. 3: 286. 1828. Shrub or tree 5–20 m tall, glabrous; leaves long-acuminate, coriaceous, petiole to 1 cm long, lateral veins ca. 10–15 pairs; flowers and filaments white, to 4 cm long; inflorescence usually terminal; flowers fragrant; fruit rosescented, ca. 3 cm diameter, white, pinkish, or yellowish, the pulp white. Escaping from cultivation, 50–500 m; Bolívar (scattered). Widespread elsewhere in Venezuela; native to the Indo-Malaysian region, cultivated and naturalizing pantropically. ◆Fig. 101.

Syzygium malaccense (L.) Merr. & Perry, J. Arnold Arb. 19: 215. 1938. —Eugenia malaccensis L., Sp. Pl. 470. 1753. —Pomagás. Tree to 30 m tall, glabrous; leaves large, 20–30 cm long, broadest at middle or above, lateral veins 10–15 pairs; inflorescence axis woody, from stout branches within the canopy; flowers rose or reddish purple; filaments pink or rose, 2–3 cm long; fruit obovoid, ca. 7 cm long, the pulp white. Escaping from cultivation, flooded forests, ca. 100 m; Delta Amacuro (Río Cuyubini), Amazonas (San Carlos de Río Negro). Widespread elsewhere in Venezuela; native to the Indo-Malaysian region, cultivated pantropically. ◆Fig. 103.

17. UGNI Turcz., Bull. Soc. Imp. Naturalistes Moscou 21(1): 579. 1848. [Subtribe Myrtinae]. by Leslie Landrum Shrubs to ca. 3 m tall; trichomes unicellular, simple or dibrachiate. Leaves opposite, persistent, coriaceous, mainly elliptic to lanceolate. Inflorescence a solitary, nodding flower, the peduncles not superimposed, borne in the axils of leaves or bracts. Flowers usually 5-merous, less often both 4- and 5-merous on a single plant; corolla campanulate, the stamens not exceeding the petals, the filaments short, the anthers elongate, ± sagittate, introrsely dehiscent; bracteoles persistent usually until fruit matures, linear to narrowly elliptic. Ovary usually 3-locular, the ovules biseriate, 4-seriate, or multiseriate. Fruit a berry. Seeds few to many, shiny light brown, small, ca. 1–2 mm long; embryo C-shaped, the cotyledons almost as long as the hypocotyl, scarcely if at all folded back against themselves. High-elevation areas of Mexico (north to Oaxaca), Central America, Colombia, Venezuela, Ecuador, Peru, Chile (where most diverse), Argentina; 4 species, 1 in Venezuela. Ugni myricoides (H.B.K.) O. Berg, Linnaea 27: 391. 1856. —Myrtus myricoides H.B.K., Nov. Gen. Sp. 6 (quarto ed.): 131. 1823. Myrtus stenophylla Oliv., Trans. Linn. Soc. London, Bot. 2: 273. 1887. —Ugni stenophylla (Oliv.) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15: 507. 1941. —Myrtus myricoides var. stenophylla (Oliv.) Steyerm., Fieldiana, Bot. 28: 1022. 1957. —Ugni myricoides f. stenophylla (Oliv.) McVaugh, Mem. New York Bot. Gard. 18(2): 266. 1969. Myrtus roraimensis N.E. Br., Trans. Linn. Soc. London, Bot. 6: 26. 1901. —Ugni

roraimensis (N.E. Br.) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15: 507. 1941. —Myrtus myricoides var. roraimensis (N.E. Br.) Steyerm., Fieldiana, Bot. 28: 1022. 1957. —Ugni myricoides var. roraimensis (N.E. Br.) McVaugh, Mem. New York Bot. Gard. 18(2): 265. 1969. Ugni angustifolia Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15: 507. 1941. Ugni vaccinioides Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15: 506. 1941. Myrtus myricoides var. turumiquirensis Steyerm., Fieldiana, Bot. 28: 1022. 1957. Ugni myricoides f. bifaria McVaugh, Mem. New York Bot. Gard. 18(2): 267. 1969.

Ugni 99

Ugni myricoides f. grossa McVaugh, Mem. New York Bot. Gard. 18(2): 266. 1969. Ugni myricoides f. oligandra McVaugh, Mem. New York Bot. Gard. 18(2): 266. 1969. Shrub 1–2 m tall or less often a small tree to 5 m tall; leaves small, stiff-coriaceous, margins revolute, the secondary venation inconspicuous; inflorescences 1-flowered, the flowers nodding; fruits white(?) at maturity. High-elevation shrublands and open areas on tepuis, 1900–2800 m; widespread in Bolívar and Amazonas. Mérida, Sucre, Táchira; southern Mexico, Central America, Colombia, Guyana, Ecuador, Peru, Brazil (Amazonas: Serra da Neblina; Roraima: Cerro Roraima). ◆Fig. 104. This is a variable species in the Guayana Highland. Six of the varieties and forms recognized by McVaugh (Mem. New York Bot. Gard. 18(2): 263–267) are reported for our area. This variation is of interest and persons wishing to recognize intraspecific taxa may want to consult his work.

Fig. 104. Ugni myricoides

100

N AJADACEAE

NAJADACEAE by Robert R. Haynes and Lauritz B. Holm-Nielsen Annual or rarely perennial aquatic herbs, submersed in fresh or brackish waters, glabrous. Stems slender, much-branched, rooting at the lower nodes, sometimes armed with prickles on the internodes. Leaves subopposite or appearing whorled due to reduced internode length, sessile, each divided into blade and sheath; blade linear and flattened, 1-veined, sometimes dorsally armed with prickles on the midrib; margins usually serrulate, or serrate to dentate, with 5–100 teeth per side, the apex acute to acuminate, with 1–3 teeth per side, the teeth unicellular or else multicellular, then formed by layers of cells that decrease in cell number outward, terminated by a large, sharp-tipped cell. Leaf sheaths variously shaped, each enclosing a pair of tiny hyaline scales, the margins usually toothed with 1–15 teeth per side. Flowers unisexual (plants monoecious or dioecious), axillary, sessile or short-pedunculate, solitary or clustered, often subtended by an involucre; involucre clear, bronze, brown, light green, purple, or red-purple. Staminate flowers subtended by a membranous involucre, or the involucre rarely absent, each consisting of a single stamen; peduncle at first short, elongating at anthesis, pushing the anther through the involucre; anther sessile, 1- or 4-locular, dehiscing irregularly; pollen 3-celled, globular or ellipsoid, densely filled with starch, monocolpate, the wall with shallow reticulations, thin, not divided into exine and entine. Pistillate flowers sessile, 1locular, 1-ovuled; involucre absent or rarely present, ending in a short style with 2– 4 branches. Fruit 1-seeded, dehiscing by decay of the gynoecial wall, which is extremely delicate and closely envelopes the seed. Seeds fusiform to obovate, occasionally asymmetrical at apex or recurved, with a basal raphe; testa hard, brittle, 3- or several-cell-layered, pitted or smooth; areolae formed by outer 2 layers of testa, variously rectangularly shaped, irregularly arranged in 15–60 rows, the end walls often raised, giving the testa a papillose appearance; endosperm lacking; embryo elongate, without lateral enlargements of cotyledons. Nearly cosmopolitan; 1 genus and ca. 40 species, 1 species in the flora area. 1. NAJAS L., Sp. Pl. 1015. 1753; Gen. Pl. ed. 5. 445. 1754. The genus with the characteristics of the family and distribution of the family; ca. 40 species, 3 in Venezuela, 1 of these in the flora area. Seed characters in Najas are very important for distinguishing the species.

Fig. 105. Najas wrightiana

N Y C TA G I N A C E A E 101

Najas wrightiana A. Braun, Sitzungsber. Ges. Naturf. Freunde Berlin 1868: 17. 1868. Plant 5–45 cm long. Submersed in lakes

and ponds, 800–1000 m; Bolívar (Río Kukenán, Santa Elena). U.S.A. (Florida), Central America, Colombia, Ecuador, Peru, Brazil. ◆Fig. 105.

NYCTAGINACEAE by Julian A. Steyermark and Gerardo A. Aymard C. Annual or perennial herbs, shrubs, or trees, sometimes climbing, the stems frequently swollen at the nodes, sometimes armed with spines, dichotomously or trichotomously branched. Leaves simple, alternate, opposite, or whorled, entire, without stipules, often marked with shortly linear raphides. Flowers in terminal or axillary inflorescences, perfect or unisexual and dioecious, actinomorphic, hypogynous, cymose or sometimes solitary, usually with 1–3 enlarged bracts or involucre, the involucre persistent or deciduous, often colored and petaloid of free or connate segments and enclosing 1 or more flowers. Perianth usually colored and petaloid but apetalous, tubular, urceolate, campanulate, funnelform, or salverform, involute-plicate in aestivation, base of the tube persistent in fruit, the limb truncate or 3–7toothed or -lobed, persistent or deciduous. Stamens 1–30, usually as many as the perianth lobes and alternate with them, but also fewer or more numerous; filaments unequal, or equal, distinct or connate at the base, included or exserted; anthers dorsifixed near the base, longitudinally dehiscent; pollen tricolpate to pantoporate. Ovary 1-locular, sessile or stipitate, with one basal stalked ovule; ovule campylotropous or infrequently hemitropous; style slender, short or elongate, sometimes absent; stigma, simple, capitate, peltate, or fimbriate. Fruit a fleshy, coriaceous, or woody, indehiscent achene or nut, often enclosed within the persistent indurated base of the perianth to form an anthocarp. Seed with scant or abundant endosperm and straight or strongly curved embryo. Tropical and subtropical regions of the New World and Old World (mainly New World), a few species in temperate regions; 34 genera and ca. 350 species, 6 genera and 43 species in the flora area. Key to the Genera of Nyctaginaceae 1. 1. 2(1). 2. 3(2). 3. 4(3).

4.

Leaves alternate ................................................................... 2. Bougainvillea Leaves opposite, subopposite, or whorled ................................................. 2 Plants with spines ............................................................................ 6. Pisonia Plants without spines ................................................................................ 3 Herbs or suffruticose plants ...................................................................... 4 Shrubs, vines, or trees ............................................................................... 5 Flowers surrounded by 5-parted calyx-like whorl of united bracts; perianth tube 1–5 cm long, brightly colored, and uniformly and corolla-like ................................................................................................... 4. Mirabilis Flowers not surrounded as above; the subtending bracts inconspicuous and separate; perianth tube 1.5–5 mm long, separated into a lower calyx-like base and an upper corolla-like portion ................. 1. Boerhavia

102

5(3).

5.

N YCTAGINACEAE

Staminate flowers with exserted stamens, the perianth campanulate; pistillate flowers with the stigma exserted at anthesis; pistil borne on a short stipe; fruit usually black or dark purple, usually longitudinally ridged when dry ......................................................................... 3. Guapira Staminate flowers with included stamens, the perianth tubular to ellipsoid; pistillate flowers with the stigma rarely exserted at anthesis; pistil sessile or merely narrowed at the base; fruit often yellow, orange, or purple, sometimes longitudinally striate when dry ....................... 5. Neea

1. BOERHAVIA L., Sp. Pl. 3. 1753, spelling variant: Boerhaavia. Annual or perennial herbs, sometimes ligneous at base, with erect or prostrate, usually much branched, pubescent, sometimes glandular stems, frequently viscous at the nodes. Leaves opposite or subopposite, frequently unequal, entire, or undulate, bearing conspicuous raphides seen as surface irregularities upon drying. Inflorescence terminal, paniculiform or racemiform, many-flowered. Flowers small, bisexual, umbellate, cymose, capitate, or racemose, bracteate with 1–3 small, slender bracteoles subtending each flower. Perianth corolla-like, the tube campanulate or nearly rotate, persistent, constructed above the ovary, the limb upwardly shallowly 5-lobed. Stamens 1–5, included or exserted; filaments filiform, unequal, connate at the base. Ovary stipitate; style slender; stigma peltate or capitate, usually exserted. Fruit obovate, subellipsoid, or obpyramidal, 3–5-angled or rarely winged, glabrous or glandular-pubescent. Widely distributed in the tropics and subtropics of both the Old and New Worlds; ca. 40 species, 4 in Venezuela, 2 of these in the flora area. Key to the Species of Boerhavia 1. 1.

Inflorescence usually small, few-branched; flowers in clusters of 3–12; leaves and stems generally densely viscid-pubescent ............. B. coccinea Inflorescence frequently large, diffuse and much branched; flowers in clusters of 2–5; leaves and stems not densely viscid-pubescent, often with thin hairs 0.1–1 mm long .................................................... B. diffusa

Boerhavia coccinea Mill., Gard. Dict. ed. 8, Boerhavia no. 4. 1768. —Hierba de boca. Boerhavia caribaea Jacq., Observ. Bot. 4: 5. 1771. Decumbent or sprawling herbaceous plant to 0.8 m tall and 1.2 m long; flowers 2–3 mm long, rose-purple to deep red or red-purple. Disturbed areas near habitations, alluvial soils along streams, 100–200 m; Bolívar (Río Cuyuní at San Martin de Turumbán, Río Yuruani at Santa María del Capos). Scattered in northern and western Venezuela; widely distributed in Central America, South America, Asia, and Africa. ◆Fig. 106. Boerhavia diffusa L., Sp. Pl. 3. 1753. Boerhavia paniculata Rich., Actes Soc.

Hist. Nat. Paris 1: 105. 1792. —Tostón. Prostrate or ascending, much-branched, herbaceous plant to 1 m tall; leaves strongly unequal, the lower surface usually white; flowers about 2 mm long, pink, red, or purplish; fruit narrowly obovoid, thickest near apex. Weedy plants in disturbed areas in villages, along roadsides, fence rows, airports, and open soils, near sea level to 200 m; Delta Amacuro (Pedernales, Tucupita), Bolívar (Ciudad Bolívar, Jabillal on middle Río Caura, La Vergareña, San Martin de Turumbán to Casa Blanca road), Amazonas (Puerto Ayacucho, San Fernando de Atabapo). Common elsewhere in Venezuela; widely distributed from the southern U.S.A., Mexico, Central America, West Indies, South America, and the Old World tropics.

Bouganvillea 103

Fig. 106. Boerhavia coccinea

Fig. 107. Bouganvillea spectabilis

104

N YCTAGINACEAE

2. BOUGAINVILLEA Comm. ex Juss., Gen. Pl. 91. 1789, nom. cons., “Buginvillaea.” Shrubs, small trees, or most frequently woody vines, frequently armed with spines. Leaves alternate. Flowers perfect, without an involucre, usually in a 3-flowered axillary inflorescence of 3 large, colored, persistent bracts, one flower borne on the inner surface of each bract with its pedicel merging with the midrib of the bract. Perianth tubular, the limb small, 5-lobed, the lobes induplicate-valvate; tube terete or 5-angled. Stamens 5–10; filaments somewhat unequal, connate at base. Ovary stipitate, fusiform, slightly compressed laterally; style short, included. Anthocarp coriaceous, fusiform, 5-costate. Venezuela, Ecuador, Peru, central Brazil, Paraguay, northwest Argentina, cultivated in the New World tropics and subtropics; 14 species, 3 in Venezuela, 2 of these in the flora area. Because of its widespread cultivation, Bougainvillea is a difficult genus to study taxonomically. The species most commonly cultivated in Venezuela are Bougainvillea glabra Choisy and B. spectabilis Willd. Key to the Species of Bougainvillea 1. 1.

Young stems and leaves glabrous to puberulent; perianth tube with curving hairs 0.3–0.6 mm long ......................................................... B. buttiana Young stems and leaves pilose; perianth tube with erect hairs ca. 0.1 mm long ......................................................................................... B. spectabilis

Bougainvillea buttiana Holttum & Standl., Publ. Field Mus. Nat. Hist, Bot. Ser. 23: 44. 1944. —Trinitaria. Spiny shrub or climber; stems with straight thorns, mostly glabrous; leaves ovate-rounded or broadly elliptic-ovate, truncate or broadly rounded at base; bracts red or orange. Roadsides, ca. 100 m; Bolívar (lower Río Cuchivero basin). Native of Peru; rarely escaped and persisting.

Bougainvillea spectabilis Willd., Sp. Pl. 2: 348. 1789. —Trinitaria. Spiny shrub or climber; stems with straight thorns, pilose; bracts red. Roadsides, 100–200 m; Bolívar (Guayapo, lower Río Cauca). Native of Brazil, cultivated in Venezuela, the tropics, and subtropics. ◆Fig. 107.

3. GUAPIRA Aubl., Hist. Pl. Guian. 308, t. 119. 1775. Torrubia Vell., Fl. Flum. 139. 1825; Fl. Flum. Icon. 3: t. 150. 1827. Dioecious trees and shrubs. Leaves opposite or subopposite, sometimes whorled, often unequal, entire, pinnately veined. Inflorescence terminal or lateral pedunculate cymes with the branching opposite or subopposite, densely minutely brownish red or ferruginous-puberulent. Flowers small, reddish, greenish, greenish yellow, whitish, not involucrate, sessile or pedicellate, usually in groups of 2 or 3 on the ultimate branches of the inflorescence, subtended by 1–3 minute bracts. Staminate perianth obconic, campanulate, or funnelform, shallowly 5-lobed or dentate, the teeth induplicate-valvate. Stamens 5–10; filaments slender, unequal, united at the base to the stipe of the pistillode; anthers versatile, exserted. Pistillate perianth tubular, shallowly 5-dentate; staminodes with large sterile anthers, included within the perianth tube. Pistil narrowly ovoid with a basal stipe adnate to the filament tube; style 1; stigma fimbriate or penicillate. Fruit red or black, drupaceous,

Guapira 105

the exocarp fleshy, the anthocarp formed by enclosure of the ovary within the succulent perianth tube, usually longitudinally striate in the dried state; embryo straight. Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay; ca. 50 species, 16 in Venezuela, 13 of these in the flora area The genus Guapira is much in need of revision. Key to the Species of Guapira 1. 1. 2(1). 2.

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

6.

7(5). 7. 8(7). 8. 9(8).

9.

Leaves 25–30 × > 15 cm; fruits ca. 2 cm long ............................ G. sipapoana Leaves 0.1–15 × 0.5–15 cm; fruits < 1.5 cm long ....................................... 2 Inflorescence globose or subhemispheric, densely flowered ..... G. ferruginea Inflorescence variously branched, the flowers usually pedicellate, or if sessile, the leaves rounded or obtuse at apex and 1.8 cm wide or less ................................................................................................................ 3 Lower and/or upper surface of leaves, or lower midrib, pilosulous with lax, spreading, or divaricate hairs ........................................................ 4 Lower surface of leaves, including midrib, glabrous or pubescent, but the indument not spreading or divaricate .................................................. 5 Peduncle 7–9.5 cm long; petiole and young stems densely pubescent with spreading hairs 0.2–0.5 mm long ................................... G. marcano-bertii Peduncle (1–)3.5–5 cm long; petiole and young stems with hairs < 0.1 mm long ........................................................................................... G. rusbyana Lower surface of leaves, midrib, or veins with a minute tomentum of nonspreading hairs ................................................................................ 6 Lower surface of leaves glabrous or essentially so, the midrib or veins with scattered microscopic tomentum .................................................. 7 Leaves coriaceous, lower surface covered completely by dense rufo-ferruginous tomentum, the tertiary venation on both surfaces very conspicuous or elevated ........................................................ G. sancarlosiana Leaves chartaceous to subcoriaceous, lower surface sparsely rufous-tomentose to glabrescent, but never covered with dense rufo-ferruginous tomentum; tertiary venation either not evident, inconspicuous, or not elevated ................................................................................... G. cuspidata Peduncle and petiole with dense ferruginous tomentum ...... G. amacurensis Peduncle and petiole glabrous or sparsely pubescent .............................. 8 Peduncle and/or axes of inflorescence sparsely to moderately puberulent ................................................................................................................ 9 Peduncle and/or axes of inflorescence glabrous ...................................... 10 Leaves often broadest above the middle, conspicuously venose; lateral veins conspicuous, 9–11 on each side, subelevated or impressed on both sides, conspicuously anatomosing with the tertiary veinlets, ascending to an angle of 45° or more; tertiary veinlets forming a prominent network ............................................................................. G. fragrans Leaves often broadest near the middle, not venose, opaque; lateral veins inconspicuous, 5 or 6 on each side, impressed, divaricately spreading at an angle of 15–30°; tertiary veinlets obsolete or inconspicuous ................................................................................................ G. eggersiana

106

N YCTAGINACEAE

10(8). Tertiary veinlets prominent and finely reticulate on both sides of leaf blades ................................................................................................... 11 10. Tertiary veinlets, if present, inconspicuous and subreticulate .............. 12 11(10). Leaves 7–18 × 3–8 cm; apex acute to acuminate; petiole 0.8–2 cm long; staminate inflorescence penducle 4–8 cm long .................. G. bolivarensis 11. Leaves 1–6 × 0.5–2.5 cm; apex rounded or manifestly obtuse; petiole 0.1– 0.5 cm long; staminate inflorescence penducle not longer than 3 cm .............................................................................................. G. microphylla 12(10). Leaves oblanceolate to elliptic-lanceolate; staminate perianth cylindrictubular, 1.8–2 mm wide; stamens 8 ...................................... G. glabriflora 12. Leaves ovate or elliptic-ovate; staminate perianth ± funnel-shaped, 3.2 mm wide; stamens 10 .................................................................. G. neblinensis Guapira amacurensis Steyerm., Ann. Missouri Bot. Gard. 74: 615. 1987. —Casabe, Casabito, Mijarro. Tree 15–25 m tall; leaves inequilateral at base, dark brown to blackish upon drying; staminate perianth pale green, 7 mm long. Seasonally dry evergreen forests, ca. 300 m; Bolívar (east northeast of El Palmar, Represa Guri, east of Río Grande). Endemic. Guapira bolivarensis Steyerm., Ann. Missouri Bot. Gard. 74: 616. 1987. Tree 6–15 m tall; leaves 6–17 × 3–8 cm, glabrous on both sides; peduncle of staminate inflorescence 4–7.8 cm long, peduncle of pistillate inflorescence 1.8–3.3 cm long; stamens 10 in the staminate inflorescence. Seasonally dry evergreen forests, 500–1100 m; Bolívar (Amaruay-tepui). Brazil (Maranhão). ◆Fig. 108. Guapira cuspidata (Heimerl) Lundell, Wrightia 4: 80. 1968. —Pisonia cuspidata Heimerl, Bot. Jahrb. Syst. 21: 628. 1896. —Torrubia cuspidata (Heimerl) Standl., Contr. U.S. Nat. Herb. 18: 100. 1916. —Casabe, Casabe marrón, Cazabe, Gregarito, Ihchan-tó-pa-mëntu-kinke-tipeñ (Panare), Manteco, Merewadi (Yekwana), Palo de mono. Guapira ayacuchae Steyerm., Ann. Missouri Bot. Gard. 74: 615. 1987. Shrub or tree 4–25 m tall; leaves 5.5–15 × 3–8.5 cm, broadly ovate, acute to cuspidate at apex, rounded to acute at base; petioles rufous; peduncle brick-red; fruit red. Exposed igneous outcrops on dry forested slopes, seasonally dry semi-evergreen forests, gallery

forests, semi-deciduous forests, riparian forests, 50–600 m; Delta Amacuro (east-northeast of El Palmar, east of Río Grande), Bolívar (Altiplanicie de Nuria, vicinity of Ciudad Bolívar, region of El Manteco, El Perú, Guaniamo, vicinity of Panare and Túriba, Río Asa, lower Río Caura, Río Orocopiche near Ciudad Piar, affluent of Río Paragua, San Félix, Tumeremo), Amazonas (vicinity of Puerto Ayacucho and San Carlos, Río Coromoto, Río Cunucunuma, Samariapo). Monagas, Sucre; Colombia, Trinidad, Tobago, Guyana, Suriname. ◆Fig. 109. Guapira ferruginea (Klotsch ex Choisy) Lundell, Wrightia 4: 80. 1968. —Pisonia ferruginea Klotsch ex Choisy in A. DC., Prodr. 13(2): 445. 1849. —Torrubia ferruginea (Klotsch ex Choisy) Standl., Contr. U.S. Nat. Herb. 18: 100. 1916. —Casabe, Casabito. Torrubia cepalantha Standl. ex Pittier, Man. Pl. Usual. Venez. 177. 1926. Guapira davidsei Steyerm., Ann. Missouri Bot. Gard. 74: 617. 1987. Deciduous shrub or tree 3–15 m tall; young branchlets densely rufous-tomentulose; young leaves ferruginous-pubescent, mature ones glabrate to glabrous; perianth greenish to yellow or green with brown margins; filaments white, anthers cream-colored; fruit 7 mm long. Exposed igneous outcrops in deciduous or semi-deciduous forests, sometimes bordering streams, 100–400 m; Delta Amacuro (east-southeast of Los Castillos de Guayana), Bolívar (Altiplanicie de Nuria, valley of Caño Colorado east of Arekuna,

Guapira 107

southeast of El Callao, Represa Guri, Río Maniapure, Sierra de Imataca). Aragua, Carabobo, Falcón, Guárico, Lara, Miranda, Portuguesa, Yaracuy. ◆Fig. 110. Guapira fragrans (Dum.-Cours.) Little, Phytologia 17: 368. 1968. —Pisonia fragrans Dum.-Cours., Bot. Cult. ed. 2, 7: 114. 1814. Torrubia flagrans Dum.-Cours., Bot. Cult. ed. 2, 7: 114. 1814. Pisonia eucalyptifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 7: 197. 1825. Tree 5–15 m tall; leaves glabrous on both sides, obovate-oblong, oblanceolate, obovate, or rhombic-elliptic; inflorescence rather few-flowered with puberulent or glabrate branches; perianth puberulent to nearly glabrous. Gallery forests, riparian rainforests, dry forests, 100–200 m; Bolívar (northwestern portion, Represa Guri, Río Villacoa), Amazonas (San Carlos de Río Negro). Widely distributed in northern and central Venezuela, also Apure and Nueva Esparta; West Indies, northern Colombia. Guapira glabriflora Steyerm., Ann. Missouri Bot. Gard. 74: 617. 1987. Small tree with branchlets, petioles, and leaves glabrous; stamens 8. Evergreen lowland forests, 100–200 m; Amazonas (vicinity of San Carlos). Endemic. Guapira guianensis Aubl., Hist. Pl. Guian. 308, t. 119. 1775. —Casabe morado, Cazabe, Cazabito, Ishakyek (Pemón). Shrub or tree 3–15 m tall, with glabrous or sparsely puberulent branchlets; leaves chiefly glabrous; inflorescence on elongate peduncles, glabrous or sparsely puberulent; flowers greenish; fruit dark purple. Seasonally dry to evergreen lowland and montane forests, gallery forests, rainforests, 100– 1300 m; Delta Amacuro (Caño Guayamuni, east-northeast of El Palmar, east of Río Grande), Bolívar (common in the eastern sector from the Altiplanicie de Nuria south to Santa Elena de Uairén). Anzoátegui, Aragua, Miranda, Monagas, Nueva Esparta, Sucre; Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil (Amapá). Guapira guianensis has been confused with G. olfersiana (Link, Klotzsch & Otto)

Lundell, a species of southern Brazil, Ecuador, and Bolivia. Guapira olfersiana has a broader perianth more widely spreading above and a more ferruginous-pubescent pistillate perianth. Guapira marcano-bertii Steyerm., Ann. Missouri Bot. Gard. 74: 618. 1987. —Cazabe, Gregorito. Tree with densely pubescent young stems; petioles, peduncles and leaves; petioles 1–2 cm long; leaf blades 7–12.5 × 3–7 cm, broadly elliptic-ovate; infructescence subumbellate to irregularly paniculate; fruit 9–10 mm long. Seasonally dry evergreen forests, ca. 200 m; Delta Amacuro (northeast of El Palmar, east of Río Grande), Bolívar (Temblador on lower Río Caura). Endemic. Guapira microphylla (Heimerl) Lundell, Wrightia 4: 82. 1928. —Pisonia microphylla Heimerl, Oesterr. Bot. Z. 56: 425. 1906; Heimerl in Urb., Symb. Antill. 7: 215. 1912. —Torrubia microphylla (Heimerl) Standl., Contr. U.S. Nat. Herb. 18: 100. 1916. —Casabe, Palo de mono. Shrub or small tree to 10 m tall; with ferruginous-puberulent branchlets; leaf blades and petioles shorter than in the other Guayanan species; petioles 1.5–5 mm long; inflorescence small, 7–15 mm long, 11–20 (–30) mm broad, sparsely appressed-pubescent; flowers sessile or nearly so; staminate perianth 3.5–4 mm long, pistillate perianth 2–2.2 mm long. Gallery forests, savanna-dry forest ecotone, 100–200 m; Bolívar (Altiplanicie de Nuria, Guaniamo, lower Río Caroni, Río Chaviripa, affluent of middle Río Orinoco). Anzoátegui, Apure, Aragua, Carabobo, Falcón, Miranda, Nueva Esparta, Sucre, Táchira; West Indies. Guapira neblinensis Maguire & Steyerm., Mem. New York Bot. Gard. 51: 129. 1989. Small tree to 5 m tall with mainly glabrous leaves and peduncles; inflorescence relatively few-flowered; staminate perianth 5.5 mm long, pistillate, 7–7.5 mm long. Evergreen lowland to montane forests, 100–700 m; Amazonas (Piedra Nunca north of Piedra Cocuy, Sierra de la Neblina). Endemic.

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Fig. 108. Guapira bolivarensis

Fig. 109. Guapira cuspidata

Mirabilis 109

Fig. 110. Guapira ferruginea

Guapira rusbyana (Heimerl ex Standl.) Lundell, Wrightia 4: 83. 1968. —Pisonia rusbyana Heimerl ex Standl., Field Mus. Nat. Hist., Bot. Ser. 11: 85. 1931, nom. nud. —Torrubia rusbyana Heimerl ex Standl., Field Mus. Nat. Hist., Bot. Ser. 11: 85. 1931. Shrub or tree to 8 m tall; with densely ferruginous-pubescent branches; leaves opposite or whorled, 6–14 × 2–6.5 cm, lanceolateoblong, elliptic-oblong, or elliptic; staminate perianth 3.5–4 mm long, minutely puberulent to glabrous; stamens ca. 8. Riparian forests, 50–100 m; Delta Amacuro (lower Río Orinoco), Bolívar (Río Botanamo, region of Tumeremo to Los Castillos de Guayana). Distrito Federal. Guapira sancarlosiana Steyerm., Ann. Missouri Bot. Gard. 74: 619, fig. 2. 1987. —Congrio, Palo chaparro, Palo chaparro banero.

Tree 4–12 m tall with dense rufous-ferruginous pubescence on young branchlets; leaves coriaceous, 7.5–14.5 × 4.5–7.5 cm, the lower surface with dense rufous-ferruginous pubescence; lateral veins 9–11 each side; staminate perianth 4–4.8 mm long; stamens 5 or 6. White-sand scrub (bana), low Amazonian caatinga forests, 100–200 m; Amazonas (region of San Carlos de Río Negro). Endemic. Guapira sipapoana Steyerm., Ann. Missouri Bot. Gard. 74: 620, fig. 3. 1987. Shrub or small tree with densely rufoustomentose young branches; leaves ellipticovate, rounded at base, the lower surface puberulous, prominently reticulate; peduncle 3.8 cm long, densely rufous-tomentose; fruit linear-ellipsoid, ca. 2 cm long, rufous-tomentose. Lowland riparian forests, 100–200 m; Amazonas (Río Sipapo). Endemic. Guapira sipapoana has the largest leaves of any of the Venezuelan species of Guapira.

4. MIRABILIS L., Sp. Pl. 177. 1753; Gen. Pl. no. 39. 1737. Annual or perennial herbs or small subshrubs, sometimes woody below, usually branched and often smaller at the nodes. Leaves opposite, simple, entire or undulate, pinnately veined cystoliths, without stipules. Inflorescence usually of small cymes or clusters, terminal or axillary. Flowers bisexual, usually 1–several subtended by united leaf-like bracts forming a 5-parted involucre. Perianth campanulate, funnelform, or salverform, 5-lobed, the lobes induplicate-valvate in bud, de-

110

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ciduous after anthesis. Stamens 3–5, usually exserted; filaments slender, unequal, shortly connate at base. Ovary ovoid or subglobose; style filiform; stigma capitate, long-papillate. Fruit (anthocarp) globose to cylindric or obovoid, terete, 5-angled or 5-sulcate, rugose or tuberculate, glabrous or puberulent, mucilaginous when wet; embryo sharply curved. Tropical and temperate parts of the Americas, 1 species in southeastern Asia; ca. 60 species, 3 in Venezuela, 1 of these in the flora area. Mirabilis jalapa L., Sp. Pl. 177. 1753. —Buenas tardes, Don diego de noche, Jasmín colorado, Jasmín de cafetal, Jasmín de tarde, Pasaña. A much-branched annual or perennial herb to 1.3 m tall; leaf blades triangular to ovate-deltoid with petioles 0.8–6 cm long; flowers usually rose-purple or magenta, but also pink, white, yellow, or orange, or variously striped, very fragrant, especially at night. Waste and cultivated ground, some-

times in cocoa plantations, 50–200 m; Delta Amacuro (Tucupita). Bolívar (north of El Manteco). Throughout Venezuela, where planted for its oranamental flowers; originally native apparently to Mexico, but cultivated in most tropical countries and grown as a summer garden flower in temperate regions, where the English name is usually Four-O’clock. ◆Fig. 111.

Fig. 111. Mirabilis jalapa

5. NEEA Ruiz & Pav., Fl. Peruv. Prodr. 52. 1794. Trees or shrubs. Leaves opposite or in whorls, sometimes subopposite or alternate. Inflorescence terminal, sometimes axillary or cauliflorous, cymose, paniculate, or thyrsiform. Flowers small, white, green, or reddish, unisexual, dioecious, sessile or pedicellate, often in groups of 3s and 1–3-bracteolate at the base of the perianth. Staminate perianth ellipsoid, ovoid, or urceolate, 4- or 5-lobed or -dentate at the apex. Stamens usually 8, sometimes 5–10, included within the perianth tube; filaments unequal in length, coalesced at the base; pistillode present. Pistillate perianth urceolate or tubular-cylindric, constructed below the 4 or 5 teeth or lobes, usually smaller than the staminate perianth; staminodes present, lacking nonfunctional anthers. Pistil often exserted; stigma penicillate or fimbriate. Fruit a fleshy anthocarp, deep red, purple, pink, or yellow, ellipsoid or linear-oblong, smooth ex-

Neea 111

cept for the longitudinal ribs when dry. Seed hard, striate or costate with little endosperm; embryo straight. Neotropics; ca. 85 species, 30 in Venezuela, 24 of these in the flora area. The genus is greatly in need of revision. The Amazonian species of Neea are well known as “palo de culebra,” and the crushed bark has a good reputation as an antidote to snake bites. Key to the Species of Neea 1.

Leaves sessile or nearly so, usually obtuse at base, the petiole 1 mm long; young stem and petiole hirtellous with subspreading rufous-brown hairs; inflorescence both axillary and terminal on the stem ... N. ignicola 1. Leaves petiolate, generally acute to acuminate at base; the petiole 2– 35 mm long; young stem and petiole glabrous or pubescent with appressed hairs; inflorescence either cauliflorous on the old stem or terminal on the branches ........................................................................... 2 2(1). Inflorescence cauliflorous on the old stem ................................................ 3 2. Inflorescence terminating the stem or its branches ................................. 7 3(2). Leaves oblanceolate-elliptic or lance-elliptic, 8–12.5(–15) × 2.5–4.5(–5.5) cm .................................................................................................. N. clarkii 3. Leaves mainly obovate, oblong, or oblong-ovate, 16–30 cm long ............. 4 4(3). Principal secondary leaf veins 6–8 each side, slightly elevated on lower surface ......................................................................... N. brevipedunculata 4. Principal secondary leaf veins 8–12 each side, conspicuously elevated on lower surface .......................................................................................... 5 5(4). Lower leaf surface glabrous, but midrib and secondary veins with minute spreading hairs; flowering peduncle 0.5–0.6 cm long ............... N. liesneri 5. Lower leaf surface and veins completely glabrous; fruiting peduncle 1– 3.5 cm long ............................................................................................. 6 6(5). Flowers sessile; inflorescence dichotomous or with short axes on an elongated rachis ......................................................................... N. davidsei 6. Flowers pedicellate; inflorescence a panicle, not dichotomous, branched into 3–8 secondary axes ......................................................... N. floribunda 7(2). Leaves broadly rounded at the apex, obovate or elliptic-obovate ..................................................................................................... N. obovata 7. Leaves mainly acute to acuminate at the apex, of shapes other than above ...................................................................................................... 8 8(7). Main secondary veins of leaf blades 15–25 each side, subhorizontal or ascending at an angle < 20°, relatively close together, 3–4 mm apart .................................................................................................. N. ovalifolia 8. Main secondary veins of leaf blades generally fewer than 12 each side (fainter intermediate veins may be present), ascending at an angle usually > 30°, or if more than 12 veins or at a smaller angle, then the veins 4 mm apart ................................................................................... 9 9(8). Peduncle 6–11 cm long ............................................................................ 10 9. Peduncle 1–5 cm long .............................................................................. 11 10(9). Stem, peduncle, and axes of inflorescence densely ferruginous-tomentose; perianth 3 × 1.5 mm, densely ferruginous-tomentose ........... N. bernardii

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N YCTAGINACEAE

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(15). 19. 20(19).

Stem glabrous, peduncle and axes of inflorescence minutely puberulous or glabrescent; perianth 8–9 × 3 mm, glabrous ......................... N. grandis Peduncle and axes of inflorescence glabrous throughout or essentially so .............................................................................................................. 12 Peduncle and axes of inflorescence sparsely to densely ferruginous or puberulent, sometimes with glandular trichomes ................................. 15 Staminate perianth slightly or moderately ferruginous-pubescent toward base ................................................................................................ N. clarkii Staminate perianth glabrous or sometimes sparsely puberulous near base, or minutely papillate apically .................................................... 13 Staminate perianth 6–6.5 × 3.5–4 mm; inflorescence paniculately and irregularly branched, large and conspicuous with elongated rachis and axes, 6–15 cm long, 5–10 cm wide; peduncle 3–5 mm diameter; principal secondary veins 9–12 each side; tertiary veinlets conspicuously reticulate and subelevated on upper leaf surface ..................... N. robusta Staminate perianth 2–3.5 × 0.8 mm; umbellately or trichotomously branched, 1–2 cm long (or high), 2.5–4 cm wide; peduncle 0.8–1.5 mm diameter; principal secondary veins 5–7 each side; tertiary veinlets obsolete in upper surface ..................................................................... 14 Upper surface of leaf lustrous, lower surface with subelevated, subreticulate tertiary venation ± manifest; leaves ovate to subovate, obtuse or rarely rounded at apex; peduncle (pistillate) 1–1.5 mm diameter .................................................................................. N. tepuiensis Upper leaf surface dull; lower leaf surface with obsolescent tertiary venation; leaves oblanceolate, obtusely acute to acute at apex; peduncle (staminate) 0.8–1 mm diameter .......................................... N. subglabrata Junction of lowest axes of inflorescences with summit of peduncle enlarged, 2.5–4 mm wide ........................................................................ 16 Junction of lowest axes of inflorescence with summit of peduncle not manifestly enlarged, 0.8–2 mm wide .................................................. 19 Perianth 3–4.5 mm long, 1.5 mm wide; petiole and young stem sparsely puberulent .............................................................................. N. sebastianii Perianth 6–8 mm long, 2–3.5 mm wide; petiole and young stem glabrous or glabrescent ....................................................................................... 17 Tertiary venation manifest, elevated on lower surface, impressed on upper surface; flowers in small cluster at or near the end of the ultimate axes ........................................................................................ N. neblinensis Tertiary venation obsolete or scarcely manifest; flowers, if clustered, mainly scattered along the length of the ultimate axes .................... 18 Perianth glabrous without; leaves drying blackish; principal secondary veins 9–12 each side ..................................................... N. huachamacarae Perianth with minute, sparse to moderate ferruginous indument; leaves drying blackish brown; principal secondary veins 6–8 each side ........................................................................................... N. mapourioides Peduncle usually with paired ferruginous bracts .................................. 20 Peduncle ebracteate ................................................................................. 21 Primary axes of inflorescence 4–7, umbellate; pistillate perianth ferruginous-tomentose; main lateral veins of leaf blades sharply ascending at 50°–60° ..................................................................................... N. bracteosa

Neea 113

20.

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

22.

23(22).

23.

24(22).

24.

25(24).

25.

Primary axes of inflorescence 3, not umbellate; staminate perianth glabrous except for papillate exterior of lobes; main lateral veins of leaf blades shallowly ascending at 15°–25° ........................... N. amaruayensis Perianth glabrous or glabrescent ........................................... N. marahuacae Perianth variously pubescent .................................................................. 22 Ultimate axes of inflorescence racemose with alternately arranged flowers; upper and lower midribs of the leaf blade with minute, ferruginous indument .............................................................................................. 23 Ultimate axes of inflorescence cymose or corymbiform, or with fasciculately arranged flowers; upper midrib of leaf blades glabrous, the lower midrib glabrous or the basal part sparsely puberulent ........... 24 Lower leaf surface covered with an appressed dense indument; perianth (staminate) broadly urceolate; inflorescence much branched; perianth and inflorescence axes densely ferruginous-tomentose with some glandular hairs; peduncle to 1.2 cm long .................................... N. parimensis Lower leaf surface glabrous or glabrescent; perianth (pistillate) subfunnelform; inflorescence sparsely branched; perianth and inflorescence axes rather sparsely ferruginous-tomentose without glandular hairs; peduncle 2–4 cm long .......................................................... N. tristis Petiole and upper part of stem with pale, minute, spreading papilla-like trichomes; perianth with vermiform, short, appressed hairs; lower leaf surface not densely dark-punctate ........................................ N. cedenensis Petiole glabrous or sometimes minutely glandular; stem glabrous; perianth either minutely appressed puberulent or densely glandular; lower leaf surface densely dark-punctate ........................................... 25 Tertiary venation scarcely evident on any part of the leaves; axes and peduncle of inflorescence densely ferruginous-pubescent; fruit 3.5–5 mm wide ............................................................................................... N. clarkii Tertiary venation grossly reticulate, subelevated and manifest on both sides of leaves; axes and peduncle of inflorescence sparsely to moderately glandular-pubescent; anthocarp 6–7 mm wide ..... N. guaiquinimae

Neea amaruayensis Steyerm., Ann. Missouri Bot. Gard. 74: 623. 1988. Shrub or tree 2.5–5 m tall; leaves mainly opposite, 6.5–15 × 2.5–7.5 cm, elliptic-ovate or oblong- or lance-elliptic, glabrous on both sides; inflorescence small; staminate perianth greenish white, 3–3.5 mm long. Montane forested slopes of seasonally dry evergreen forests, 600–900 m; Bolívar (Amaruaytepui, drainage of Río Acanán). Endemic. Neea bernardii Steyerm., Ann. Missouri Bot. Gard. 74: 623. 1988. Shrub or tree to 40 m tall; with densely ferruginous-tomentose branches, peduncle, and inflorescence; leaves opposite, 5.5–11.5 × (2.7–)3–6.5 cm, broadly ovate, elliptic-ovate, or oblanceolate-elliptic; staminate perianth 3–4 mm long. Seasonally dry evergreen ri-

parian forests, 300–900 m; Bolívar (vicinity of Santa Elena de Uairén and El Palmar). Endemic. Neea bracteosa Steyerm., Ann. Missouri Bot. Gard. 74: 624. 1987. Tree ca. 8 m tall; leaves opposite or in 3s at the upper nodes, 6–9 × 2–3.5 cm, elliptic or lance-elliptic, the adult glabrous; inflorescence umbelliform with 4–7 slender main axes bearing 2–5 umbellately disposed secondary axes. Dry forests bordering savannas, outcrops, 200–300 m; Bolívar (region of Represa Guri northeast of Ciudad Piar). Endemic. Neea brevipedunculata Steyerm., Ann. Missouri Bot. Gard. 24: 624, fig. 4. 1987. —Dukatu (Yekwana), Palo de culebra.

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Tree 6–20 m tall; leaves coriaceous, large, obovate, broadly oblanceolate or elliptic-oblong, abruptly obtusely acute or rarely rounded at apex; cauliflorous inflorescences at the usually thickened woody nodes; peduncle 1–1.5 cm long with widely divaricate primary axes of the inflorescence; staminate perianth 3.7 mm long, pistillate 3.5 mm. Lowland rainforests, white-sand scrub forests, 100–200 m; Bolívar (upper Río Caura), Amazonas (Culebra to Paso del Diablo, upper Río Orinoco, vicinity of San Carlos de Río Negro). Endemic. The bark of Neea brevipedunculata is crushed and used to treat snake, spider, and ant bites. Neea cedenensis Steyerm., Ann. Missouri Bot. Gard. 24: 625. 1987. —Morichalero. Shrub to small tree 5 m tall; flowers sessile, green. Seasonally dry evergreen forests, semi-evergreen forest, 100–200 m; Bolívar (Caicara-El Burro road, around Caicara del Orinoco, Chiviripa). Endemic. Neea clarkii Steyerm., Ann. Missouri Bot. Gard. 24: 625. 1987. —Palo de cachicamo, Palo de culebra. Shrub ca. 3 m tall, with glabrous branches; leaves 8–12.5(–15) × 2.5–5.5 cm, lanceolate-elliptic or oblanceolate-elliptic; lateral and tertiary venation scarcely evident; staminate perianth 4 mm long, minutely sparsely pubescent. White-sand scrub forests (bana), 100–200 m; Amazonas (vicinity of San Carlos de Río Negro, Tama Tama, Yavita to Maroa road). Endemic. Neea davidsei Steyerm., Ann. Missouri Bot. Gard. 74: 627. 1987. —Casabe, Ekedek (Pemón), Medewadi (Yekwana). Tree ca. 8 m tall; leaves opposite, broadly obovate or oblong-elliptic; flowers sessile; cauliflorous inflorescence without peduncle or peduncle 1–3.5 cm long; fruit 1.2 cm long, not ribbed. Seasonally dry evergreen forests, semi-evergreen to nonflooded forests, 200– 600 m; Delta Amacuro (Río Toro, vicinity of Sierra Imata and Rio Orocoima). Bolívar (south of El Dorado, El Plomo, middle Río Caura, Río Karún, mouth of Río Nichare). Endemic. Neea floribunda Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 46. 1838. —Casabe, Educadu

(Yekwana), Ekedek (Pemón), Ma-hanahana (Yekwana). Neea cauliflora Heimerl, Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 16c: 129. 1934. Tree 10–30 m tall; flowers and inflorescences pink; fruits green. Lowland nonflooded forests, 100–500 m; Amazonas (Cerro Huachamacari, Culebra, Río Cunucunuma, upper Río Orinoco), Bolívar (Río Ariza, upper Río Caura, middle Río Paragua). Colombia, Suriname, Ecuador, Peru, Brazil (Acre, Amazonas). Neea grandis Maguire & Steyerm., Mem. New York Bot. Gard. 51: 109. 1989. Tree to 10 m tall; leaf blades 8–23 × 5–12 cm, elliptic-oblong or elliptic-ovate, abruptly acute to subacuminate, glabrous throughout; pistillate peduncle 6–11 cm long. Montane forests, 1300–2000 m; Amazonas (Sierra de la Neblina). Endemic. Neea guaiquinimae Steyerm., Ann. Missouri Bot. Gard. 74: 627. 1987. Shrub or small tree 2–3 m tall; leaves 5–12.5 × 2–6 cm, elliptic-ovate, obtusely acute, the lower surface densely dark glanddotted; pistillate inflorescence trichotomously branched with 4 main axes; pistillate perianth 4 mm long. Montane forests bordering upland savannas, open rocky sandstone plateaus, 1500–1700 m; Bolívar (summit of Cerro Guaiquinima). Endemic. Neea huachamacarae Steyerm., Ann. Missouri Bot. Gard. 74: 628. 1987. —Jadajada (Yekwana), Palo de culebra. Tree ca. 10 m tall; leaves opposite, oblanceolate or oblong-elliptic, glabrous; pistillate perianth 7.5–8 mm long. Evergreen lowland to montane forested slopes, ca. 200–300 m; Amazonas (base of Cerro Huachamacari, Yavita to Maroa road). Endemic. Neea ignicola Steyerm., Ann. Missouri Bot. Gard. 74: 628. 1987. Shrub 2–3 m tall; leaves membranaceous, 3.5–6 × 1.3–2.5 cm, ovate-lanceolate, both surfaces glabrous; inflorescence ferruginoushirtellous with ferruginous subspreading hairs. Igneous outcrops among shrubby vegetation, ca. 100 m; Amazonas (region north of Puerto Ayacucho). Endemic. Neea liesneri Steyerm., Ann. Missouri Bot. Gard. 74: 629, fig. 5. 1987.

Neea 115

Tree 5 m tall; leaf blades large, broadly obovate, obtusely acute or rounded at the apex; cauliflorous staminate inflorescence many-branched, many-flowered from a short peduncle 0.5–0.6 cm long; staminate perianth greenish brown, 6 mm long; fruit oblong-ellipsoid, 20 mm long. Evergreen lowland forests, 50–100 m; Amazonas (base of Sierra de la Neblina). Brazil (Serra da Neblina). Neea mapourioides Steyerm., Ann. Missouri Bot. Gard. 74: 630. 1987. —Palo culebra. Shrub 3 m tall; leaves opposite, coriaceous, 5.5–17.5 × 4–7 cm, elliptic-obovate; inflorescence terminal with spreading primary axes; staminate perianth whitish, ellipsoid, 6.5–7 mm long. Wet forests, 100–200 m; Amazonas (Río Mawarinuma, vicinity of San Carlos de Río Negro). Endemic. The bark of Neea mapourioides is used to treat snake bite. Neea marahuacae Steyerm., Ann. Missouri Bot. Gard. 74: 631. 1987. Shrub 2 m tall; leaves 7.5–10.5 × 2.5–4.5 cm, elliptic-oblanceolate, acuminate, glabrous; staminate inflorescence terminal, subumbellately 5-branched, sparsely appressed-rufous brown pubescent. Montane forests near streams, ca. 1000 m; Amazonas (Cerro Marahuaka). Endemic. ◆Fig. 112. Neea neblinensis Maguire & Steyerm., Mem. New York Bot. Gard. 51: 112. 1989. —Palo de culebra, Palo perro de agua. Tree to 5 m tall; stems glabrous; leaves with manifest venation on lower surface. Montane forests, white-sand savannas, 100– 1300 m; Amazonas (Caño Colorado 55 km southeast of Puerto Ayacucho, La Esmeralda, San Carlos de Río Negro, Sierra de la Neblina). Endemic. Neea obovata Spruce ex Heimerl., Beitr. Syst. Nyctag. 38. 1897. —Arepillo, Cumello, Cumello banero, Cupi hoja fina. Shrub or small tree 2–5 m tall; leaves opposite, obovate, broadly rounded or submarginate at summit; inflorescence very slender; perianth subcampanulate, 3 mm long, densely rufous-tomentose; stamens 6. White-sand dwarf shrub thickets (bana), shrub islands in white-sand savannas, gallery forests, dwarf forested slopes overlying

sandstone (caatinga and bana), 100–300 m; Amazonas (frequently encountered from base of Cerro Aracamuni north to Río Sipapo east to Cerro Yapacana, Cerro Duida, Río Atabapo, Río Guainía, Río Negro, Río Orinoco, Río Sipapo, lower Río Ventuari, San Carlos de Río Negro, Santa Bárbara del Orinoco, Yavita). Amazonian Colombia, Peru, and Brazil. ◆Fig. 113. Unlike most of the species of Neea, the anthocarps of N. obovata are usually costate or striate longitudinally. Neea ovalifolia Spruce ex Schmidt in Mart., Fl. Bras. 14(2): 368. 1872. —Curamita, Curamito. Shrub or small tree 1–4 m tall; leaves chiefly 5–7.5 × 2.5–5 cm, oval, shortly acuminate or acute; inflorescence divaricately branched; peduncle sometimes red; pistillate perianth pale green, 2 mm long, ferruginoustomentose; mature fruit purple. Seasonally flooded forests near black-water rivers, riparian forests on sandy soils and abandoned cut-over land, ca. 100 m; Amazonas (Macabana on Río Ventuari, vicinity of Puerto Ayacucho and Isla Ratón, Río Atacavi, Río Emoni in lower Río Siapa basin, San Carlos de Río Negro, 32 km south of San Carlos de Río Negro, 15 km southeast of San Fernando de Atabapo). Apure, Falcón, Táchira; Colombia, Guyana, French Guiana, Amazonian Brazil, Bolivia. Neea parimensis Steyerm., Ann. Missouri Bot. Gard. 74: 631. 1987. Tree 10 m tall; leaves membranous, 4–7 × 1.5–2.5 cm, oblanceolate; staminate perianth urceolate, 4 mm long; stamens 7. Gallery forests, ca. 800 m; Amazonas (Río Matacuni, Sierra Parima). Endemic. ◆Fig. 114. Neea robusta Steyerm., Ann. Missouri Bot. Gard. 74: 632. 1987. —Bejuco ciruelo, Palo de culebra. Tree 3–5 m tall; glabrous throughout; leaves coriaceous, 13–14.5 × 6.5–9 cm; staminate perianth suburceolate, 6–6.5 cm long; stamens 9–10. White-sand savannas, scrub forests (bana), flooded forests, 100–200 m; Amazonas (Caño Cotúa, Caño San Miguel, Caño Yagua, base of Cerro Yapacana, Río Caname, affluent of Río Orinoco, Río Pasimoni, Río Temi, region of San Carlos de Río Negro, Yavita). Adjacent Brazil (near base of Serra da Neblina).

116

N YCTAGINACEAE

Fig. 112. Neea marahuacae

Fig. 113. Neea obovata

Fig. 114. Neea parimensis

Pisonia 117

Neea sebastianii Steyerm., Ann. Missouri Bot. Gard. 74: 633. 1987. Tree ca. 8 m tall; leaf blades 7.5–14 × 3.5– 6.5 cm, ovate-elliptic, obtusely acute or rounded at apex; staminate inflorescence densely ferruginous-tomentose. Seasonally flooded forests, 100–200 m; Amazonas (Isla Sebastián along Río Casiquiare). Endemic. Neea subglabrata Steyerm., Ann. Missouri Bot. Gard. 74: 633. 1987. Tree ca. 5 m tall; mainly glabrous throughout; leaves 4–9.5 × 2.5–4 cm, narrowly cuneate at base; stamens 6. Forested slopes in seasonally dry evergreen forests bordering savannas, gallery forests, and montane forests, 300–1300 m; Bolívar (El Dorado, Gran Sabana, Río Aponguao, Ueitepui), Amazonas (slopes of Cerro Yutajé). Táchira; Guyana. Neea subglabrata has been confused with N. spruceana of Peru and Brazil, which has larger staminate flowers in a more widely

branched inflorescence and differently shaped leaves with more numerous, spreading, lateral veins. Neea tepuiensis Steyerm., Ann. Missouri Bot. Gard. 74: 634. 1987. Tree 4–5 m tall; glabrous throughout except for the glandular-pubescent terminal bud; leaves (2.7–4.3)7–9.5 × (1.5–3)3.5–5.5 cm; inflorescence and young fruit dull red; fruit ellipsoid-oblong, 9 mm long. Montane forested slopes overlying sandstone and bordering wet savannas, 1300–2000 m; Bolívar (Macizo del Chimantá [middle slopes Agparamántepui, summit Toronó-tepui]). Endemic. Neea tristis Heimerl, Notizbl. Bot Gar. Berlin-Dahlem 6: 131. 1914. Tree 3–4 m tall; fruits yellow-green. Lowland nonflooded forests, 100–200 m; Amazonas (Tamatama, Yapacana Sabana at the base of Cerro Yapacana). Ecuador, Brazil (Acre, Amazonas).

6. PISONIA L., Sp. Pl. 1026. 1753. Calpidia Thouars, Pl. Iles Afrique Austr. 37. 16 Apr 1804. Rockia Heimerl, Oesterr. Bot. Z. 63: 289. 1913. Heimerlia Skottsb., Svensk Bot. Tidskrift 30: 738. 1936. Heimerliodendron Skottsb., Svensk Bot. Tidskrift 35: 364. 23 Dec 1941. Small trees, shrubs, or climbers, glabrous or puberulent, with or without short, axillary, straight or curved spines; leaves opposite, subopposite, or occasionally alternate or whorled; blade decurrent on the petiole, simple and entire; venation pinnate, often developing from sides of the spines; flowers unisexual or occasionally bisexual, inflorescences solitary or fasciculate, axillary or terminal on highly modified axillary short shoots, the flowers often borne in cymes but the cymes in umbelliform, capitate, or corymbiform clusters; bracteoles 2–4, minute; flowers greenish, subtended by bracteolate pedicels; staminate flowers with a campanulate perianth tube, the tube with 5 obscure distal lobes or teeth, stamens 6–8(–10), exerted at anthesis, filaments slender, united near the base to form a small tube and adnate to the base of the pistillode; pistillate flowers with a narrow perianth tube, staminodes often in the form of a dentate disk adnate to the narrowed base (stipe) of the pistil, ovary 1-locular and with 1 ovule, style narrow and fimbriate stigma. Fruit an anthocarp formed by the union of perianth tube and pistil, narrowly elongate, terete or 5-angled in cross section, with longitudinal rows of stalked glands in 1 or several ranks, the fruit at maturity usually borne on greatly elongated peduncle and pedicels. Tropical and subtropical regions of the world, mainly America and Southeast Asia, with few species in Africa and Australia; ca 40 species, 3 in Venezuela, 1 of these in the flora area.

118

N YCTAGINACEAE

Fig. 115. Pisonia aculeata

Pisonia aculeata L., Sp. Pl. 1026. 1753. Tree or scandent shrub to 8 m tall; with spines on stems; ca. 300 m; Bolívar (Represa Guri), Amazonas (Río Ocamo–Raudal Arata). Aragua, Carabobo, Distrito Federal, Falcón, Guárico, Lara, Miranda, Portuguesa, Sucre,

Zulia; U.S.A. (Florida, Texas), southern Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina, Asia (Philippines, Taiwan), Africa (Ghana, South Africa). ◆Fig. 115.

NYMPHAEACEAE by John H. Wiersema Aquatic, rhizomatous herbs, rooted in substrate. Leaves arising directly from the rhizome, alternate, long-petiolate, simple, the mature blades mostly floating on the water surface (sometimes submersed or emergent), ovate-lanceolate to orbicular, peltate or more commonly with basal sinus. Flowers solitary, axillary, or extra-axillary, long-pedunculate, borne at or above the water surface (occasionally submersed), bisexual, radially symmetrical, usually showy. Sepals 4–6(–14), free, mostly greenish (flora area species) or yellow and ± petaloid; petals 4–many (flora area species) or rarely absent, free, often transitional to stamens. Stamens numer-

Nymphaea 119

ous, mostly spirally arranged, usually grading from outer laminar forms to inner forms with more distinctive filament and anther. Carpels 3–many, at least partially united, the ovary superior to inferior, with 3–many locules and 2–many ovules per locule with laminar placentation; ovules pendulous, anatropous (flora area species) or orthotropous; stigmas radiate on distal surface of ovary. Fruits irregularly dehiscent, berry-like or capsule-like, usually retracted beneath the water surface. Seeds several to numerous, operculate, arillate (flora area species) or exarillate, with a minute embryo, little endosperm, and abundant perisperm. Cosmopolitan; 6 genera and ca. 70 species, 1 genus and 7 species in the flora area. 1. NYMPHAEA L., Sp. Pl. 510. 1753, nom. cons. Perennial herbs. Rhizomes erect or horizontal, ovoid to cylindric; elongate stolons present or absent. Mature leaf blades mostly floating, elliptic-ovate to orbicular, entire to spinose-dentate, acute-tapered to truncate at apex, cordate to sagittate at base, with few to numerous trichosclereids and occasionally with needle-like sclereids embedded in mesophyll and forming tiny prominences on upper surface of dried leaves; venation mostly radiate but with a prominent midvein. Flowers with peduncles mostly stouter than petioles, especially when emergent. Sepals normally 4, hypogynous, often containing sclereids; petals 7–40, white (flora area species), blue, red, or yellow, in various shades, inserted in several series, hypogynous to perigynous, occasionally containing sclereids. Stamens numerous, multiseriate and appearing spiraled adaxially, perigynous to epigynous, often containing sclereids, the connectives with or without terminal appendage. Carpels 5–47, forming a ring embedded in cup-shaped receptacular and appendicular tissue, free or coherent laterally, the upper surface of each forming a ray of stigmatic tissue, the latter usually terminating abaxially in a free, upwardly incurved carpellary appendage, the appendage triangular, tapered, linear, or clavate; ovules numerous in each locule. Fruits irregularly dehiscent, berry-like or capsule-like, ripening under water. Seeds with a floating membranous aril, often with ridges or hair-like papillae on outer surface. Distributed in tropical to temperate habitats on all continents except Antarctica; ca. 45–50 species, 9 in Venezuela, 7 of these in the flora area. Most species of Venezuelan Nymphaea are night-blooming species which appear to be pollinated exclusively by scarab beetles of Cyclocephala Latreille. Key to the Species of Nymphaea 1.

1. 2(1).

2.

Mature leaf blades sinuate to strongly dentate, rarely entire; flowers usually elevated above water surface, opening diurnally or nocturnally; seeds ≥ 1.5 mm long ............................................................................... 2 Mature leaf blades entire; flowers floating at water surface, opening nocturnally; seeds always < 1.4 mm long ................................................... 3 Flowers opening diurnally, carpellary appendages short-triangular, outer connectives projecting as much as 3 mm beyond anthers; sepals with dark markings abaxially ................................................. N. pulchella Flowers opening nocturnally, carpellary appendages clavate, outer connectives projecting < 2 mm beyond anthers; sepals lacking dark markings abaxially .................................................................. N. rudgeana

120

3(1).

3.

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

5.

6(5).

6.

N YMPHAEACEAE

Sepals usually with purple or black flecks abaxially; leaves with ring of pubescence at apex of petiole; leaf blades and perianth lacking needlelike sclereids ......................................................................... N. amazonum Sepals uniformly green abaxially; leaves lacking ring of pubescence at apex of petiole; leaf blades and/or perianth containing numerous needle-like sclereids .............................................................................. 4 Mature leaf blades sagittate (later becoming elliptic), > 1.5 times as long as wide ................................................................................. N. potamophila Mature leaf blades elliptic-ovate to suborbicular, ≤ 1.5 times as long as wide ........................................................................................................ 5 Leaf veins lacking evident cross veins centrally, the venation radiate; stigmatic rays not containing needle-like sclereids; seed testa smooth (not appearing areolate in whole mount at 40×) ............... N. glandulifera Leaf veins with slightly to strongly evident cross veins centrally, forming web-like venation pattern; stigmatic rays with projecting needle-like sclereids; seed testa finely granulate (appearing areolate in whole mount at 40×) ......................................................................................... 6 Plants producing stolons only during germination of tubers; flowers autogamous, the fruits always maturing; lower surface of leaves uniformly green ........................................................................................... N. conardii Plants stoloniferous throughout growing season; flowers not autogamous, the fruits seldom maturing; lower surface of leaves usually mottled with rusty brown pigment .................................... N. gardneriana

Nymphaea amazonum Mart. & Zucc., Abh. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. 1: 363. 1832 [1830]. —Nymphaea blanda var. amazonum (Mart. & Zucc.) Planch., Fl. Serres Jard. Eur. 11: 21. 1856. —Nymphaea rudgeana var. amazonum (Mart. & Zucc.) Griseb., Fl. Brit. W. I. 12. 1864. —Leuconymphaea amazonum (Mart. & Zucc.) Kuntze, Revis. Gen. Pl. 1: 11. 1891. Floating-leaved aquatic herb. Rooted in rich organic substrate of slow-moving or stagnant, fresh or slightly brackish water, to 1.5 m deep, near sea level to 100 m; Delta Amacuro (Tucupita). Apure, Aragua, Guárico, Miranda, Monagas, Sucre; southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia, Brazil, Paraguay, Argentina. ◆Fig. 118. Nymphaea conardii Wiersema, Brittonia 36: 213. 1984. Floating-leaved aquatic herb. Rooted in mucky substrate of small freshwater streams, temporary ditches, or ponds to 1 m deep, near sea level to 200 m; northeastern Bolívar (Upata). Apure, Carabobo, Cojedes,

Portuguesa, Táchira, Zulia; southern Mexico, Central America, Greater Antilles, northern Colombia, Guyana(?), northeastern Brazil. ◆Fig. 117. Nymphaea gardneriana Planch., Fl. Serres Jard. Eur. 8: 120. 1852. Floating-leaved aquatic herb. Rooted in mucky substrate of stagnant or slow-moving fresh water to 1 m deep, 100–300 m; Bolívar (Guri, Río Suapure, Sabana Cardona). Apure, Barinas, Cojedes, Guárico, Portuguesa; Guyana, Brazil, eastern Bolivia, Paraguay, northeastern Argentina. ◆Fig. 121. Nymphaea glandulifera Rodschied, Med. Bem. Kol. Rio Essequibo 76. 1796 [1794]. —Nymphaea blanda G. Mey., Prim. Fl. Esseq. 201. 1818, nom. superfl. —Castalia blanda (G. Mey.) G. Lawson, Proc. & Trans. Roy. Soc. Canada 6: 117. 1888. —Leuconymphaea blanda (G. Mey.) Kuntze, Revis. Gen. Pl. 1: 11. 1891. Floating-leaved aquatic herb. Rooted in mucky substrate of stagnant or slow-moving water, near sea level to 300 m; Delta Amacuro (Güiniquina, Caño Pedernales,

Nymphaea 121

Serranía de Imataca), eastern Bolívar (El Foco near Upata, Río Uiri-yuk near Represa Guri, Río Urbani in lower Río Caura basin). Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, northern Peru, Bolivia, western Brazil. ◆Fig. 119. Nymphaea potamophila Wiersema, Sida 10: 195. 1984. Floating-leaved aquatic herb. Black-water swamps, along river margins, 50–200 m; Delta Amacuro (San Fernando de Atabapo), Bolívar (Los Garzones, Río Sipapo), Amazonas (Caño Yapacana, Raudal de Atures, Río Baría, Río Yaciba in Río Yatúa basin). Colombia(?), Guyana, Brazil. The relationship between Nymphaea potamophila and the related N. belophylla Trickett in interior South America remains to be clarified. Both sagittate-leaved species may be present in the region, as N.

belophylla, which differs in being stoloniferous and retaining sagittate leaves throughout the growing season, likely occurs in Barinas and eastern Colombia. Nymphaea pulchella DC., Syst. Nat. 2: 51. 1821. Nymphaea ampla auct. South America. Floating-leaved aquatic herb. Rooted in rich organic substrate of eutrophic and usually stagnant, often slightly brackish water, to 1.5 m deep, near sea level to 100 m; Delta Amacuro (between Tucupita and Caño Cocuina). Widespread elsewhere in Venezuela; western and southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 116. This species has generally been treated as Nymphaea ampla (Salisb.) DC., a distinct species that does not occur in South America.

Fig. 116. Nymphaea pulchella

122

N YMPHAEACEAE

Fig. 117. Nymphaea conardii

Fig. 118. Nymphaea amazonum

Fig. 119. Nymphaea glandulifera

Nymphaea 123

Fig. 120. Nymphaea rudgeana

Fig. 121. Nymphaea gardneriana

124

N YMPHAEACEAE

Nymphaea rudgeana G. Mey., Prim. Fl. Esseq. 198. 1818. Floating or occasionally submersed aquatic herb. Rooted in rich organic substrate in still- or slow-moving water, sometimes brackish, Mauritia palm swamps, near sea level to 900 m; Delta Amacuro

(widespread), Bolívar (northeast and San Ignacio de Yuruaní, Urimán). Anzoátegui, Miranda, Monagas, Zulia; Nicaragua, Cuba, Jamaica, Guadeloupe, Martinique, northeast Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 120.

OCHNACEAE by Claude Sastre Trees, treelets, shrubs, or herbs, generally glabrous. Leaves alternate, stipulate, rarely sessile, simple (compound in Rhytidanthera); margins generally toothed; secondary veins parallel at least in the proximal 1/2. Inflorescences generally terminal (axillary in Ouratea sect. Ouratella), generally compound with large panicles, rarely reduced to a solitary flower (Sauvagesia tenella); bracts caducous or persistent. Flowers actinomorphic or zygomorphic (Luxemburginae); pedicel articulate. Sepals generally 5(8–12 in Blastemanthus), often scarious, free, except in Perissocarpa and Ouratea sect. Kaieteurea; petals 4 or 5, free. Staminodes in 0–2 cycles, filamentous or petaloid; fertile stamens 5–many, free; filaments short (except Elvasia); anthers with longitudinal or poricidal dehiscence. Ovary superior, 2–12carpellate, the carpels fused (secondarily free in Ouratea and related genera); ovules 1–many, placentation parietal to axile, rarely basal; style single, gynobasic in Ouratea and related genera, stigmas 1–5, punctate to slightly lobed. Fruits various, capsular or indehiscent (dry or fleshy berries or drupes). Seeds albuminous (Sauvagesioideae) or exalbuminous (Ochnoideae). Pantropics; ca. 30 genera and ca. 400 species, 12 genera and 120 species in the flora area. Key to the Genera of Ochnaceae 1. 1. 2(1). 2. 3(2). 3. 4(1). 4. 5(4). 5. 6(5). 6.

Carpels 1-ovulate; fruit indehiscent, dry or separating into mericarps as fleshy berrylets or drupelets; seeds without endosperm ...................... 2 Carpels many-ovulate; fruit capsular; seeds with endosperm ................. 4 Leaves with secondary veins curved close the the margin; fruits with free mericarps on a swollen receptacle; style gynobasic ................... 6. Ouratea Leaves with secondary veins ± straight; carpels united in the fruit; style apical ...................................................................................................... 3 Flowers pedicellate; sepals free, petals yellow; stamens 5–25; style elongate or subulate; fruits sometimes ± lobed ................................. 5. Elvasia Flowers subsessile; sepals fused for most of their length; petals white; stamens 5; style conical; fruits not lobed ................................ 7. Perissocarpa Plants herbs to treelets; flowers actinomorphic, with petaloid staminodes ................................................................................................................ 5 Plants treelets to trees; androecium and gynoecium zygomorphic; petaloid staminodes absent .......................................................................... 8 Plants herbs to shrubs; staminodes without a keel; seeds not winged ....... 6 Plants shrubs to treelets; staminodes adaxially keeled; seeds winged ....... 7 Androgynophore present; stipule margins glandular; seeds muriculate ................................................................................................. 2. Adenarake Androgynophore absent; stipule margins with long awns; seeds alveolate

Adenarake 125

............................................................................................. 10. Sauvagesia Sepals ciliate-glandular; carpels 2 ............................................. 1. Adenanthe Sepals not ciliate-glandular; carpels 3 .......................................... 11. Tyleria Stipules persistent, > 20 mm long; carpels 5; staminodes lacking; capsules 5-valved ......................................................................... 4. Cespedesia 8. Stipules caducous or < 10 mm long; carpels 2 or 3; filamentous staminodes many or lacking; capsules 2- or 3-valved .......................... 9 9(8). Sepals 8–12; fertile stamens 10 ...........................................3. Blastemanthus 9. Sepals 5; fertile stamens 5 ....................................................................... 10 10(9). Petals white to pink; inflorescence axillary; capsules woody, globuliform, valves separating completely; seeds not winged .................... 12. Wallacea 10. Petals yellow; inflorescence terminal; capsule leathery, fusiform, valves remaining connate at the base; seeds winged .................................... 11 11(10). Stamens partly fused at base and congested on the adaxial side of the flower; anthers subsessile, staminodes absent; capsules opening from the base ...................................................................................... 8. Philacra 11. Stamens free, filaments conspicuously curving toward the adaxial side during anthesis; staminodes in 2 series; capsules opening from the apex ....................................................................................... 9. Poecilandra 7(5). 7. 8(4).

1. ADENANTHE Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 10(4): 19. 1961. Shrubs. Leaves sessile; stipules coriaceous, ovate-triangular 1–1.5 × 0.5–0.7 mm; blade coriaceous, oblong to lanceolate, 3–6 × 1.5–2.5 cm, the apex obtuse; margin with numerous small teeth near the apex; secondary veins thin, parallel, ascending at an 80° angle. Panicle terminal, 15–30 cm long; bracts persistent, ca. 2 mm long; pedicel < 1 cm long. Sepals 5, ovate to lanceolate, 8–13 × 3–5 mm, glandular; petals 5, obovate, 18–22 × 8–12 mm. Fertile stamens 5, subsessile, 5–6 mm high, poricidal; staminodes 10, 1-seriate in a 2 mm high corona, the 5 staminodes alternate with the fertile stamens narrowly spathulate, 8–9 mm high, the 5 opposite the stamens narrowly lanceolate, 9–10 mm high. Ovary 2-carpellate, 1-locular; ovules numerous. Capsule ovate, 2-valved, 8–10 mm long. Seeds winged, 3–3.5 mm long. Endemic to the Guayana Shield in southeastern Venezuela and western Guyana; 2 species, 1 of these in Venezuela. The second species in the genus, Adenanthe ciliata Sastre, is known from Mount Ayanganna, Guyana. Adenanthe bicarpellata Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 10(4): 19, fig. 27A, B. 1961. Shrub 0.3–4 m tall; margin of the leaf blade compactly ciliate at the middle and the

base; bracts glandular; petals white to pink. Savannas and rock outcrops on tepui summits, 1900–2300 m; Bolívar (Macizo del Chimantá). Guyana. ◆Fig. 122.

2. ADENARAKE Maguire & Wurdack, Mem. New York Bot. Gard. 10(4): 15. 1961. Shrubs. Leaves petiolate; stipules ± persistent, 1–5 mm long, linear or auriculate; blade membranaceous, elliptic, the apex acuminate; margin denticulate; secondary veins curved and parallel. Panicle 15–22 cm long; bracts persistent, glandular. Sepals 5; petals 5; androgynophore present, 1.5 mm long. Fertile stamens 5, subsessile; petaloid stamens fused at the base, in 2 series, the 10 outer scale-like, the 5 inner larger, contorted, enclosing completely the ovary and fertile stamens. Ovary 3-carpellate, 1-locular; ovules ca. 20. Capsule 3-valved. Seeds elliptic, muriculate, not winged.

126

O CHNACEAE

Fig. 122. Adenanthe bicarpellata

Fig. 123. Adenarake muriculata

Endemic to the Guayana Shield in southern Venezuela and northern Brazil; 2 species, both in the flora area. Key to the Species of Adenarake 1. 1.

Leaf blades 5–8 cm long, blade and sepal margin glandular-ciliate; capsule 12–13 mm long ........................................................ A. muriculata Leaf-blades 8–9 cm long, blade and sepal margin not glandular-ciliate; capsule 24–25 mm long ....................................................... A. macrocarpa

Adenarake macrocarpa Sastre, Novon 8: 301. 1998. Herbaceous shrub, woody at base, 1–3 m

tall. Granitic outcrops, ca. 1500 m; Amazonas (Serranía Sipapo-Guayapo 115 km southeast of Puerto Ayacucho). Endemic.

Blastemanthus 127

Adenarake muriculata Maguire & Wurdack, Mem. New York Bot. Gard. 10(4): 15, fig. 27C. 1961. Shrub 1–5 m tall; petals pink, bracts 2–5 mm long; outer staminodes 10, lanceolate, 3– 3.5 × 1 mm; inner staminodes 5, petaloid,

spathulate, 6–7 × 2 mm. Escarpments and scrub forests on tepui summits and slopes, 1500–2800 m; Amazonas (Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). ◆ Fig. 123.

3. BLASTEMANTHUS Planch., London J. Bot. 5: 644. 1846. Treelets or shrubs. Leaves petiolate, coriaceous, elliptic, the apex often emarginate and acuminate; secondary veins thin, numerous, and parallel; stipules ± deciduous. Panicles terminal. Flowers zygomorphic only at the androecium and gynoecium level. Sepals 8–12, unequal, the outer ones short and coriaceous, deciduous at anthesis, the inner ones longer and subcoriaceous; petals 5, equal, yellow. Staminodes many, thread-like, 4–5 mm long; fertile stamens 10, a few curved to the lower part of the flower; filaments short; anthers dehiscing by 2 pores. Ovary 3-carpellate, 3-locular, curved to the lower part of the flower as in the stamens; placentation parietal; ovules numerous. Capsule coriaceous, fusiform; style persistent. Seeds not winged, 1 or 2 per locule. Colombia, Venezuela, Guyana, Brazil; 2 species, 1 in Venezuela. Blastemanthus gemmiflorus (Mart.) Planch., London J. Bot. 5: 645. 1846. —Godoya gemmiflora Mart., Nov. Gen. Sp. Pl. 1: 118. t. 74. 1824 [1826]. Treelet or shrub 2–7 m tall; petiole 0.5–1.5 mm long; inflorescence 5–20 cm long; capsule 12–14 × 4–5 mm; seeds 5 × 0.5 mm. 100–200 m. Colombia, Venezuela, Guyana, Brazil; 2 subspecies, both in the flora area. Intermediates between the 2 subspecies occur along the Rio Negro in Amazonas, Brazil. Key to the Subspecies of B. gemmiflorus 1. Leaf blades 15–19 × 6–1.5 cm; secondary veins anastomosed ................................... ............................ subsp. gemmiflorus 2. Leaf blades 5–7 × 2–3 cm; secondary veins parallel ........................ subsp. sprucei B. gemmiflorus subsp. gemmiflorus Blastemanthus paniculatus Tiegh., Ann. Sci. Nat. Bot. sér. 8, 19: 68. 1904. Seasonally flooded forests along rivers and open areas. Amazonas (Río Pasimoni, near San Carlos de Río Negro). Colombia (Vaupés), Guyana, Brazil (Amazonas, Pará). B. gemmiflorus subsp. sprucei (Tiegh.) Sastre, comb. nov. —Blastemanthus sprucei Tiegh., Ann. Sci. Nat. Bot. sér. 8, 19: 68. 1904. —Palo de rábano. Blastemanthus densiflorus Hallier, Recueil Trav. Bot. Néerl. 10: 354. 1913.

Fig. 124. Blastemanthus gemmiflorus subsp. sprucei

128

O CHNACEAE

Riverbanks and savannas, on sandy soil. Amazonas (base of Cerro Yapacana, Río Atacabi, Río Baría, Río Caname,Río Casiquiare,

Río Guainía, Río Guayapo, Río Orinoco, Río Pasimoni, Río Temi). Colombia (Vaupés), Brazil (Amazonas: Rio Negro). ◆Fig. 124.

4. CESPEDESIA Goudot, Ann. Sci. Nat. Bot. sér. 3, 2: 368. 1844. Fournieria Tiegh., Ann. Sci. Nat. Bot. sér. 8, 19: 53. 1904, non Fourniera J. Bommer ex E. Fourn. 1873, nec Fourniera Scribn. 1897 (the latter two pteridophytes). Cespedezia Dwyer, Lloydia 9: 54. 1946. Shrubs or trees; trunks with buttresses at the base. Leaves petiolate; stipules coriaceous, sublinear to rectangular, 21–70 × 7–16 mm, ciliate at the base in the interior face; petiole 1–4 cm long; blade coriaceous, spatulate to ovate, 30–100 × 8–15 cm, the apex obtuse; margin toothed; secondary veins curved and parallel. Panicle large, terminal, 30–120 cm long; bracts caducous; pedicel 1–1.5 cm long. Flowers zygomorphic at the gynoecium level. Sepals 5, united at the base, obovate, 2–4 × 2–4 mm; petals 5, yellow, obovate, 1–2 × 0.8–1 cm. Stamens 60–80, articulate; filaments ca. 1 cm long; anthers dehiscent by 2 pores, 5–6 mm long. Ovary 5-carpellate, 1-locular, curved at the inferior part of the flower. Capsule coriaceous, fusiform, 6–7 × 1 cm, 5-valved. Seeds winged.

Fig. 125. Cespedesia spathulata

Elvasia 129

Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil (Mato Grosso); 1 species. Cespedesia spathulata (Ruiz & Pav.) Planch., London J. Bot. 5: 647. 1846. —Godoya spathulata Ruiz & Pav., Fl. Peruv. Prodr. 58, t. 11. 1794. —Aroiwaray-yek (Arekuna). Cespedesia bonplandii Goudot, Ann. Sci. Nat. Bot. sér. 3, 2: 370. 1844. Cespedesia sprucei Tiegh., Ann. Sci. Nat. Bot. sér. 8, 19: 51. 1904.

Tree 4–30 m tall; trunk diameter 0.5–0.7 m above the buttresses; flowers yellow. Forests, often disturbed areas, 100–1000 m; Delta Amacuro (Caño Cariabo in the Río Amacuro basin), Bolívar (Gran Sabana), Amazonas (Cerro Huachamacari, Cerro Sipapo, Cerro Yutajé, Río Cunucunuma). Sucre, Zulia; other distribution as in genus. ◆ Fig. 125.

5. ELVASIA DC., Ann. Mus. Natl. Hist. Nat. 17: 422. 1811. Hostmannia Planch. in Hook., Icon. Pl. 8: pl. 709. 1848. Trichovaselia Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 411. 1902. Vaselia Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 409. 1902. Tree or shrubs. Leaves subsessile to short-petiolate; stipules deltoid, united at the base; blades coriaceous, ± elliptic, acuminate; margin entire, slightly undulate with minute teeth; secondary veins thin, numerous and parallel. Inflorescence terminal or apparently axillary, 2–20 cm long; pedicels generally 5–10 mm long. Flower buds 3–4 mm long. Sepals 3–5, the outer ones coriaceous, the inner ones membranous; petals 3–5, membranous. Stamens 5–25; filaments 1–2.5 mm long; anthers ovate, 1–1.5 mm long, dehiscent by 2 subterminal pores. Ovary 2–6(7)-carpellate, the carpels fused, forming 2–6(7) locules; ovule 1 per locule. Fruit ligneous, globulose or forming a small star. Seeds 1, rarely 2. Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil; 9 species, 4 in Venezuela, all in the flora area. Key to the Species of Elvasia 1. 1. 2(1). 2. 3(2). 3.

Ovary 2-locular; filaments (at late anthesis) exceeding the anthers in length ..................................................................................... E. elvasioides Ovary 4- or 5-locular; filaments (at late anthesis) not exceeding the anthers in length ....................................................................................... 2 Stamens 20–25; stigma 1 ................................................... E. brevipedicellata Stamens 7–10; stigmas 4 or 5, fimbriate, radial ...................................... 3 Leaf blades cinereo-canescent on upper surface, 5–7 × 2–2.5 cm; fruits small and star-shaped ............................................................. E. canescens Leaf blades dull brown and glabrous on upper surface, 10–15 × 3–5 cm; fruits lobed .......................................................................... E. quinqueloba

Elvasia brevipedicellata Ule, Notizbl. Ann. Sci. Nat. Bot. sér. 8, 16: 411. Königl. Bot. Gart. Berlin 6: 339. 1915. 1902. Tree 5–15 m tall; leaf blades oblong, 6–8 × Small to medium-sized tree 2–10 m tall; 2.5–3.5 cm; fruit unknown. Tepui forests, fruit 6–7 mm diameter, 4- or 5-lobed; young 1800–1900 m; Bolívar (Cerro Jaua, Roraima- stem and peduncle shortly pubescent. Seatepui). Endemic. sonally flooded forests mostly along blackwater rivers, 100–200 m; Amazonas (Río Elvasia canescens (Tiegh.) Gilg in Engl. & Atabapo basin, Río Baría, Río Casiquiare baPrantl., Nat. Pflanzenfam. ed. 2, 21: 77. sin). Brazil (Amazonas: upper Rio Negro; 1925. —Trichovaselia canescens Tiegh., Mato Grosso: Rio Xingu). ◆Fig. 128.

130

O CHNACEAE

Elvasia elvasioides (Planch.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. III. 6: 145. 1893. —Hostmannia elvasioides Planch. in Hook., Icon. Pl. 8: pl. 709. 1845. —Laurel. Elvasia hostmannia Planch., London J. Bot. 5: 648. 1846. Hostmannia sagoti Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 414. 1902. —Elvasia sagoti (Tiegh.) Lemée, Fl. Guyane Fr. 3: 13. 1954. Elvasia caurensis Pittier, Bol. Soc. Venez. Ci. Nat. 6: 17. 1940. Tree 3–20 m tall, sometimes with buttresses; fruit 1.5–2 cm diameter. Edges of rivers, riparian forests, 100–400 m; Bolívar (Río Caroní, Río Caura, Río Suapure), Amazonas

(Puerto Ayacucho, Río Sipapo). Distrito Federal, Miranda; Colombia, Guyana, Suriname, French Guiana, Brazil (Pará). ◆Fig. 126. Elvasia quinqueloba Spruce ex Engl. in Mart., Fl. Bras. 12(2): 353. 1876. —Vaselia quinqueloba (Spruce ex Engl.) Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 409. 1902. Tree 3–6 m tall; fruit 1–1.5 cm diameter. River edges, 100–200 m; Amazonas (base of Cerro Yapacana, Río Manapiare, Río Temi). Colombia (Vaupés), Brazil (Amazonas: upper Río Negro). ◆Fig. 127.

Fig. 126. Elvasia elvasioides

Fig. 128. Elvasia canescens

Fig. 127. Elvasia quinqueloba

Ouratea 131

6. OURATEA Aubl., Hist. Pl. Guiane 397, t. 152. 1775. Trees, treelets, or shrubs. Leaves petiolate or rarely sessile, stipulate, coriaceous to membranaceous, elliptic to ovate or obovate; apex often acuminate; margin denticulate sometimes crenulate or toothed; secondary veins subequal, curved along margin or unequal, some arcuate and ascending along margin between smaller, fine, parallel, and straight veins; tertiary veins parallel and straight or anastomosed with the secondary veins; secondary and tertiary veins salient or impressed. Inflorescences terminal or axillary, paniculate or spicate, short (2–3 cm) to long (15 cm), bracteate or not, axes pubescent or not. Flowers actinomorphic. Sepals 2–5, coriaceous, united in 2 or 3 parts or free; petals 5, free, membranaceous, yellow; staminodes absent; stamens 10, sessile; anthers dehiscing by 2 pores. Ovary 5–10carpellate, carpels initially united, later free, uniovulate. Fruit in two parts (carpophore and carpels); carpophore (torus) fleshy, red; carpels 1–10, fleshy, 1–seeded, black at maturity. Neotropics; ca. 200 species, 72 in Venezuela, 67 of these in the flora area. Key to the Species of Ouratea 1. 1. 2(1). 2. 3(2).

Flowers with 2–4 sepals united into 2 or 3 parts ..................................... 2 Flowers with 5 distinct sepals ................................................................. 12 Leaves 12–22 × 4.5–6 cm ................................................................. O. engleri Leaves 2–10(–14) × 0.7–5.5 cm.................................................................. 3 Flowers with 6 or 7 carpels; sepals sometimes persistent in the old fruits ................................................................................................. O. articulata 3. Flowers with 5 carpels; sepals never persistent ....................................... 4 4(3). Leaves orbicular ......................................................................................... 5 4. Leaves not orbicular .................................................................................. 6 5(4). Fruiting pedicels 0.5 mm long, 2.2–3.5 × 1.8–2.5 cm; torus rhomboid, 5– 6 mm long ................................................................................... O. medinae 5. Fruiting pedicels ca. 1 cm long, 3–4.5 × 1.5–3 cm; torus subspheric, 4– 5 mm diameter .............................................................................. O. clarkii 6(4). Leaves subsessile, base cordate ....................................... O. brevipedicellata 6. Leaves petiolate, base not cordate ............................................................ 7 7(6). Leaves 2.5–3.5 × 0.7–1.5 cm ................................................... O. steyermarkii 7. Leaves 6–10(–14) × 2–5.5 cm .................................................................... 8 8(7). Inflorescence 2–5 cm long; leaves pruinose ................................. O. lajaensis 8. Inflorescence 5–15 cm long; leaves not pruinose ...................................... 9 9(8). Sepals 4–4.5 mm long .............................................................................. 10 9. Sepals 6–7 mm long ................................................................................. 11 10(9). Shrub 2–4 m tall; secondary veins of leaf blades subequal ...... O. brevicalyx 10. Tree 15–25 m tall; secondary veins of leaf blades unequal ....................... ......................................................................................... O. arbobrevicalyx 11(9). Shrub or treelet 1–4 m tall; leaves elliptic, base rounded ........ O. polyantha 11. Tree 4–15 m tall; leaves obovate, base rounded to cordate ...... O. thyrsoidea 12(1). Fruits with the carpels horizontal ............................................... O. oligantha 12. Fruits with the carpels vertical ............................................................... 13 13(12). Carpels 8–10; fruits with a plane torus ..................................... O. discophora 13. Carpels 5; fruits with a globular torus .................................................... 14

132

O CHNACEAE

14(13). 14. 15(14). 15. 16(15). 16. 17(15). 17. 18(17). 18. 19(17). 19. 20(19). 20. 21(19). 21. 22(21). 22. 23(22). 23. 24(14). 24. 25(24). 25. 26(25). 26. 27(26). 27. 28(26). 28. 29(28). 29. 30(29). 30. 31(30). 31. 32(25). 32. 33(32). 33. 34(32). 34. 35(34). 35. 36(35).

Sepals persistent in fruit ......................................................................... 15 Sepals caducous in fruit ........................................................................... 24 Margin of the leaf blades spinulose ......................................................... 16 Margin of the leaf blades not spinulose .................................................. 17 Principal vein salient on lower surface from base to the apex; leaf blades 11–26(–38) × 3.5–9(–12) cm ...................................................... O. angulata Principal vein scarcely salient only near the base; leaf blades 6–13 × 2– 4 cm ..................................................................................... O. castaneifolia Leaf blades with secondary veins unequal ............................................. 18 Leaf blades with secondary veins subequal ............................................ 19 Leaves 18–22 cm long, sepals 9–10 mm long ........................... O. guianensis Leaves 5–9 cm long, sepals 5–7 mm long ..........................O. marahuacensis Leaves 12–19 cm long .............................................................................. 20 Leaves 3–12 cm long ................................................................................ 21 Leaf blades oblong, veins on the lower surface impressed ................ O. tatei Leaf blades elliptic-ovate, veins not impressed .......................... O. paratatei Leaf margins clearly undulate and toothed .................................. O. superba Leaf margins few-toothed ........................................................................ 22 Leaf blades 6–12 × 3–5 cm ......................................................... O. spruceana Leaf blades 3–6 × 1.7–2.7 cm .................................................................. 23 Leaf blades subamplexicaul; sepals 5–6 mm long ......... O. subamplexicaulis Leaf blades ovate; sepals 10 mm long ........................................ O. longistyla Inflorescences axillary, rarely terminal .................................................. 25 Inflorescences always terminal ............................................................... 39 Inflorescences spicate, unbranched to few-branched ............................. 26 Inflorescences paniculate ........................................................................ 32 Inflorescences 11–30 cm long .................................................................. 27 Inflorescences 2–10 cm long .................................................................... 28 Leaves 10–15 × 4–6.5 cm ............................................................... O. croizatii Leaves 6–10 × 3–6 cm ................................................................ O. grosourdyi Margin of leaves revolute, few-toothed ..................................... O. yapacanae Margin of leaves flat, toothed .................................................................. 29 Leaves 20–60 cm long ....................................................................... O. ornata Leaves < 15 cm long ................................................................................. 30 Leaf blades 3–5 × 2–5 cm ................................................ O. guaiquinimensis Leaf blades 6–15 × 1.5–7 cm ................................................................... 31 Leaves membranaceous; inflorescence 8–10 cm long ............ O. soderstromii Leaves coriaceous; inflorescence 2–8 cm long .............................O. guriensis Leaf blade 1–2.5 times as long as wide ................................................... 33 Leaf blade > 2.5 times as long as wide .................................................... 34 Leaf blade 3–5.5 × 1.5–2.5 cm ......................................... O. pseudoguildingii Leaf blade 6–15 × 3.5–6 cm .................................................... O. ramosissima Axes of the inflorescence pilose ......................................... O. heterobracteata Axes of the inflorescence glabrous .......................................................... 35 Inflorescences multibracteate, multibranched; flowers very small (3– 4 mm long) ...................................................................... O. multibracteata Inflorescences not bracteate, few branched; flowers 6–8 cm long ......... 36 Leaf blades with the secondary and tertiary veins anastomosed .......... 37

Ouratea 133

36 Leaf blades with the secondary and tertiary veins not anastomosed ... 38 37(36). Leaf blades 5.5–8 × 2–3 cm; young stems without scales; inflorescence axes papillate ........................................................................... O. papillata 37. Leaf blades 10–14 × 4–6 cm; young stems with scales; inflorescence axes not papillate ............................................................................ O. squamata 38(36). Secondary veins subequal .......................................................... O. pisiformis 38. Secondary veins unequal ................................................................. O. duidae 39(24). Inflorescences spicate, unbranched to few branched ............................. 40 39. Inflorescences paniculate ........................................................................ 46 40(39). Axes of inflorescences ferruginous-pubescent .......................... O. ferruginea 40. Axes of inflorescences glabrous ............................................................... 41 41(40). Leaves > 8 cm long, apex acuminate ....................................................... 42 41. Leaves < 8 cm long, apex mucronate ...................................................... 44 42(41). Secondary veins unequal, margin not serrulate ............................ O. orisina 42. Secondary veins subequal, margin serrulate ......................................... 43 43(42). Inflorescences 4–6 cm long ...................................................... O. culminicola 43. Inflorescences 7–16 cm long .................................................... O. chaffanjonii 44(41). Inflorescences 5–11 cm long ............................................................. O. rigida 44. Inflorescences 3–4 cm long ...................................................................... 45 45(44). Leaf blade 1.7–2.8 × 0.6–0.9 cm ....................................................... O. huberi 45. Leaf blade 5.5–7 × 0.8–1.1 cm ..................................................... O. deminuta 46(39). Leaf blades > 12 cm long ......................................................................... 47 46. Leaf blades < 12 cm long ......................................................................... 54 47(46). Leaf blades > 4 times as long as wide ..................................... O. megaphylla 47. Leaf blades < 4 times as long as wide ..................................................... 48 48(47). Leaf blades < 2 times as long as wide ..................................................... 49 48. Leaf blades > 2 times as long as wide ..................................................... 50 49(48). Base of the leaf blade cordate; lateral veins salient on lower surface ............................................................................................... O. sipapoensis 49. Base of the leaf blade obtuse; lateral veins impressed on lower surface ..................................................................................................O. paruensis 50(48). Flower buds 4–5 mm long ........................................................................ 51 50. Flower buds 6–8 mm long ........................................................................ 52 51(50). Secondary veins of the leaves subequal ........................................ O. coccinea 51. Secondary veins of the leaves unequal ............................................ O. rorida 52(50). Secondary veins subequal .......................................................... O. fusiformis 52. Secondary veins unequal ......................................................................... 53 53(52). Leave blades oblong, base obtuse ................................................. O. maguirei 52. Leave blades elliptic, base acute ................................................... O. leblondi 54(46). Leaf blades < 2.8 times as long as wide .................................................. 55 54. Leaf blades > 2 times as long as wide ..................................................... 60 55(54). Leaf blades obovate ......................................................................... O. obovata 55. Leaf blades not obovate ........................................................................... 56 56(55). Leaf blades 3.5–6 cm long ........................................................................ 57 56. Leaf blades 6–12 cm long......................................................................... 58 57(56). Secondary and tertiary veins salient on both surfaces ................ O. davidsii 57. Secondary and tertiary veins not salient on both surfaces ........................ ......................................................................................... O. ptaritepuiensis

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O CHNACEAE

58(56). Leaf blade bases acute ................................................................. O. roraimae 58. Leaf blade bases cordate .......................................................................... 59 59(58). Leaf blade bases amplexicaul; secondary veins salient ............................. ..................................................................................... O. subamplexicaulis 59. Leaf blade bases not amplexicaul; secondary veins impressed on upper surface ................................................................................. O. maigualidae 60(54). Leaf blades narrowly ovate ............................................................. O. evoluta 60. Leaf blades not narrowly ovate ............................................................... 61 61(60). Leaf blades conduplicate ......................................................................... 62 61. Leaf blades not conduplicate ................................................................... 63 62(61). Flower buds 5–6 mm long; venation not anastomosed .............. O. attenuata 62. Flower buds ca. 10 mm long; venation anastomosed .......... O. rotundipetala 63(61). Axes of the inflorescences pilose ............................................................. 64 63. Axes of the inflorescences glabrous ......................................................... 65 64(63). Lateral veins salient on lower surface, sulcate on upper surface .............................................................................................. O. saldariagae 64. Lateral veins scarcely salient on lower surface, not sulcate on upper surface ...................................................................................... O. papillata 65(63). Flower buds 7–8 mm long ......................................................... O. grandiflora 65. Flower buds 5–6 mm long ........................................................................ 66 66(65). Leaf blades pulverulent on upper surface ............................. O. pulverulenta 66. Leaf blades not pulverulent ..................................................................... 67 67(66). Inflorescences with primary and secondary axes elongate, 6–8 cm long ................................................................................................... O. aquatica 67. Inflorescences with axes not elongate ..................................................... 68 68(67). Leaves shortly petiolate; petiole 3–4 mm long ................................ O. asisae 68. Leaves not shortly petiolate; petiole 6–8 mm long ................................. 69 69(68). Leaf blade lateral veins subequal .............................................. O. pisiformis 69. Leaf blade lateral veins unequal ............................................................. 70 70(69). Lateral veins salient on upper surface; buds 6–7 mm long .......... O. leblondi 70. Lateral veins not salient on upper surface; buds 4–5 mm long .... O. liesneri Ouratea angulata Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 259. 1902. —Sierrito. Tree 3–15 m tall; leaves coriaceous, 11– 26(–38) × 3.5–9(–12) cm; margin spinulose; secondary veins unequal; inflorescences terminal, paniculate, 13–25 cm long; torus subspherical, 7–8 mm diameter. Evergreen lowland to lower montane forests and gallery forests, 100–900 m; Delta Amacuro (Cano Orocoima), Bolívar (Ciudad Bolívar, Río Aro), Amazonas (Puerto Ayacucho, Río Ventuari). Zulia; Colombia (Vichada), Suriname, Brazil (Amapa, Amazonas, Goias, Matto Grosso, Minas Gerais, Pará). Ouratea aquatica (H.B.K.) Engl. in Mart., Fl. Bras. 12(2): 343. 1876. —Gomphia aquatica H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 14. 1823, non Maguire & Steyerm. 1989.

Ouratea pumila Maguire & Steyerm., Mem. New York Bot. Gard. 51: 86. 1989. Ouratea pumila var. heterodoxa Maguire & Steyerm., Mem. New York Bot. Gard. 51: 87. 1989. Shrub or treelet 0.5–4 m tall; leaves coriaceous, 2.5–9 × 1–3.5 cm; secondary veins unequal, variously prominent; inflorescence terminal, paniculate, few-branched, sometimes almost spicate, 6–10 cm long; torus obconic, 5–6 mm long. White-sand savannas, ca. 100 m; Amazonas (Caño Yagua, Cerro Yapacana base, Río Atabapo, Río Guayapo, Río Pasimoni, Yavita). Endemic. Ouratea arbobrevicalyx Sastre, Novon 11: 107. 2001. Tree 15–25 m tall; leaves chartaceous, 7– 14 × 3–4.2 cm; secondary veins unequal; in-

Ouratea 135

florescences terminal, paniculate, 10–13 cm long; fruit unknown. Along rivers, 100–200 m; Amazonas (Puerto Ayacucho, Río Casiquiare, Río Siapa). Endemic. Ouratea articulata Sastre, Phytologia 64: 247. 1988. Shrub 0.5–2 m tall; leaves coriaceous, 2.5– 4.5 × 1.5–2.5 cm; secondary veins unequal; inflorescences axillary, usually paniculate, 4–6 cm long; fruiting torus subspherical, 3–5 mm diameter, carpels 6–7 long. On quartzitic rocks, 1200–1400 m; Bolívar (Gran Sabana). Endemic. Ouratea asisae Maguire & Steyerm., Mem. New York Bot. Gard. 51: 62. 1989. Shrub 1–3 m tall; leaves coriaceous, 4–6.5 × 1.5–3.5 cm; secondary veins unequal; inflorescences terminal, paniculate, few-flowered, 2.5–5.5 cm long; fruit unknown. Understory of montane forests, ca. 1200; Amazonas (Cerro Asisa, Cerro Parú). Endemic. Ouratea attenuata Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 259. 1902. Treelet; leaves coriaceous, 9–10 × 3–4 cm; secondary veins subequal; inflorescence terminal paniculate, to 9 cm long; fruit unknown. Riparian forests, ca. 100 m; Amazonas (Río Casiquiare, San Carlos de Río Negro). Endemic. Ouratea brevicalyx Maguire & Steyerm., Mem. New York Bot. Gard. 51: 62. 1989. Shrub or treelet 2–5 m tall; leaves subchartaceous, 7–10 × 3–4.5 cm; secondary veins subequal; inflorescences terminal, paniculate, 11–15 cm long. Savannas, granitic and sandstone outcrops, 100–1000 m; Amazonas (Cerro Parú, Cerro Yutajé, Río Ventuari). Endemic. Ouratea brevipedicellata Maguire & Steyerm., Mem. New York Bot. Gard. 51: 64. 1989. Shrub or treelet 0.3–3 m tall; leaves coriaceous, 3.5–7 × 2.2–4 cm; secondary veins unequal; inflorescences terminal, sometimes on short stems, 4–10 cm long, few-branched; torus rhomboid, 4–5 mm high. Evergreen lowland forests and forest edges, 50–200 m; Amazonas (Río Guainía, Río Negro–Río Casiquiare junction). Endemic.

Ouratea castaneifolia (DC.) Engl. in Mart., Fl. Bras. 12(2): 309. 1876. —Gomphia castaneaefolia DC., Ann. Mus. Nat. Hist. 17: 417, pl. 11. 1811. —Ají de paloma. Shrub, treelet, or tree, 3–20 m tall; leaves coriaceous, 6–13 × 2–4 cm; secondary veins unequal; inflorescences terminal, paniculate, 8–21 cm long; torus turbinate, 8–9 mm high, 10–11 mm diameter. Forests, 100–800 m; Delta Amacuro, Bolívar (Ciudad Bolívar, Represa Raul Leoni, lower Río Caura), Amazonas (Raudal de Atures, Río Ventuari, Sierra Parima). Anzoátegui, Portuguesa, Táchira; Colombia (Amazonas, Antioquia, Vichada), Guyana, Suriname, Brazil (Amazonas, Maranhão, Pará, Roraima), Bolivia (La Paz). Ouratea chaffanjonii (Tiegh.) Sastre, Ernstia 56: 23. 1989. —Cercouratea chaffanjonii Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 272. 1902. Ouratea ayacuchae Maguire & Steyerark, Mem. New York Bot. Gard. 51: 62. 1989. Shrub 2–4 m tall; leaves chartaceous, 7– 16 × 3.5–6 cm; secondary veins subequal; inflorescences terminal, spicate, 7–16 cm long; torus conical, ca. 4 mm high. Granitic outcrops, ca. 100 m; Bolívar (Urbana), Amazonas (Puerto Ayacucho). Endemic. Ouratea clarkii Sastre, Bull. Mus. Hist. Nat. (Paris) sér. 4, 10, sect. B1: 51. 1988. —Ají de paloma. Tree 6–33 m tall; leaves coriaceous, 3–4.5 × 1.5–3 cm, base subcordate; secondary veins unequal; inflorescences terminal, paniculate, 5–10 cm long; torus subspherical, 4–5 mm diameter, mature carpels not seen. Evergreen lowland forests, white-sand scrub (bana), low- to medium-height forests on white sand, ca. 100 m; Amazonas (Río Casiquiare, San Carlos de Río Negro, road between Yavita and Maroa). Endemic. Ouratea coccinea Mart. ex Engl. in Mart., Fl. Bras. 12(2): 310, pl. 62. 1876. Tree 3–6 m tall; leaves coriaceous, 14– 17(–25) × 4.5–5.5(–9.5); secondary veins subequal, finely impressed; inflorescences terminal, paniculate, 9–13 cm long with 5 or 6 closely flowered axes, 5–13 cm long; torus globose, 3–4 mm diameter. Forests on white sand, ca. 100 m; Amazonas (Río Casiquiare,

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San Carlos de Río Negro). Peru (Loreto), Brazil (Amazonas, Pará). Ouratea croizatii Maguire & Steyerm., Mem. New York Bot. Gard. 51: 65. 1989. Shrub or treelet to 3 m tall, leaves coriaceous, 10–19 × 4–7.5 cm, base rounded; secondary veins unequal, tertiary veins anamastosed; inflorescence axillary at the top of 1- or 2-leaved stem, spicate or 1branched, 14–20 cm long; torus pyriform, straight or curved, mature carpel elongate, 12–15 × 5–7 mm. Along rivers, forests in rocky places, 100–200 m; Amazonas (Río Casiquiare). Endemic. Ouratea culminicola Maguire & Steyerm., Mem. New York Bot. Gard. 51: 65. 1989. Treelet 2–3 m tall; leaves coriaceous, 13– 18 × 4–7; secondary veins subequal; inflorescences terminal, spicate, 5–7 cm long; torus turbinate, 7 mm long. Low forests in rocky and sandy places, 700–1100 m; Bolívar (Amaruay tepui, Cerro Guaiquinima). Endemic. Ouratea davidsii Sastre, Phytologia 64: 365. 1988. Tree 10–25 m tall, branches pilose; leaves coriaceous, 1.5–3.5 × 1.5–3 cm; secondary veins subequal, anastomosed with the tertiary; inflorescences terminal, paniculate, 3– 9 cm long; sepals pilose; fruit unknown. Riparian forests, 100 m; Bolívar (Caicara). Anzoátegui, Apure (Río Arauca). Ouratea deminuta Maguire & Steyerm., Mem. New York Bot. Gard. 51: 66. 1989. Shrub 0.4–0.5 m tall; leaves coriaceous, 6.5–7.5 × 0.8–1.6 cm; secondary veins subequal faintly evident; inflorescences terminal, spicate, 7–8.5 cm long; torus conical elongate, 4–5 × 1–2 mm. White-sand savannas, 50–200 m; Amazonas (Cano Cumare, Minicio, San Fernando de Atabapo). Endemic. Ouratea discophora Ducke, Arq. Inst. Biol. Veg. 4(1): 53. 1918. Ouratea discophora subsp. pervenulosa Maguire & Steyerm., Mem. New York Bot. Gard. 51: 67. 1989. Medium-sized tree 4–12(–22 m in Brazil) m tall; leaves coriaceous, 8–19 × 4–7.5 cm;

secondary veins subequal; inflorescences terminal and axillary, paniculate, 4–10(–12 cm in Brazil) cm long; torus disciform, lobulate. Granitic outcrops, mixed montane forests, 100–600 m; Bolívar (Serranía de los Pijiguaos), Amazonas (Cerro Sipapo, near Puerto Ayacucho, road between Yavita and Maroa). Brazil (Acre, Amazonas, Mato Grosso, Rondônia). ◆Fig. 129. Ouratea duidae Steyerm., Fieldiana, Bot. 28: 369. 1952. Shrub 1–3 m tall; leaves coriaceous; secondary veins unequal; inflorescences axillary, at the top of 0–3-leaved stem, paniculate, few-branched, 1.5–9 cm long; fruit unknown. Upland savannas, 800–1200 m; Amazonas (Cerro Duida). Endemic. Ouratea engleri Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 259. 1902. Treelet 5 m tall; leaves coriaceous; secondary veins unequal; inflorescences terminal, 7–9 cm long, paniculate with 1–4 ramifications; fruits unknown. Lowland forests, ca. 100 m; Amazonas (Río Casiquiare). Brazil (Amazonas: Manaus-Caracarai road). Ouratea evoluta Maguire & Steyerm., Mem. New York Bot. Gard. 51: 67. 1989. Shrub or slender tree 1–3 m tall; leaves coriaceous, 7–15 × 3.5–4 cm; secondary veins unequal; inflorescences terminal, paniculate, loosely, 15–33 cm long; torus subglobose, 4–5 mm high, 5–8 mm wide. Riparian forests, ca. 100 m; Amazonas (base of Cerro Yapacana, Río Atabapo, near San Fernando de Atabapo). Endemic. Ouratea ferruginea Engl. in Mart., Fl. Bras. 12(2): 335, pl. 66. 1876. —Cercouratea ferruginea (Engl.) Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 276. 1902. Shrub or treelet 1–7 m tall; leaves membranaceous to chartaceous, oblong, 15–25 × 4–8 cm; secondary veins unequal; inflorescence terminal, racemiform, pendulous; torus turbinate, 10–15 mm long. Evergreen forests, 100–900 m; Bolívar (near Auyántepui, Cerro Guaiquinima, Río Caura, Santa Maria de Erebato), Amazonas (base of Cerro Duida, Cerro Huachamacari, Cerro Sipapo, Puerto Ayacucho, Río Atabapo, Río Pasimoni, San Carlos de Río Negro). Zulia; Colombia (Bolívar, Santander, Vaupés, Vichada), Bra-

Ouratea 137

zil (Amapa, Amazonas, Maranhão, Pará, Rondônia), Peru (Huánuco, Loreto). ◆ Fig. 134. Ouratea fusiformis Sastre, Novon 11: 113, fig. 13. 2001. Ouratea coccinea auct. non Engl. 1876: sensu Maguire & Steyerm., Mem. New York Bot. Gard. 51: 64. 1989, pro parte. Shrub or tree 2–15 m tall; leaves coriaceous, 23–29 × 7–9.5; secondary veins subequal; inflorescences terminal, paniculate, 10–25 cm long; torus conical, papillate, 10–11 mm high. Riparian forests, 100–200 m; Amazonas (Río Baría and adjacent Río Orinoco, Río Yatua). Endemic. Ouratea grandiflora (DC.) Engl. in Mart., Fl. Bras. 12(2): 336. 1876. —Gomphia grandiflora DC., Ann. Mus. Hist. Nat. 17: 420, pl. 17. 1811. Ouratea vasivae Spruce ex Engl. in Mart., Fl. Bras. 12(2): 336. 1876. Treelet 3–8 m tall; leaves subcoriaceous, 4–10 × 1–4 cm; secondary veins subequal; inflorescence terminal, shortly paniculate, 2.5– 6 cm long; torus turbinate, 9–12 mm high. Riparian forests, ca. 100 m; Amazonas (Río Atabapo, Río Casiquire, Río Guainía, Río Negro, Río Pasimoni). Brazil (Amazonas: Rio Negro). Ouratea grosourdyi (Tiegh.) Steyerm., Fieldiana, Bot. 28: 370. 1952. —Cercouratea grosourdyi Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 278. 1902. Ouratea pittieri Sleumer, Notizbl. Bot. Gart. Berlin-Dahlem 13: 351. 1936. Shrub or treelet 1–5 m tall; leaves coriaceous, 6–10 × 2–4.5 cm; secondary veins subequal; inflorescences axillary at the top of few-leaved stem, spicate, sometimes sparsely branched; torus pyriform, 3–5 mm long. Savanna and rocky formations, in thickets, semidecidous forests, 50–400 m; Bolívar (Caicara, Ciudad Bolívar, Ciudad Guyana, lower Río Caroní). Anzoátegui, Apure, Guárico, Miranda, Monagas. Ouratea guaiquinimensis Sastre, Phytologia 64: 365. 1988. Shrub 0.5–2 m tall; leaves coriaceous, 2–4 × 1.5–3 cm; secondary veins subequal; inflorescences axillary, spicate, few-flowered; torus subglobose, elongated, 9–10 mm long.

Tepui scrub, 1200–1800 m; Bolívar (Cerro Guaiquinima, Cerro Guanacoco). Endemic. Ouratea guianensis Aubl., Hist. Pl. Guiane 397, pl. 152. 1775. Ouratea longifolia auct. non (Lam.) Engl.: sensu Maguire & Steyerm., Mem. New York Bot. Gard. 51: 1989. Treelet or shrub 6–16 m tall; leaves coriaceous, 18–22(–33) × 4–8 cm; secondary veins unequal; inflorescences terminal, 1–25 cm long, paniculate. Lowland forests, 100–200 m; Delta Amacuro, Bolívar (Distrito Roscio). Guyana, Suriname, French Guiana, Brazil (Maranhão). Ouratea guriensis Sastre, Novon 5: 197. 1995. Shrub or treelet 1–4 m tall; leaves coriaceous, assymetric, 6–9.5 × 1.5–4 cm; secondary and tertiary veins anastomosed; inflorescences terminal or axillary, spicate, 2–8 cm long; torus discoidal, 3 × 8 mm. Semideciduous forests, 200–700 m; Bolívar (Cerro Toribio, El Pao, Represa Guri). Endemic. Ouratea heterobracteata Sastre, Novon 11: 108, fig. 5. 2001. Treelet 3 m tall, stems pilose; leaves membranaceous, 25–34 × 4.5–8 cm; secondary veins unequal; inflorescences axillary, paniculate, 12–16 cm long, axis pilose, bracteate at the base, axillary paired scale-like bracts on the secondary branches; fruit unknown. White-sand avannas, ca. 100 m; Amazonas (Río Guainía). Endemic. Ouratea huberi Maguire & Steyerm., Mem. New York Bot. Gard. 51: 74. 1989. Shrub 0.3–1 m tall; leaves coriaceous, ascending, 1.7–2.8 × 0.4–1 cm; secondary veins unequal; inflorescences terminal, spicate, few-flowered, 3–6 cm long; torus obconic, 5 mm long. White-sand savannas, ca. 100 m; Amazonas (Canaripó, Río Ventuari). Endemic. Ouratea lajaensis Sastre, Novon 11: 105, fig. 1. 2001. Treelet ca. 3 m tall; leaves coriaceous, 4.5– 10 × 2.5–4.5 cm, base obtuse or subcordate; secondary veins unequal; inflorescences terminal, paniculate, 3–4 cm; fruit unknown. Granitic outcrops, ca. 100 m; Amazonas (Puerto Ayacucho). Endemic.

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Ouratea leblondi (Tiegh.) Lemée, Fl. Guyane Fr. 3: 8. 1953. —Camptouratea leblondi Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 205. 1902. —Ajicito, Guayabito. Camptouratea sagotii Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 211. 1902. —Ouratea sagotii (Tiegh.) Lemée, Fl. Guyane Fr. 3: 9. 1953. Shrub or treelet 2–6 m tall; leaves coriaceous, 8–26 × 4–10.5 cm; secondary veins unequal; inflorescence terminal, paniculate, 6– 20(–26) cm long; torus subsphaeroidal, 4–9 mm high. Riparian forests and rocky formations, 50–600 m; Delta Amacuro (Río Grande), Bolívar (Canaima, El Dorado, Serranía de Imataca). Guyana, Suriname, French Guiana, Brazil (Amapa, Maranhão). ◆Fig. 133. Ouratea liesneri Sastre, Phytologia 64: 366. 1988, “liesnerii.” Shrub 3–4 m tall, stem hairy; leaves subcoriaceous, velvety on lower surface, 6–10 × 2–5 cm, lateral veins unequal; inflorescence terminal, paniculate, hairy, 6–10 cm long; mature fruits unknown. On rock outcrops, 50–200 m; Amazonas (Río Chimoni, Río Mamurividi on middle Río Pasimoni). Endemic. Ouratea longistyla Maguire & Steyerm., Mem. New York Bot. Gard. 51: 78. 1989. Shrub to 3 m; leaves coriaceous, 4–6.5 × 1.5–3.5 cm; secondary veins subequal; inflorescence terminal, paniculate, 8–10 cm long; mature fruit unknown. Sandy lowland savannas, ca. 100 m; Amazonas (lower Río Siapa). Endemic. Ouratea maguirei Sastre, Novon 11: 115, fig. 15. 2001. Ouratea coccinea auct. non Engl. 1876: senus Maguire & Steyerm., Mem. New York Bot. Gard. 51: 64. 1989, pro parte. Treelet ca. 5 m tall; leaves coriaceous, (13–) 22–27 × 6–8.5 cm, undulate, margin denticulate; secondary veins unequal; inflorescences terminal, paniculate, 7–22 cm long, axis papillate; fruits unknown. Riparian forests, ca. 100 m; Amazonas (San Fernando de Atabapo). Endemic. Ouratea maigualidae Sastre, Novon 11: 117, fig. 18. 2001. Shrub or treelet 1.5–4 m tall; leaves coriaceous, 6–9 × 4–7 cm; secondary veins un-

equal, anastomosed with the tertiary; inflorescence terminal, paniculate, 8–15 cm long. Low forests on granitic outcrops, 1700– 2000 m; Amazonas (Sierra de Maigualida). Endemic. Ouratea marahuacensis Maguire & Steyerm., Mem. New York Bot. Gard. 51: 80. 1989. Treelet ca. 7 m tall; leaves subcoriaceous, 5–9 × 2.5–4 cm; secondary veins subequal; inflorescences terminal, paniculate, 6–8 cm long. Montane forests along rivers, 1100– 1200 m; Amazonas (Cerro Marahuaka). Endemic. Ouratea medinae Maguire & Steyerm., Mem. New York Bot. Gard. 51: 81. 1989. Tree; leaves coriaceous, 2.2–3.5 × 1.8–2.5 cm; secondary veins unequal; inflorescences terminal, paniculate, 4–6 cm long; flowers unknown. Along rivers, ca. 100 m; Amazonas (Río Atabapo, San Fernando de Atabapo). Endemic. Ouratea megaphylla Sastre, Novon 11: 113, fig. 12. 2001. Treelet 5–10 m tall; leaves coriaceous, 35– 45 × 8–12.5 cm; secondary veins subequal; inflorescence terminal, paniculate, 23–30 cm long; torus globose, 7 mm diameter. Lowland evergreen forests, ca. 100 m; Amazonas (Puerto Ayacucho, Río Cataniapo). Endemic. Ouratea multibracteata Steyerm. ex Sastre, Novon 11: 108, fig. 4. 2001. Tree 6–8 m tall; leaves chartaceous, 10–15 × 4–6 cm; secondary veins unequal; inflorescence axillary, paniculate, 15–20 cm long, with many persistent, strongly appressed bracts; torus disciform, 10-lobed. Along rivers, ca. 100 m; Amazonas (Río Autana). Endemic. Ouratea obovata Sastre, Novon 11: 111, fig. 11. 2001. Shrub 3 m tall; leaves coriaceous, 7–12 × 4.5–6 cm; secondary veins subequal; inflorescences terminal, paniculate, 4–6 cm long; flowers unknown; torus conical 13–14 mm high. Along streams, 1500–1700 m; Amazonas (Cerro Sipapo). Endemic. Ouratea oligantha Steyerm. ex Sastre, Novon 5: 194. 1995.

Ouratea 139

Shrub 1.5–4 m tall; leaves coriaceous, 7–9 × 3–3.5 cm; secondary veins unequal; inflorescences usually axillary and spicate, 4–10 cm long; carpels horizontal; mature fruits unknown. Forests on sandstone, ca. 400 m; Bolívar (upper Río Cuyuní). Endemic. Ouratea orisina Sastre, Novon 11: 111, fig. 8. 2001. Tree; leaves membranaceous, 10.5–17 × 3–5.5 cm; secondary veins unequal; inflorescence terminal, spiciform, 8 cm long; flowers unknown; torus subspheric, 5 mm diameter. Riparian forests, lowland slope forests, 200– 300 m; Bolívar (Río Oris in upper Río Paragua basin). Endemic. Ouratea ornata Maguire & Steyerm., Mem. New York Bot. Gard. 51: 82. 1989. Shrub ca. 1.5 m tall, stems ferruginousverruculose; leaves coriaceous, 15–30 × 4–7 cm; secondary veins unequal; inflorescences axillary at the top of 1-leaved stem, spicate 4–4.5 cm long, on stems 9 cm long; flowers unknown; torus subglobulose, ca. 4 × 8 mm. Evergreen lowland to mixed montane forests, 100–900 m; Amazonas (Cerro Sipapo, Río Cataniapo). Endemic. Ouratea papillata Maguire & Steyerm., Mem. New York Bot. Gard. 51: 82. 1989. Shrub 0.5–2 m tall; leaves coriaceous, 5.5– 8 × 1.8–2.5 cm; secondary veins unequal, not apparent on lower surface; inflorescences terminal and axillary, paniculate, 5–18 cm long; torus turbinate to subglobulose, 2–6 mm high. White-sand savannas, 50–200 m; Amazonas (base of Cerro Yapacana, Río Pasimoni, Río Siapa, Río Yatua). Endemic.

long; fruits unknown. Montane meadows, scrub forests, 1600–2000 m; Amazonas (Cerro Parú, Serranía Uasadi). Endemic. Ouratea pisiformis Engl. in Mart., Fl. Bras. 12(2): 343. 1876. Ouratea crenata Spruce ex Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 259. 1902. Tree 2–12 m tall; leaves membranaceous, 8–12 × 2.5–4 cm; secondary veins subequal; inflorescence terminal and axillary, paniculate, 7–18 cm long; torus pyriform to conical, 3–4 mm high. Riparian forests, 100–400 m; Bolívar (Río Paragua), Amazonas (Puerto Ayacucho, Río Atabapo, Río Emoni in lower Río Siapa basin, Río Guainía, Río Negro, Río Sipapo, San Carlos de Río Negro). Apure, Táchira; Guyana, Colombia (Guainía), Brazil (Amazonas: Rio Negro). Ouratea polyantha (Triana & Planch.) Engl. in Mart., Fl. Bras. 12(2): 317. 1876. —Gomphia polyantha Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 18: 274. 1862. Tree, treelet, or shrub, 1–10 m tall; leaves chartaceous, 7–15 × 3–5.5 cm; secondary veins unequal; inflorescences terminal, paniculate, 9–19 cm long; torus spheroid, 4 mm high, shorter than the mature carpels, 8 mm long. Riparian forests, shrub savannas, 100– 200 m; Bolívar (Caicara, Río Parguaza), Amazonas (base of Cerro Yapacana, Puerto Ayacucho, Río Manapiare, Río Orinoco, Río Sipapo). Apure; Colombia (Meta, Vaupés, Vichada). ◆Fig. 130.

Ouratea paratatei Sastre, Novon 5: 196. 1995. Tree ca. 7 m tall; leaves coriaceous, 14–19 × 4.5–7 cm; secondary veins subequal; inflorescences terminal, paniculate, ca. 12 cm long; flowers unknown; torus conical, 5–6 mm high. Semideciduous forests, ca. 100 m; Bolívar (north of Upata). Endemic.

Ouratea pseudoguildingii Sastre, Novon 5: 197. 1995. Shrub or slender tree 2–5 m tall; leaves coriaceous, 3–5.5 × 1.5–2.5 cm; secondary veins subequal, tertiary veins anastomosed; inflorescences paniculate, 5–10 cm long at the top of short stem with 1 or 2 leaves at base; torus pyriform 5 mm long. Wet places in savannas, palm swamps, and along rivers, 100–300 m; Bolívar (Cerro Carichana, Cerro Gavilán, Ciudad Bolívar, Puerto Ordaz, Río Horeda near Ciudad Bolívar). Endemic.

Ouratea paruensis Maguire & Steyerm., Mem. New York Bot. Gard. 51: 83. 1989. Shrub 1.5–4 m tall; leaves coriaceous, 13– 16 × 4–7 cm; secondary veins unequal, anastomosed with the tertiary veins impressed; inflorescences terminal, paniculate, 7–14 cm

Ouratea ptaritepuiensis Steyerm., Fieldiana, Bot. 28: 370. 1952. Ouratea propingua Maguire & Steyerm., Mem. New York Bot. Gard. 51: 85. 1989. Shrub 0.5–2.5 m tall; leaves coriaceous, 2.5–6(–8) × 1.5–3.5(–4) cm; secondary veins

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subequal; inflorescence terminal, paniculate, 3–6 cm long; torus obconic, 6–8 mm long. Elfin forests, rocky savannas, 1500–2000 m; Bolívar (Cerro Guanacoco, Cerro Jaua, Macizo del Chimantá, Ptari-tepui). Endemic. Ouratea pulverulenta Sastre, Novon 11: 111, fig. 10. 2001. Tree ca. 5 m tall; leaves coriaceous, powdered on upper surface, pruinose on lower surface, 6–9 × 2–3 cm; secondary veins subequal; inflorescences terminal, paniculate, 4–6 cm long; flowers unknown; torus subcylindric, mamillate, 2 mm high, 3 mm diameter. Riparian lagoons, 100–200 m; Amazonas (Río Ocamo). Endemic. Ouratea ramosissima Maguire & Steyerm., Mem. New York Bot. Gard. 51: 87. 1989. Shrub or treelet 0.3–7 m tall; leaves coriaceous, 6–18(–20) × 5–10.5 cm; secondary veins unequal; inflorescence axillary, paniculate, 7–19 cm long; torus elongate-turbinate, 5–7 mm long. On rocky quartzitic slopes, 400–1600 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Cerro Ichún, Río Caroní, Río Paragua), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Marahuaka, Cerro Yapacana, Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina, Serra Aracá). ◆ Fig. 131. Ouratea rigida Engl. in Mart., Fl. Bras. 12(2): 307. 1876. Villouratea spiciformis Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 240. 1902. Small tree 2–5 m tall, stems villous; leaves coriaceous, 3–6 × 1.5–3 cm; secondary veins subequal, anastomosed with the tertiary veins; inflorescence terminal, spicate, 5–12 cm long; torus turbinate, 8–9 mm diameter, 4–6 mm high. Semi-evergreen forests on soils with iron, ca. 700 m; Bolívar (Cerro Toribio). Guyana. Ouratea roraimae Engl. in Mart., Fl. Bras. 12(2): 308. 1876. —Copeycito. Shrub or treelet 1–6 m tall; leaves coriaceous, 6–14 × 3.5–8.5 cm; secondary veins subequal; inflorescence terminal, paniculate, 7–25 cm long; torus conical, > 1 cm long. Low savanna-forests, 100–900 m; Bolívar (Cerro Bolívar, Cerro Guaiquinima, Gran Sabana, Guri, upper Río Caroní, Río Paragua),

Amazonas (Cerro Yapacana, Río Siapa, Río Ventuari, Sierra Unturán). Guyana, Brazil (Amazonas, Pará, Roraima). ◆Fig. 137. Ouratea rorida Sastre, Novon 11: 115, fig. 16. 2001. Ouratea guianensis auct. non Aubl. 1775: sensu Maguire & Steyerm., Mem. New York Bot. Gard. 51: 73. 1989, pro parte. Tree 3–15 m tall; leaves membranous, 18– 23 × 5.5–8.5 cm, epidermis minutely tuberculate; secondary veins unequal; inflorescences terminal, paniculate, 13 cm long; torus globular, 3–4 mm diameter. Riparian forests, 200–600 m; Bolívar (Río Caura, middle Río Paragua, Sierra de Lema), Amazonas (Cerro Yutajé). Endemic. Ouratea rotundipetala Maguire & Steyerm., Mem. New York Bot. Gard. 51: 90. 1989. Treelet 2–5 m tall; leaves coriaceous, 7–10 × 2.5–4 cm; secondary veins subequal, reticulate with the tertiary; inflorescence terminal, paniculate, few-flowered, 1–2.5 cm long; mature fruit unknown. Riparian forests, 100– 200 m; Amazonas (base of the Cerro Yapacana, Río Casiquiare). Endemic. Ouratea saldariagae Sastre, Novon 11: 111, fig. 9. 2001. Tree 12 m tall; leaves chartaceous, 5.5–9.5 × 1.7–4 cm; secondary veins unequal; inflorescence terminal, paniculate or spicate, axes papillate, few-flowered, 2.5–3 cm long; flower unknown; torus subcylindric, 5–8 mm high, 9–11 mm diameter. Evergreen lowland forests, 100–200 m; Amazonas (Brazo Casiquire, near San Carlos de Río Negro). Endemic. Ouratea sipapoensis Maguire & Steyerm., Mem. New York Bot. Gard. 51: 92. 1989. Shrub 2–4 m tall; leaves coriaceous, 10–20 × 6–10 cm; secondary veins unequal; inflorescence terminal and axillary, paniculate, 10–25 cm long; torus turbinate, 7–10 mm long. Montane forests, 1400–1800 m; Amazonas (Cerro Sipapo). Endemic. ◆Fig. 136. Ouratea soderstromii Sastre, Phytologia 64: 248. 1988. Treelet or tree 4–20 m tall; leaves membranous, 6–15 × 2.5–5 cm; secondary veins unequal; inflorescence axillary, spicate, pen-

Ouratea 141

dulous, 6–8 cm long; torus subconic, 7–9 mm long. Swamp and riparian forests, ca. 400 (550–900 in Guyana) m; Bolívar (Río Caroní). Guyana. Ouratea spruceana Engl. in Mart., Fl. Bras. 12(2): 337. 1876. Treelet or shrub 1–10 m tall; leaves coriaceous, 6–12(–16) × 3–5(–6) cm; secondary veins subequal; inflorescences terminal, paniculate, 4–15 cm long; fruit included in the sepals. Savannas on white sand and quartzitic outcrops, 100–400 m; Bolívar (mouth of Río Paragua), Amazonas (Cerro Yapacana, Río Casiquiare, Río Siapa, Río Sipapo, San Carlos de Río Negro, San Fernando de Atabapo). Colombia (Amazonas, Vaupés), Guyana, Brazil (Amazonas, Pará). ◆ Fig. 132. Ouratea squamata Sastre, Novon 5: 199. 1995. Tree ca. 10 m tall; leaves coriaceous, 10– 14 × 4–6 cm; secondary veins subparallel; inflorescences axillary, paniculate, 16–26 cm long, axes with scales; flowers unknown; torus conical, 5–6 × ca. 5 mm. On quartzitic outcrops, ca. 1200 m; Bolívar (Cerro Marutaní). Endemic. Ouratea steyermarkii Sastre, Phytologia 64: 247. 1988. Ouratea macurucoensis Maguire & Steyerm., Mem New York Bot. Gard. 51: 80. 1989. Shrub 0.1–3 m tall; leaves coriaceous, 3–5 × 1–2.2 cm; secondary veins unequal; inflorescences terminal on short stems; torus elongate, 7–10 mm long, carpels ellipsoid, 6.5–7 × 4–5 mm. White-sand savannas, 100– 200 m; Amazonas (base of Cerro Yacapana, upper Río Orinoco, Río Ventuari, San Fernando de Atabapo). Endemic. Ouratea subamplexicaulis Maguire & Steyerm., Mem. New York Bot. Gard. 51: 96. 1989. Shrub or treelet 1–5 m tall; leaves subcoriaceous, 7–15 × 2–5 cm, serrulate; secondary veins subequal; inflorescence terminal, paniculate, few-branched, 5–10 cm long. Open formations on rocks, 400–500 m; Bolívar (base of Cerro Venamo). Endemic. Ouratea superba Engl. in Mart., Fl. Bras. 12(2): 334. 1876.

Ouratea inundata Spruce ex Engl. in Mart., Fl. Bras. 12(2): 335. 1876. Ouratea odora Poepp. ex Engl. in Mart., Fl. Bras. 12(2): 334. 1876. Cercouratea laxa Tiegh., Ann. Sci. Nat. Bot. sér. 8, 16: 273. 1902. Ouratea inconformis Maguire & Steyerm., Mem. New York Bot. Gard. 51: 75. 1989. Ouratea wurdackiana Maguire & Steyerm., Mem. New York Bot. Gard. 51: 100. 1989. Shrub or treelet 2–10 m tall; leaves chartaceous, 5–12 × 2–5 cm; secondary veins subequal; inflorescences terminal and axillary on short stem, paniculate, 5–10 cm long; fruiting torus sphaeroidal, 5–7 mm high. Riversides, 50–400 m; Bolívar (Cerro Guaiquinima, Río Caroní, upper Río Paragua), Amazonas (Río Atacavi, Río Sipapo, Río Ventuari, San Fernando de Atabapo). Apure, Guárico; Colombia (Amazonas, Putumayo), Guyana, Suriname, Peru (Loreto), Brazil (Amapá, Amazonas, Mato Grosso do Norte, Pará, Rondônia). Ouratea tatei Gleason, Bull. Torrey Bot. Club 56: 398. 1929. Tree 4–10 m tall; leaves coriaceous, (9–) 12–18 × 3–6.5 cm; secondary veins subequal; inflorescences terminal, paniculate, 14–21 cm long; fruiting torus rhomboid, 8–10 cm high. Sandstone areas, 1000–1700 m; Bolívar (Gran Sabana, Macizo del Chimantá, Roraima-tepui). Guyana (Mount Roraima). Ouratea thyrsoidea Engl. in Mart., Fl. Bras. 12(2): 313. 1876. Ouratea venezuelensis Steyerm., Fieldiana, Bot. 28: 370. 1952. Tree 4–15 m tall; leaves chartaceous, 6–13 × 2.5–5 cm; secondary veins subequal; inflorescences terminal, paniculate, 14–21 cm long; fruiting torus sphaeroidal, 5–6 mm long. Riversides, 50–200 m; Amazonas (Río Casiquiare, Río Guainía, Río Orinoco, Río Ventuari, Río Yatua). Brazil (Amazonas, Pará). Ouratea yapacanae Sastre, Phytologia 64: 248. 1988. Few-branched shrub 0.2–1 m tall; leaves rigid coriaceous, 9–13 × 2–3 cm; secondary veins subequal; inflorescence axillary, on short and thick stems without leaves, fewflowered; torus subspherical, 7–8 mm diameter. Sandy savannas, 100–200 m; Amazonas (Cerro Yapacana base). Endemic. ◆Fig. 135.

142

O CHNACEAE

Fig. 129. Ouratea discophora

Fig. 130. Ouratea polyantha

Fig. 131. Ouratea ramosissima

Ouratea 143

Fig. 132. Ouratea spruceana

Fig. 133. Ouratea leblondi

Fig. 134. Ouratea ferruginea

144

O CHNACEAE

Fig. 135. Ouratea yapacanae

Fig. 136. Ouratea sipapoensis

Fig. 137. Ouratea roraimae

Perissocarpa 145

Fig. 138. Perissocarpa steyermarkii

Fig. 139. Perissocarpa umbellifera

7. PERISSOCARPA Steyerm. & Maguire, Ann. Missouri Bot. Gard. 71: 319. 1984. Shrubs or trees to 15 m tall. Leaves alternate, sometime pseudo-opposite or pseudoverticillate, petiolate; stipules coriaceous 1–1.5 mm long; blades coriaceous, oblong to elliptic, 2.5–18 × 1.8–10 cm; margins with small black teeth; secondary veins thin and straight, parallel. Inflorescence terminal or subterminal, paniculately or umbellately branched, axis densely flowered, 2–15 cm long; bracts triangular, 1–2 mm long; pedicels ca. 1 mm long. Flowers actinomorphic. Sepals 5, free or fused for most of their length, splitting into 2 or 3 segments, suborbicular, often reflexed, 1.5–4 mm long; petals 5, white, suborbicular, longer than the sepals. Stamens

146

O CHNACEAE

5; filaments short, 0.3–0.5 mm long; anthers ovate, 1.5–1.8 mm long, opening by 2 apical pores. Ovary of 2 or 3 united carpels, 2- or 3-locular; ovules 1 per locule; style conical, 0.5–0.8 mm long. Fruit subglobose-pyriform, leathery. Seed 1. Colombia, Venezuela, Peru, Brazil; 3 species, 2 in Venezuela, both in the flora area. Key to the Species of Perissocarpa 1.

1.

Inflorescences pedunculate, the branches alternately arranged along the main axis, calyx lobes > 0.6 mm long; petiole > 10 mm long .............................................................................................. P. steyermarkii Inflorescences sessile, umbellately branched; calyx lobes < 0.2 mm long; petiole 1–6 mm long ...............................................................P. umbellifera

Perissocarpa steyermarkii (Maguire) Steyerm. & Maguire, Ann. Missouri Bot. Gard. 71: 319. 1984. —Elvasia steyermarkii Maguire, Acta Bot. Venez. 3: 297. 1968. Small or medium-sized tree to 15 m tall; calyx lobes evident. Montane forests, 1500– 1900 m; Bolívar (Cerro Jaua), Amazonas (Cerro Duida, Sierra de Maigualida). Aragua, Monagas, Táchira; Brazil (Amazonas). ◆Fig. 138.

Perissocarpa umbellifera Steyerm. & Maguire, Ann. Missouri Bot. Gard. 71: 320. 1984. Shrub to small tree 1.5–10 m tall; leaf blades 2.5–20 × 1.8–10 cm; apex of the petals subtruncate. Dwarf forests along streams and in rocky areas on tepui slopes and summits, 800–2000 m; Bolívar (Cerro Sarisariñama), Amazonas (Cerro Duida, Cerro Sipapo). Colombia (Amazonas), Brazil (Amazonas: Serra Aracá). ◆ Fig. 139.

8. PHILACRA Dwyer, Brittonia 5: 124. 1944. Treelets or shrubs 2–8 m tall. Leaves sessile, coriaceous; blades elliptic, obovate, or sometimes falciform, 6–22 cm long, the apex acute, base attenuate or auriculate; margins serrate with small teeth 3–4 mm long; secondary veins parallel, ascending. Inflorescence terminal, paniculately branched or not; bracts minute, caducous; pedicel articulate. Flowers zygomorphic at the androecium and gynoecium level. Sepals 5, unequal, the outer smaller, 4–6 mm long, coriaceous, deciduous at anthesis, subrotund to oblong; petals 5, yellow, equal, obovate-oblong, ca. 18 × 10– 12 mm, the apex round-obtuse. Stamens (4–10)20–25, subsessile, incompletely cyclic, aggregated into a mass that incompletely surrounds the pistil, arranged in three ranks, the inner ones longer than the outer ones, 6–7 mm long, dehiscing by subterminal pores. Pistil arcuate, opposite the stamens; ovary 3-angled, fusiform, 7 mm long; style rostrate. Capsule thin-coriaceous, subfalcate and fusiform, dehiscing from the base, remaining attached at the apex, 1–2 cm long. Seeds smooth, terete, 1.2–1.5 mm long, the wing 0.5–0.7 mm long at each end. Endemic to the Guayana Shield in southern Venezuela and northern Brazil; 4 species, all in the flora area. Key to the Species of Philacra 1. 1. 2(1). 2.

Flowers few (5–15) on a short inflorescence ca. 1 cm long; pedicels 1–4 mm long ................................................................................................. P.duidae Flowers numerous (> 30) on a large inflorescence 10–17 cm long ........... 2 Inflorescence branched ............................................................ P. steyermarkii Inflorescence not branched ........................................................................ 3

Philacra 147

Fig. 140. Philacra duidae

Fig. 141. Philacra auriculata

Fig. 142. Philacra steyermarkii

148

3(2). 3.

O CHNACEAE

Leaf blades auriculate at base .................................................... P. auriculata Leaf blades attenuate at base ....................................................... P. longifolia

Philacra auriculata Dwyer, Brittonia 5: 125. 1944. Tree or shrub 2–8 m tall. Upper montane woodland and escarpments, 800–1700 m; Amazonas (Sierra de la Neblina). Brazil (Amazonas: Serra Imeri). ◆ Fig. 141. Philacra duidae (Gleason) Dwyer, Brittonia 5: 125. 1944. —Luxemburgia duidae Gleason, Bull. Torrey Bot. Club 58: 389. 1931. Shrub ca. 2 m tall. Tepui summits, ca. 1600 m; Amazonas (Cerro Duida). Endemic. ◆ Fig. 140.

Philacra longifolia (Gleason) Dwyer, Brittonia 5: 127. 1944. —Luxemburgia longifolia Gleason, Bull. Torrey Bot. Club 58: 390. 1931. Shrub or tree 3–6 m tall. Bluffs and escarpments, 200–2000 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Parú, Cerro Sipapo, Río Cunucunuma). Brazil (Amazonas: Serra Aracá). Philacra steyermarkii Maguire, Acta Bot. Venez. 2(5–8): 241. 1967. Shrub ca. 2.5 m tall. Dwarf forests along streams, 1600–1800 m; Bolívar (Auyántepui). Endemic. ◆ Fig. 142.

9. POECILANDRA Tul., Ann. Sci. Nat. Bot. sér. 3, 8: 342. 1847. Treelets or shrubs, rarely subshrubs, bark black. Leaves subsessile, coriaceous; secondary veins thin, numerous and parallel; stipules deciduous. Panicles terminal on secondary stems. Flowers zygomorphic only at the androecium and gynoecium level. Sepals 5, coriaceous, unequal, the outer smaller; petals 5, membranaceous, equal, yellow. Outer staminodes numerous, snake-like, short, inner staminodes (except 5 short ones) numerous, longer and thread-shaped; fertile stamens 5, curved toward the upper part of the flower, filaments articulated with the anther, dehiscence by one pore. Ovary 3-carpellate, 1-locular, curved; placentation parietal; ovules numerous. Capsule coriaceous, 3-valved, opening from the apex. Seeds winged. Endemic to the Guayana Shield in eastern Colombia, southern Venezuela, Guyana, and northern Brazil; 2 species, both in the flora area. Key to the Species of Poecilandra 1. 1.

Subshrub to shrub 0.5–2 m tall; leaf blades 1.5–4.5 × 0.6–1.5 cm, apex acuminate ...................................................................................... P. pumila Shrub to treelet 1.5–12 m tall; leaf blades 4–12 × 2–6 cm, apex truncate to emarginate ..................................................................................... P. retusa

Poecilandra pumila Steyerm., Fieldiana, Bot. 28: 371, fig. 75. 1952. Shrub or subshrub 0.5–2 m tall. Upland savannas, 1000–1600 m; Bolívar (Gran Sabana). Guyana. ◆ Fig. 144. Poecilandra retusa Tul., Ann. Sci. Nat. Bot. sér 3, 8: 342. 1847. Treelet or shrub 1.5–12 m tall. Savannas, shrub thickets, on sandstone and white sand. Colombia, Venezuela, Guyana, Brazil; 2 varieties, both in the flora area.

Intermediates between the two varieties occur in the Gran Sabana. Key to the Varieties of P. retusa 1. Leaf blades plane, only the margins revolute; inflorescence large, 6–15 cm long; apex of anthers without excrescence ........................................... var. retusa 1. Leaf blades strongly revolute; inflorescence short, 5–6 cm long; apex of anthers with excrescence ...... var. sclerophylla

Poecilandra 149

Fig. 143. Poecilandra retusa var. sclerophylla

Fig. 144. Poecilandra pumila

Fig. 145. Poecilandra retusa var. retusa

150

O CHNACEAE

P. retusa var. retusa 500–2000 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Cerro Venamo, Gran Sabana, Río Caroní, Roraima-tepui), Amazonas (Cerro Duida, Cerro Sipapo, Cerro Yapacana). Colombia (Vaupés), Guyana (Kaieteur Plateau, Mount Roraima), Brazil (Roraima). ◆Fig. 145.

P. retusa var. sclerophylla (Ule) Sastre, Mém. Soc. Biogéogr. sér. 3, 4: 7. 1994. —Poecilandra sclerophylla Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 341. 1915. Montane forests on tepui slopes and summits, 1500–2800 m; Bolívar (tepuis of the Gran Sabana). Endemic. ◆Fig. 143.

10. SAUVAGESIA L., Sp. Pl. 203. 1753. Leitgebia Eichler in Mart., Fl. Bras. 13(1): 413. 1871. Pentaspatella Gleason, Bull. Torrey Bot. Club 58: 388. 1931. Roraimanthus Gleason, Phytologia 1: 39. 1933. Annual or perennial herbs, subshrubs, or shrubs. Leaves alternate, sessile or petiolate; stipules free, sometimes reduced to a scale, margins with long awns; blades membranous or coriaceous, oblong to ovate or sometimes falciform, generally persistent, apex obtuse; margins crenulate; secondary veins parallel, arcuate and ending tangentially to the margin at a tooth. Inflorescence axillary or terminal, ± compound. Flowers actinomorphic. Sepals 5, green, persistent, equal or unequal; petals 5, white or reddish, membranous. Fertile stamens 5, dehiscence poricidal or longitudinal, outer staminodes 0–many, serpentiform, apex obtuse or reniform, (in S. deflexifolia and S. linearifolia, staminodes in 2 series of 5, inner ones petaloid or sometimes reduced to short scales between the fertile stamens), sometimes forming a corona (S. erioclada). Ovary of 2 or 3 united carpels; placentation parietal, rarely basal or semibasal; ovules few to many (when few, placentation basal). Capsules 2or 3-valved. Seeds alveolate, not winged. Pantropics (predominantly Neotropics); 34 species, 22 in Venezuela, 21 of these in the flora area. Key to the Species of Sauvagesia 1. 1. 2(1). 2. 3(2). 3. 4(2). 4. 5(4). 5. 6(5). 6. 7(6). 7. 8(1). 8.

Inflorescence axillary ................................................................................. 2 Inflorescence terminal ............................................................................... 8 Flower 1 in a reduced inflorescence on expanded axes with 1–4 bracts (2– 4 mm long) ............................................................................................. 3 Flowers 1–7 in a botryoidal inflorescence, bracts none or 1, less than 1 mm long ............................................................................................... 4 Axes with 1 or 2 bracts ............................................................... S. guianensis Axes with 3 or 4 bracts ............................................................ S. imthurniana Herbs .................................................................................................. S. erecta Shrubs or subshrubs .................................................................................. 5 Leaves and sepals falciform; 1 sepal longer than the others ..... S. falcisepala Leaves and sepals ovate to lanceolate; sepals equal ................................. 6 Leaves lanceolate, caducous ...................................................... S. nudicaulis Leaves ovate, persistent ............................................................................ 7 Leaves coriaceous, 1.6–4.3 times as long as wide ........................ S. erioclada Leaves membranous, 4–8 times as long as wide .............................. S. aliciae Outer staminodes in 2 series of 5, petaloid, the series opposite each other ................................................................................................................ 9 Outer staminodes lacking, or 50–70 in 1–3 series, serpentiform or reniform, variously arranged ..................................................................... 10

Sauvagesia 151

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

Leaves elliptic, recurved, 2–4 times as long as wide; placentation parietal ................................................................................................ S. deflexifolia Leaves linear, plane, 6–15 times as long as wide; placentation sub-basal ................................................................................................S. linearifolia Sepals unequal ......................................................................................... 11 Sepals equal ............................................................................................. 13 Placentation basal; outer staminodes never present, inner staminodes reduced ......................................................................................... S. ramosa Placentation parietal; outer staminodes usually present, inner staminodes petaloid ...................................................................................... 12 Leaves ovate, recurved ................................................................... S. amoena Leaves lanceolate, plane .............................................................. S. sprengelii Outer staminodes absent ......................................................................... 14 Outer staminodes present ....................................................................... 15 Subshrub 10–65 cm tall ........................................................... S. roraimensis Annual herb 4–15 cm tall ................................................................. S. tenella Annual herb ............................................................................ S. ramosissima Perennial herb or subshrub ..................................................................... 16 Fleshy herb; inner staminodes spatulate ..................................... S. longipes Erect ligneous herb; inner staminodes ovate ......................................... 17 Ovary 2-carpellate; placentation sub-basal ............................ S. angustifolia Ovary 3-carpellate; placentation parietal ............................................... 18 Stipules glandular with branched cilia; leaves 2–5 times as long as wide .......................................................................................................... S. elata Stipules sparsely glandular, cilia not branched; leaves 5–10 times as long as wide .................................................................................................. 19 Leaves 2.5–9.3 cm long ................................................................ S. longifolia Leaves < 2.5 cm long ................................................................................ 20 Internodes 2–3 mm long; leaves linear to oblong ......................... S. fruticosa Internodes 3–20 mm long; leaves elliptic to lanceolate ............ S. rubiginosa

Sauvagesia aliciae Sastre, Bull. Mus. Hist. Nat. (Paris) sér. 3, 35: 35. 1978. Sauvagesia duidae auct. non Steyerm. 1952: sensu Steyerm., Mem. New York Bot. Gard. 10(4): 16. 1961. Colombia, Venezuela, French Guiana; 2 subspecies, 1 in Venezuela. The second subspecies, subsp. aratayensis Sastre, is known only from French Guiana. S. aliciae subsp. aliciae Subshrub 0.2–1 m tall; petals white. On rocks along streams, 100–200 m; Amazonas (base of Cerro Yapacana). Colombia (Amazonas, Vaupés). ◆ Fig. 146. Sauvagesia amoena Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 341. 1915. Perennial herb with woody base, 10–30 cm tall; petals pink. Swampy savannas, in

association with Brocchinia, 900–1300 m; Bolívar (Auyán-tepui, Gran Sabana). Colombia (Vaupés), Brazil (Amazonas, Pará). ◆ Fig. 152. Sauvagesia angustifolia Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 342. 1915. Sauvagesia striata Gleason & Dwyer, Brittonia 3: 169. 1939. Suffructescent, 10–60 cm tall; petals white. White-sand savannas, beaches along rivers, 400–1500 m; Bolívar (Gran Sabana, Macizo del Chimantá, Río Caroní). Guyana, Suriname. Sauvagesia deflexifolia Gardner in Hook., Icon. Pl. 5: pl. 484. 1842. Annual herb 10–60 cm tall; petals white. Savannas, ca. 100 m; Amazonas (La Esmeralda). Colombia (Vaupés, Vichada),

152

O CHNACEAE

Brazil (Amazonas, Mato Grosso), Bolivia (La Paz). ◆Fig. 151.

ligneous vegetation, 700–800 m; Amazonas (Cerro Aracamuni). Endemic.

Sauvagesia elata Benth., J. Bot. (Hooker) 4: 107. 1842. Sauvagesia elata subsp. occidentalis Maguire & Wurdack, Mem. New York Bot. Gard. 10(4): 17. 1967. Suffrutescent, 0.6–1 m tall; petals white. In clearings, along streams, on white sand, 50–100 m; Delta Amacuro (Serranía de Imataca), Amazonas (Río Sipapo). Panama, Guyana, Suriname, French Guiana, Brazil (Amazonas, Pará).

Sauvagesia fruticosa Mart., Nov. Gen. Sp. Pl. 1: 38. 1824. Sauvagesia rosacea Ule, Bot. Jahrb. Syst. 40: 432. 1908. Leitgebia colombiana R. Schult., Bot. Mus. Leafl. 16: 85. 1953. Ligneous herb to subshrub, 30–50 cm tall; petals white or pinkish. Savannas on quartzite, 1200–2300 m; Bolívar (Auyán-tepui, Cerro Jaua, Gran Sabana), Amazonas (Cerro Duida, Cerro Yapacana, Sierra Parima). Colombia (Amazonas, Vaupés), Peru, Brazil (Amazonas: Rio Negro, Serra Aracá).

Sauvagesia erecta L., Sp. Pl. 203. 1753. Neotropics from Mexico to northern Argentina, tropical Africa, Madagascar; 2 subspecies, 1 in Venezuela. S. erecta subsp. erecta Distribution as in species; 2 varieties, 1 in Venezuela. S. erecta subsp. erecta var. erecta Sauvagesia adima Aubl., Hist. Pl. Guiane 252, t. 1000, fig. a. 1793. Perennial herb 10–60 cm tall. In savannas, clearings, along streams and roads, secondary vegetation, near sea level to 1200 m; Delta Amacuro (Serranía de Imataca), widespread in Bolívar and Amazonas. Widespread elsewhwere in Venezuela; other distribution as in species (though absent from Cuba and Jamaica). ◆Fig. 148. Sauvagesia erioclada Maguire & K.D. Phelps, Bol. Soc. Venez. Ci. Nat. 14: 13. 1951. Sauvagesia duidae Steyerm., Fieldiana, Bot. 28: 372, fig. 76. 1952. Sauvagesia erioclada var. grandiflora Maguire & Wurdack, Mem. New York Bot. Gard. 10(4): 16. 1961. Shrub or subshrub 0.5–3 m tall; leaves coriaceous; petals white or pinkish, the corona purple. On quartzite outcrops, along streams, savannas, 800–1800 m; Amazonas (Cerro Coro Coro, Cerro Duida, Cerro Yutajé). Endemic. Sauvagesia falcisepala Sastre, Bull. Mus. Hist. Nat. (Paris) sér. 4, 8: 13, pl. 1. 1986. Subshrub ca. 50 cm tall; leaves coriaceous; petals white. On quartzite outcrops, on low

Sauvagesia guianensis (Eichler) Sastre, Caldasia 10: 507. 1970. —Leitgebia guianensis Eichler in Mart., Fl. Bras. 13(1): 413. 1871. Colombia, Venezuela, Guyana; 2 subspecies, 1 in Venezuela. The second subspecies, subsp. araracuarensis Sastre, occurs in Colombia. S. guianensis subsp. guianensis Leitgebia gleasoniana Lasser, Bol. Acad. Ci. Fis. 9: 246. 1945. —Sauvagesia guianensis subsp. gleasoniana (Lasser) Steyerm., Ann. Missouri Bot. Gard. 74: 101. 1987. Sauvagesia guianensis subsp. guaiquinimensis Steyerm., Ann. Missouri Bot. Gard. 74: 101. 1987. Sauvagesia guianensis subsp. sipapoensis Steyerm., Ann. Missouri Bot. Gard. 74: 101. 1987. Sauvagesia marahuacensis Steyerm., Ann. Missouri Bot. Gard. 74: 98. 1987. Subshrub 0.2–1 m tall; stem black; petals white or pale pink. On quartzite outcrops, 1100–2100 m; Bolívar (Cerro Guaiquinima, Gran Sabana), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Sipapo, Sierra de la Neblina). Guyana. ◆ Fig. 155. Sauvagesia imthurniana (Oliv.) Dwyer, Bull. Torrey Bot. Club 67: 291. 1940. —Leitgebia imthurniana Oliv., Trans. Linn. Soc. London, Bot. ser. 2, 2: 271. 1877. —Roraimanthus imthurniana (Oliv.) Gleason, Phytologia 1: 39. 1933. Sauvagesia imthurniana subsp. chiman-

Sauvagesia 153

tensis Maguire, Steyerm. & Wurdack, Mem. New York Bot. Gard. 10(4): 17. 1961. Subshrub 0.15–1 m tall; stem black; petals white or pale pink. On tepui summits, 1900–2900 m; Bolívar (Auyán-tepui, Macizo del Chimantá, Roraima-tepui), Amazonas (Cerro Marahuaka). Endemic. ◆Fig. 153. Sauvagesia linearifolia A. St.-Hil., Mém. Mus. Hist. Nat. 11: 106. 1824. Colombia, Venezuela, Peru, Brazil (Distrito Federal, Goiás, Mato Grosso, Minas Gerais); 2 subspecies, 1 in Venezuela. S.

linearifolia subsp. venezuelensis Maguire & Wurdack, Mem. New York Bot. Gard. 10(2): 18. 1961. Subshrub 0.3–1 m tall; petals white. Savannas, 100–200 m; Amazonas (Río Casiquiare, middle Río Orinoco, Río Ventuari). Endemic. ◆ Fig. 156. Sauvagesia longifolia Eichler in Mart., Fl. Bras. 13(1): 407, fig. 83, 1. 1871. Subshrub 20–80 cm tall; petals pale pink. Savannas, 100–1100 m; Bolívar (Auyántepui, Macizo del Chimantá [Churí-tepui]), Amazonas (Río Siapa). Guyana, French Guiana, Brazil (Amazonas, Bahia, Pará). Sauvagesia longipes Steyerm., Fieldiana, Bot. 28: 374, fig. 77. 1951. Perennial prostrate herb; inflorescence umbellately branched; petals whitish green. On moss-covered rocks, upper montane forests on sandstone, sometimes epiphytic, 1000–1500 m; Bolívar (Auyán-tepui, Cerro Venamo, Gran Sabana). Guyana (MazaruniCuyuní rivers, Pakaraima). Sauvagesia nudicaulis Maguire & Wurdack, Mem. New York Bot. Gard. 10(4): 17. 1961. Subshrub 15–50 cm tall; outer staminodes absent; petals white. Savannas, 100–200 m; Amazonas (Cerro Yapacana base, Río Atabapo, Río Guainía, Río Siapa, Río Ventuari). Colombia (Vaupés: Río Atabapo). ◆ Fig. 149. Sauvagesia ramosa (Gleason) Sastre, Caldasia 10: 506. 1970. —Pentaspatella ramosa Gleason, Bull. Torrey Bot. Club 58: 398. 1931.

Sauvagesia duckei Sleumer, Repert. Spec. Nov. Regni Veg. 42: 263. 1937. Subshrub or shrub 0.3–1 m tall; petals pink. Savannas, 100–300 m; Bolívar (Cusimi), Amazonas (Cerro Yapacana, La Esmeralda, Río Atabapo, Río Ventuari). Brazil (Amazonas, Pará, Roraima). ◆Fig. 147. Sauvagesia ramosissima Spruce ex Eichler in Mart., Fl. Bras. 13(1): 409, fig. 82, 2. 1871. Sauvagesia miniata Steyerm., Fieldiana, Bot. 28: 376. 1952. Much-branched annual herb, 7–37 cm tall; petals white. Sandy soils, along rivers, savannas, secondary vegetation, 100–1000 m; Bolívar (Auyán-tepui, Cerro Bolívar, Cerro Guaiquinima), Amazonas (Puerto Ayacucho, San Fernando de Atabapo). Guárico, Portuguesa, Sucre; Colombia (Meta, Tolima, Vichada), Guyana, Suriname, French Guiana, Brazil (Amazonas, Pará, Rondônia, Roraima). ◆ Fig. 157. Sauvagesia roraimensis Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 344. 1915. Sauvagesia deficiens A.C. Sm., Lloydia 2: 194. 1939. Sauvagesia smithiana Dwyer, Bull. Torrey Bot. Club 72: 536, fig. 1K, M. 1945. Perennial herb or subshrub 15–45 cm tall; petals white. Savannas on quartzite outcrop, 600–2200 m; Bolívar (Auyán-tepui, Macizo del Chimantá [Chimantá-tepui], Gran Sabana, Roraima-tepui), Amazonas (Sierra Parima). Guyana, Suriname. Sauvagesia rubiginosa A. St.-Hil., Bull. Soc. Philom. Paris 1823: 173. 1823. —Sauvagesia erecta var. rubiginosa (A. St.-Hil.) Eichler in Mart., Fl. Bras. 13(1): 409. 1871. Sauvagesia laxa Mart., Nov. Gen. Sp. Pl. 1: 38. 1824. Sauvagesia surinamensis Miq., Linnaea 18: 744. 1844. Ligneous herb or subshrub 15–80 cm; petals white or pink. Savannas, 100–1300 m; Bolívar (Ciudad Bolívar, Gran Sabana, Río Hacha, Río Paragua), Amazonas (Río Ventuari). Anzoátegui, Guárico, Monagas; Colombia (Amazonas), Trinidad, Guyana, Suriname, French Guiana, Brazil (Amapá, Amazonas: Río Branco, Maranhão, Pará).

154

O CHNACEAE

Fig. 146. Sauvagesia aliciae subsp. aliciae

Fig. 148. Sauvagesia erecta subsp. erecta var. erecta

Fig. 147. Sauvagesia ramosa

Fig. 149. Sauvagesia nudicaulis

Sauvagesia 155

Fig. 150. Sauvagesia sprengelii

Fig. 152. Sauvagesia amoena

Fig. 151. Sauvagesia deflexifolia

Fig. 153. Sauvagesia imthurniana

Fig. 154. Sauvagesia tenella

156

O CHNACEAE

Fig. 155. Sauvagesia guianensis subsp. guianensis

Fig. 156. Sauvagesia linearifolia subsp. venezuelensis

Fig. 157. Sauvagesia ramoisissima

Tyleria 157

Sauvagesia sprengelii A. St.-Hil., Bull. Soc. Philom. Paris 1823: 173. 1823. Sauvagesia erecta Aubl., Hist. Pl. Guiane 254, fig. 100b. 1775, non L. 1753. Sauvagesia serpyllifolia Mart., Nov. Gen. Sp. Pl. 1: 37. 1824. Sauvagesia kappleri Miq., Stirp. Surinam. Select. 105. 1850 [1851]. Ligneous-based herb or subshrub 10–70 cm tall; petals pink. Savannas, 50–1000 m; Bolívar (Ilú-tepui, upper Río Caroní, Río Paragua, Sierra de Lema). Trinidad, Guyana, Suriname, French Guiana, Brazil (Amapá, Amazonas: Rio Branco, Bahia, Ceará, Pará, Paraiba, Rio de Janeiro). ◆Fig. 150.

Sauvagesia tenella Lam., Tabl. Encycl. 2: 119. 1793 [1797]. Sauvagesia inconspicua Dwyer, Lloydia 2: 195. 1939. Annual herb, sometimes reduced to a small (2–15 cm tall) plant with 4 or 5 leaves and 1 flower; petals white. Savannas, 100– 200 m; northwestern Bolívar, Amazonas (Río Manapiare). Guárico, Portuguesa; Central America, Cuba, Hispaniola, Colombia, Guyana, Suriname, French Guiana, Brazil (Amazon Basin, Goiás, Bahia, Mato Grosso), Bolivia. ◆Fig. 154.

11. TYLERIA Gleason, Bull. Torrey Bot. Club 58: 391. 1931. Treelets or shrubs, 0.2–15 m tall, bark black. Leaves sessile or petiolate; secondary veins thin, numerous and parallel, apex acute (obtuse in T. grandiflora and T. spathulata); margins denticulate or toothed. Inflorescence terminal simple or branched. Flowers actinomorphic. Sepals 5, coriaceous, glandular or not; petals 5, membranaceous, white to pink. Staminodes 5, petaloid, keeled, adnate at the base (free in T. pendula, united in T. breweriana) forming a corona, with adnate appendices (free in T. pendula), outer staminodes only in T. terrae-humilis, fertile stamens 5, shorter than the staminodes (longer in T. breweriana). Ovary 3-carpellate, 1-locular, placentation parietal. Capsules coriaceous, 3-valved. Seeds winged. Endemic to the Guayana Shield in southern Venezuela and northern Brazil; 13 species, all in the flora area. Key to the Species of Tyleria 1. 1. 2(1). 2. 3(2). 3. 4(2). 4. 5(1). 5. 6(5). 6. 7(6). 7. 8(7). 8.

Leaves noticeably petiolate ....................................................................... 2 Leaves sessile, subsessile, or shortly petiolate ......................................... 5 Leaves 7–9 cm long .................................................................................... 3 Leaves 3.5–5 cm long ................................................................................. 4 Leaf blades with acumen 1.8–2.2 cm long; inflorescence pendulous ..................................................................................................... T. pendula Leaf blades with acumen 0.8–1 cm long; inflorescence erect ......... T. silvana Leaf blades elliptic, 4–5 × 1.7–2.2 cm ....................................... T. breweriana Leaf blades oblong, 3.5–4 × 1–1.2 cm ...................................... T. tremuloidea Leaf blades subsessile or shortly petiolate ............................................... 6 Leaf blades clearly sessile ....................................................................... 10 Leaf blades spathulate ............................................................... T. spathulata Leaf blades linear or oblong ...................................................................... 7 Inflorescence branched, with 10 flowers or more ..................................... 8 Inflorescence not branched, with 1–5 flowers ........................................... 9 Margin of the leaf blades plane; sepals, 2.5–4 mm long; staminodes free in 2 series .......................................................................... T. terrae-humilis Margin of the leaf blades not plane; sepals 8–9 mm long; staminodes united at base in 1 series .......................................................... T. apiculata

158

O CHNACEAE

9(7). Leaf blades oblong, aristate ............................................................ T. aristata 9. Leaf blades linear, not aristate ........................................................ T. linearis 10(5). Inflorescences branched; sepals 4–12 mm long ...................................... 11 10. Inflorescences reduced with 1–3 flowers, not branched; sepals 14–18 mm long ....................................................................................................... 12 11(10). Leaves borne at the apex of stems; sepals 4–6 mm long, subrectangular with glandular cilia ................................................................ T. spectabilis 11. Leaves borne along the stem; sepals 10–12 mm long, lanceolate, not ciliate ....................................................................................... T. floribunda 12(10). Shrubs 1–2 m tall; leaf blades rounded at apex; petals 3 cm long ............................................................................................... T. grandiflora 12. Trees 2–8 m tall; leaf blades acute at apex; petals 2 cm long ...... T. phelpsiana Tyleria apiculata Sastre, Phytologia 59: 313. 1986. Shrub or small tree 2–3 m tall. Tepui slopes, 1200–1800 m; Bolívar (Macizo del Chimantá), Amazonas (Cerro Marahuaka). Endemic. Tyleria aristata Maguire & Wurdack, Mem. New York Bot. Gard. 10(4): 13, fig. 25F. 1961. Shrub 20–80 cm tall. Meadows along streams, 1800–2000 m; Amazonas (Sierra de la Neblina). Endemic. Tyleria breweriana Steyerm., Ann. Missouri Bot. Gard. 71: 322, fig. 9. 1984. Shrub 1–1.5 m tall. Low vegetation on rocks, ca. 2000 m; Bolívar (Cerro Jaua). Endemic. Tyleria floribunda Gleason, Bull. Torrey Bot. Club 58: 392. 1931. Treelet 3–12 m tall. Open scrubby slopes, 1200–2100 m; Bolívar (Cerro Marutaní), Amazonas (Cerro Duida). Endemic. ◆Fig. 160. Tyleria grandiflora Gleason, Bull. Torrey Bot. Club 58: 393. 1931. Shrub 1–2 m tall. On ridges of tepui summits, 1800–2500 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka). Endemic. ◆Fig. 159. Tyleria linearis Gleason, Bull. Torrey Bot. Club 58: 393. 1931. Shrub, forming dense thickets. Along streams on tepui summit, 1600–2100 m; Amazonas (Cerro Duida). Endemic. Tyleria pendula Maguire & Wurdack, Mem. New York Bot. Gard. 10(4): 11, figs. 25H, 26. 1961.

Slender shrub 1–2 m tall; leaves spatulate; inflorescence nodding, reddish; petals white, basally yellow. Tepui meadows, 2000– 2100 m; Amazonas (Sierra de la Neblina). Endemic. Tyleria phelpsiana Maguire & Steyerm., Mem. New York Bot. Gard. 23: 867, fig. 9. 1972. Treelet 2–8 m tall; leaves branched at apex of stems, erect-ascending; petals white, yellow at base. Open tepui slopes, 1900–2100 m; Bolívar (Cerro Jaua). Endemic. Tyleria silvana Maguire, Mem. New York Bot. Gard. 23: 868. 1972. Treelet 1–4 m tall; corolla and stamens white. Tepui slopes, 1300–1600 m; Amazonas (Cerro Aracamuni). Brazil (Amazonas: Serra Pirapucú). Tyleria spathulata Gleason, Bull. Torrey Bot. Club 58: 394. 1931. Treelet or shrub 2–15 m tall; petals white; anthers red. Low bush with Bonnetia, or in Brocchinia stands, 1200–2000 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú). Endemic. ◆Fig. 158. Tyleria spectabilis Maguire & Wurdack, Mem. New York Bot. Gard. 10(4): 11, fig. 25E. 1961. Candelabra-like treelet 3–12 m tall; petals pink, basally yellow. Rocky, low vegetation, 1800–2200 m; Amazonas (Sierra de la Neblina). Endemic. Tyleria terrae-humilis Sastre, Novon 8: 303. 1998. Shrub ca. 2 m tall; leaf blade papery, 7–11 cm long; inflorescence 7–8 cm long; stami-

Tyleria 159

Fig. 158. Tyleria spathulata

Fig. 159. Tyleria grandiflora

Fig. 160. Tyleria floribunda

160

O CHNACEAE

nodes petaloid, 5 outer and 5 inner. In sparse lowland forests, 100–200 m; Amazonas (San Antonio to San Fernando de Atabapo). Endemic. Tyleria tremuloidea Maguire & Wurdack, Mem. New York Bot. Gard. 10(4): 14, fig. 25G. 1961. Fastigiate treelet 3–6 m tall; petals white, orange at base. An abundant or dominant element in some scrub forests, Montane thickets, 2000–2300 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 161.

Fig. 161. Tyleria tremuloidea

12. WALLACEA Spruce ex Benth. & Hook. f., Gen. Pl. 1: 320. 1862. Trees, treelets, or shrubs. Leaves short-petiolate; stipules deciduous, 2–7 cm long; blades coriaceous, elliptic to obovate, the apex truncate to emarginate; secondary veins thin, numerous, curved and parallel. Inflorescence axillary, in short racemes. Flowers zygomorphic only at androcium and gynoecium level. Sepals 5, coriaceous, subequal, ovate; petals 5, equal, elliptic, white to pinkish. Fertile stamens 5, curved to the upper part of the flower; filaments articulate with the anther, functionally dehiscent by short longitudinal slits; staminodes 0–many, thread-like. Ovary 2- or 3-carpellate, 1-locular, curved in the lower part, placentation parietal; ovules numerous. Capsule ligneous, ovoid, with short style, 2- or 3-valved, the valves separating completely. Seeds not winged. Southeastern Colombia, southern Venezuela, Brazil (Amazonas, Pará); 3 species, 2 in Venezuela, both in the flora area. Key to the Species of Wallacea 1.

1.

Petioles 0.5–1 cm long; leaf blades 4.5–15 × 2–9 cm; inflorescence axis 0.5–11 cm long, with 2–4 flowers; sepals and petals ca. 2 cm long; carpels 2 (rarely 3) ..................................................................... W. insignis Petioles 2–3 cm long; leaf blades 11–19 × 9–11 cm; inflorescence axis 10– 18 cm long, with 10–12 flowers; sepals and petals ca. 1 cm long carpels 3 (rarely 2) .............................................................................. W. multiflora

Wallacea 161

Wallacea insignis Spruce ex Benth. & Hook. f., Gen. Pl. 1: 321. 1862. Tree, treelet, or shrub 3–20 m tall; Blackwater riverbanks, white-sand savannas. Colombia, Venezuela, Brazil (Amazonas, Pará); 2 subspecies, 1 in Venezuela. W. insignis subsp. insignis. —Curame. 100–200 m; Amazonas (Brazo Casiquiare, base of Cerro Yapacana, La Esmeralda, Río Guainía, Río Negro). Southeastern Colombia,

Fig. 162. Wallacea multiflora

Brazil (Amazonas: Rio Uaupes, upper Rio Negro). Wallacea multiflora Ducke, Arq. Inst. Biol. Veg. 1: 207. 1935. Small tree or shrub 5–6 m tall; flowers pale pink. Black-water riverbanks, 100–200 m; Amazonas (Río Atabapo, Río Casiquiare, Río Negro). Brazil (Amazonas: Rio Uaupes, Rio Negro). ◆Fig. 162.

162

O LACACEAE

OLACACEAE by John M. MacDougal Trees, shrubs, or rarely lianas, sometimes with milky sap, glabrous or nearly so, less often pubescent, hermaphroditic or rarely dioecious. Plants green and photosynthetic, but many species hemiparasitic, attaching to roots of other plants. Leaves alternate, simple, evergreen (deciduous in Ximenia), petiolate, exstipulate; blades unlobed, entire (sinuate-dentate in Brachynema), often rough-textured, rarely with resin dots, pinnately veined (in most Venezuelan species) or rarely subpalmately veined (pliveined). Inflorescence axillary, fasciculate or in racemes, panicles, spikes, or rarely umbels, sometimes cauliflorous from reduced cushion-like areas, or flowers rarely solitary; bracts or bracteoles sometimes accrescent or connate to form an epicalyx or small collar below the calyx. Flowers bisexual (unisexual in an African species of Aptandra), sometimes heterostylous, actinomorphic, usually greenish, yellowish, or whitish (sometimes red in Schoepfia). Calyx small at anthesis, cupular, shallowly (3)4–6(7)-lobed, -toothed, or -crenulate, the teeth rarely obsolete, the calyx often accrescent, large, and conspicuous (sometimes becoming colored) in fruit; corolla (3)4–6(7)-merous, deciduous, usually valvate, the petals free or connate basally to form a tube (or corolla tubular with short apical lobes in Schoepfia). Nectary disk intrastaminal and surrounding the ovary (sometimes covering ovary up to the style), or extrastaminal and annular or of free glands alternate the petals or absent. Stamens as many as the petals and opposite them (antepetalous), rarely alternate with petals (antesepalous), or epipetalous and opposite or rarely alternate with the corolla lobes, sometimes 2(3) times as many as petals, in 1(–3) whorls, some occasionally staminodal (only 3 functional stamens in Dulacia), if 10 or more often of alternating lengths; filaments free or adnate basally to corolla, sometimes adnate most of length, or rarely connate around style (Aptandra); anthers opening by longitudinal slits or apical pores. Carpels (2)3(–5); ovary superior, sometimes partially inferior and immersed in the disk, rarely truly inferior and adnate to cuplike receptacle (Schoepfia), 2–5-locular basally and often 1-locular distally; ovules 1 per basal locule; style, if any, conical, columnar, or filiform; stigma 2–5-lobed, sometimes capitate. Fruit nearly always 1-seeded, a drupe or rarely nut-like, or apseudodrupe when concrescent with the cup-shaped floral axis (Schoepfia), or with the accrescent fleshy calyx (Dulacia) or accrescent fleshy disk (Cathedra). Seed with thin testa or testa absent; endosperm copious; embryo small or minute, apical in endosperm. Pantropics (a few subtropical), most diverse in Neotropics and Africa; ca. 27–29 genera and ca. 200 species, 9 genera and 31 species in the flora area. The neotropical elements of this family were monographed by Sleumer (Flora Neotropica 38: 1–159. 1984), and this treatment is largely based upon that work. Key to the Genera of Olacaceae based on flowering specimens 1.

Leaf margin shallowly sinuate-dentate, the teeth with a conspicuous central vein that is gland-tipped; petioles 5–20 cm long, of markedly different lengths on the same branch, thickened or pulvinulate both basally and apically; corolla 20–30 mm long, connate-tubular most of length; stamens 5 (rarely 4), the connective apically elongate, attenuate .............................................................................. 2. Brachynema

O L A C A C E A E 163

1.

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

6.

7(6).

7.

8(4).

8.

9(8).

9.

Leaf margin entire; petioles 0.1–2(–3) cm long, of equal or only slightly different lengths on the same branch, thickened only apically or not at all; corolla 1.5–10(–12) mm long (or in Chaunochiton, one species with corolla (30–)40–65 mm long), connate-tubular to campanulate or petals free; stamens 3–12, the connective not apically elongate .............. 2 Stamens 8–45(–60) mm long, the same number as petals (5); corolla 8–65 mm long ............................................................................. 4. Chaunochiton Stamens 0.5–5(–6) mm long, 4–10(–12) in number, the same as or twice the number of petals; corolla 1.8–7(–10 in Dulacia) mm long ............. 3 Stamens 3; staminodes 6, apically bifid; petals 6 .......................... 5. Dulacia Stamens 4–15; staminodes absent; petals or corolla lobes 4–6 ................ 4 Stamens the same number as petals ......................................................... 5 Stamens twice (or sometimes 3 times) the number of petals ................... 8 Stamens antesepalous, alternate with petals .............................. 6. Heisteria Stamens antepetalous, opposite the petals or epipetalous on corolla tube and opposite the corolla-lobes ............................................................... 6 Pedicel with 3 bracts/bracteoles at apex which are ± accrescent basally and form an acutely 3-lobed epicalyx; flower axis thickened, calyx-like, usually not hidden by the epicalyx, bearing at apex rudiments of calyx or calyx lobes, i.e., calyx adnate and continuous with the pedicel, inconspicuous to practically absent; petals 5, connate in at least the lower 2/3 to form a tube; stamens adnate to corolla tube most of their length; ovary inferior ............................................................... 8. Schoepfia Pedicel without epicalyx at apex, or epicalyx-like structure obtusely 5lobed; calyx (cup or tube with lobes or teeth) present, ± conspicuous; petals 4–6, free or eventually so; stamens free or connate into a separate tube; ovary superior or partially inferior and immersed in the disk ................................................................................................................ 7 Pedicels 1–2 mm long, the bracts connate into a cup-like (often slightly 5lobed) small collar shortly below and hiding the calyx; inflorescences of subsessile aggregates of axillary fascicles; petals (4–)6; stamens free, not connate ................................................................................ 3. Cathedra Pedicels 10–15 mm long, without bracts or bracteoles or a collar below and hiding the calyx; inflorescences of pedunculate axillary umbels; petals 4; filaments connate into a tube; anthers connate into a ring .................................................................................................. 1. Aptandra Petals remaining united into a corolla tube in the lower half; lower surface of leaf blade often puberulent on the veins and when dried darkpunctate with resinous spots; young stems and inflorescences with minute branched trichomes ................................................. 7. Minquartia Petals free or at least finally so; leaf blades glabrous, when dried light- or pellucid-punctate, not punctate, or leaves drying black; young stems and inflorescences glabrous or nearly so .............................................. 9 Petals ca. 4–6 mm long, densely barbed inside with long introrse trichomes; branchlets with spines or ending in thorns; leaves with 3– 5(–7) pairs of inconspicuous lateral veins ................................. 9. Ximenia Petals 1.5–4(–6) mm long, pubescent, short-bearded, or glabrous inside; branchlets lacking spines or thorns; leaves with (4–)6–12(–15) pairs of usually conspicuous lateral veins ............................................. 6. Heisteria

164

O LACACEAE

Key to the Genera of Olacaceae based on fruiting specimens 1.

1.

2(1).

2.

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

5(4).

5.

6(5). 6.

7(3).

7. 8(7).

8.

Leaf margin shallowly sinuate-dentate, the teeth with a conspicuous central vein that is gland-tipped; petioles 5–20 cm long, of markedly different lengths on the same branch, thickened or pulvinulate both basally and apically ............................................................. 2. Brachynema Leaf margin entire; petioles 0.1–2(–3) cm long, of equal or only slightly different lengths on the same branch, thickened or pulvinulate only apically or not at all ............................................................................... 2 Two collars present at base of or below fruit, i.e., calyx short and erect or slightly spreading, collar-like, subtended by another smaller ring of fused bracts, these often slightly adnate to base of calyx; outer fleshy layer of fruit formed by accrescent disk, covering fruit to almost the top ................................................................................................... 3. Cathedra One or no collar present at base of or below fruit, i.e., calyx or bracts collar-like, but not both, the calyx of various forms (cupular, covering fruit, winglike, or practically absent); outer layer of fruit derived from ovary or from adnate/accrescent calyx which may cover fruit almost to the top (or from floral axis and remnants of calyx in Schoepfia) ........ 3 Calyx much enlarged towards fruiting stage, either remaining partly free or enveloping the mature fruit, or completely adnate to the fruit ...... 4 Calyx hardly or not enlarged towards fruiting stage ............................... 7 Accrescent calyx finally almost covering the whole fruit, of which only the umbonate top with the style base remains free ......................... 5. Dulacia Accrescent calyx finally almost covering the fruit only in its basal or lower part, otherwise frill-like, expanding and horizontal or suberect, or expanded and reflexed .............................................................................. 5 Infructescence fasciculate, the pedicels borne from axillary cushions or glomerules; petioles usually markedly canaliculate adaxially ................................................................................................... 6. Heisteria Infructescence corymbose-paniculate, the pedicels borne from a rachis or peduncle 4–15(–30) mm long; petioles not markedly adaxially canaliculate ............................................................................................ 6 Leaf blades rounded at base; petiole well defined and usually more than 6 mm long ................................................................................ 1. Aptandra Leaf blades cuneate or narrowed at base, often attenuate or decurrent to a short and sometimes indistinct petiole, the petiole often < 5 mm long ........................................................................................... 4. Chaunochiton Pedicel with (2)3 concrescent bracteoles (epicalyx) at the very apex, persistent beneath fruit; proper calyx practically absent, i.e. adnate to the thick flower axis, its remains visible as a low crown toward apex of fruit .......................................................................................... 8. Schoepfia Pedicel without epicalyx at apex; free calyx ± conspicuous, present at base of fruit ............................................................................................ 8 Branchlets usually with spines or ending in a thorn; leaves often clustered on lateral short shoots, ± deciduous, the apices finely mucronate .................................................................................................... 9. Ximenia Branchlets without spines and never ending in a thorn; leaves not clustered on short shoots, usually persistent, the apices not mucronate ............ 9

Aptandra 165

9(8).

9.

Lower surface of leaf blade often puberulent on the veins and when dried dark-punctate with resinous dots; fruit borne from elongate spikes ............................................................................................... 7. Minquartia Lower surface of leaf blade always glabrous and when dried without dark-punctate resinous dots (sometimes pellucid-punctate or with dark linear laticifers); fruit borne from axillary subsessile cushions or glomerules ................................................................................. 6. Heisteria

1. APTANDRA Miers, Ann. Mag. Nat. Hist. ser. 2, 7: 201. 1851. Glabrous shrubs or trees, sometimes dioecious (West Africa). Leaves entire, pinnately veined, micropunctate against strong light, the laticifers narrow, hardly or not visible beneath with the naked eye. Inflorescence axillary, of pedunculate slender few- to many-flowered panicles, sometimes with small scaly and caducous prophylls on lower part of peduncle. Flowers bisexual (or unisexual), small, with filiform pedicels. Calyx patelliform, deeply 4-dentate or -lobed, small in anthesis but very much enlarged towards fruiting stage; petals 4, valvate, thin-fleshy, linearligulate, in bud coherent into a cylindrical tube which is globular-dilated distally, at full anthesis free, convolute, and revolute, pale green or white. Disk glands 4, free, petaloid, alternating with the petals, inserted on flower axis between petals and staminal tube, thickish, rounded. Stamens 4; filaments connate in a cylindrical distally thickened tube which surrounds ovary and style; anthers oblong, concrescent in a ring with the thickish connectives, each anther opening from apex to base by a membranaceous finally reflexed valve. Ovary substipitate, 2-sulcate, 2-locular below; style slender, clavate-thickened distally, as long as the staminal tube; stigma obtuse, small. Fruit a single drupe developed at end of inflorescence, rather large, included in its lower half by the much enlarged fruit-calyx, the latter cup-shaped, rather thin, its upper part erect or spreading, i.e. frill-like expanded; pericarp coriaceous; endocarp lignescent. Cotyledons orbicular, foliaceous. Amazonian Venezuela, Guyana, Peru, Brazil, Bolivia, West Africa; 4 species, 2 in Venezuela. Key to the Species of Aptandra 1.

1.

Leaves subcoriaceous, stiff, the lower surface opaque-glaucescent, base usually rounded or sometimes subcordate; corolla tube cylindric, but somewhat widened below the middle before splitting into distinct petals .................................................................................... A. liriosmoides Leaves membranous to firmly chartaceous, flexible, note opaqueglaucescent on the lower surface, base usually cuneate or rarely obtuse; corolla tube cylindric for entire length until splitting into distinct petals .......................................................................................... A. tubicina

Aptandra liriosmoides Spruce ex Miers, Ann. Mag. Nat. Hist. ser. 3, 4: 360. 1859. Aptandra benthamiana Miers, Ann. Mag. Nat. Hist. ser. 3, 4: 360. 1859. Shrub or small tree 3–8 m tall; leaves rounded or sometimes subcordate at base; flowers white; drupe subglobular, 1.5–2 cm diameter. Forest patches in white-sand sa-

vannas, seasonally flooded forests, 50–200 m; Amazonas (Caño San Miguel, base of Cerro Yapacana, Río Atabapo basin, near San Carlos de Río Negro). Guyana, Brazil (Amazonas, Roraima). ◆Fig. 163. Aptandra tubicina (Poepp.) Benth. ex Miers, Ann. Mag. Nat. Hist. ser. 3, 4:

166

O LACACEAE

360. 1859. —Heisteria tubicina Poepp., Gen. Pl. 3: 35, t. 241 (pro parte). 1843. Aptandra spruceana Miers, Ann. Mag. Nat. Hist. ser. 2, 7: 202. 1851. Chaunochiton mouriroides A.C. Sm., Brittonia 2: 148. 1936. Tree ca. 20 m tall; leaves cuneate or obtuse at base; inflorescence many-flowered; flowers greenish-white; drupe subglobular, 2–2.5 cm diameter. Evergreen lowland forest, 100–200 m; Bolívar (middle Río Erebato). Ecuador, Peru, Brazil, Bolivia.

Fig. 163. Aptandra liriosmoides

2. BRACHYNEMA Benth., Trans. Linn. Soc. London 22: 126, t. 22. 1857, nom. cons. Few-branched slender treelet or small tree. Leaves pinnately veined, shallowly sinuate-dentate, the ± conspicuous teeth always minutely glandular-tipped; petioles long, usually of markedly different lengths, pulvinulate or thickened and irregularly wrinkled (when dried) at both the base and apex. Inflorescence a short axillary ebracteate corymb, either in the upper leaf axils or else on the lower, leafless part of the stem; pedicels short, thick. Flowers bisexual, 4- or 5-merous. Calyx obscurely (4)5-dentate or subentire, accrescent towards fruiting stage; petals 4 or 5, connate almost entirely to form a tubular corolla which is purplish at least basally or with transverse purplish bands in B. ramiflorum, the free lobes valvate or slightly contorted, finally recurved-expanded, glabrous or with a central row of trichomes inside. Stamens 4 or 5, alternating with the corolla lobes, inserted at base of corolla tube with short free filaments; anthers extrorse, longitudinally dehiscent; connective conspicuously apically elongate. Disk absent. Ovary superior, sessile, (3)4- or 5locular; stigma sessile, pulviniform, hardly lobed. Drupe subglobose to obloid, apically mucronate, surrounded at base by the accrescent calyx; pericarp very thin; endocarp crustaceous, fibrous inside. Seed vertically striate, with a unilateral groove. Venezuela (Amazonas), Peru, Amazonian Brazil; 2 species, 1 in Venezuela. With its dentate leaves, long pulvinate petioles, and floral characters, Brachynema is anomalous in the Olacaceae. With further study, it will almost certainly be placed in another family where it is more consistent. Brachynema axillare was recently segregated from B. ramiflorum Benth., which has striped corolla tubes with internally pubescent lobes and flowers appearing on the lower, leafless portion of the stems.

Cathedra 167

Fig. 164. Brachynema axillare

Brachynema axillare R. Duno & P.E. Berry, Novon 5: 238. 1995. Shrub or tree 3–8 m tall; corolla narrowly tubular, 15–25 mm long, purplish basally, the free lobes white; drupe ca. 2–2.5 cm diameter, green to purplish black. Evergreen lowland to premontane forests, 100–200 m; Amazonas (Río Mawarinuma, base of Sierra de la Neblina). Peru (Loreto), Brazil (Amazonas). ◆Fig. 164.

3. CATHEDRA Miers, Ann. Mag. Nat. Hist. ser. 2, 7: 452. 1851. Shrubs or trees. Youngest branchlets minutely papillate-puberulent with simple trichomes, quickly glabrescent. Leaves entire, pinnately-veined. Inflorescence of axillary fasicles; pedicels (very) short; connate bracts or bracteoles 1 or 2 (rarely 3 or 4), cupular and calyx-like, at base and/or apex of pedicel, sometimes slightly accrescent towards fruiting stage, persistent. Flowers bisexual, small. Ca-

168

O LACACEAE

lyx fleshy, obsoletely 5- or 6-dentate, small, not or hardly accrescent towards fruiting stage; petals (4)5 or 6(7), inserted on the edge of the disk, valvate, free, triangularlanceolate, fleshy, concave inside in lower part and sheltering the anthers, bearded above the middle inside. Disk hypogynous, free, fleshy, exceeding the calyx slightly, much accrescent after anthesis to cover at least part of the fruit, often almost completely so. Stamens 5 or 6, half as long as petals and inserted in the cavity before them; filaments flat, thickish, short and broad; anthers almost tetragonous, dorsifixed, dehiscent by short vertical rims. Ovary very shortly stipitate, conical, 6– 12-sulcate when dried, attenuate to a short filiform or cylindric style, and an oblong-ovoid, hardly 3-lobed stigma, 2-locular below, 1-locular above. Fruit a pseudodrupe, subtended by 1 or 2 (rarely 3 or 4) calyx-like prophylls and the remains of the calyx, tightly covered for 1/2–4/5 its length (or sometimes completely) by the accrescent pericarp-like disk, free at least at the then umbo-like top; endocarp crustaceous. Colombia, Venezuela, Guyana, French Guiana, Peru, Brazil; 5 species, 1 in Venezuela. Cathedra acuminata (Benth.) Miers, Ann. Mag. Nat. Hist. ser. 3, 4: 361. 1859. —Diplocrater acuminatus Benth. in Hooker’s J. Bot. Kew Gard. Misc. 3: 367. 1851. —Asta blanca, Limoncillo, Yaguaza. Cathedra crassifolia Benth. ex Miers, Ann. Mag. Nat. Hist. ser. 3, 4. 361. 1859. Cathedra caurensis Pittier, Bol. Soc. Venez. Ci. Nat. 6: 5. 1939. Shrub or usually a tree (1.5–)4–15(–25) m tall; petals (4–)6, short-pubescent inside, white (to pale yellowish green); disk ur-

ceolate, 1 mm high at anthesis, later much accrescent and covering all but very apex of fruit; fruit yellow to yellow-orange. Riparian, gallery, flooded, and savanna-edge forests, Mauritia palm swamps, 50–200(–300) m; Delta Amacuro (scattered), Bolívar (Maripa to Aripao, basins of Río Caura, Río Paragua, and Río Orinoco), Amazonas (Río Cuao– Sipapo, Río Cunucunuma, Caño Surumoni west of La Esmeralda, Caño San Miguel). Apure, Distrito Federal, Guárico, Táchira; southeastern Colombia, Guyana, French Guiana, Peru, Brazil. ◆Fig. 165.

Fig. 165. Cathedra acuminata

Chaunochiton 169

4. CHAUNOCHITON Benth. in Benth. & Hook. f., Gen. Pl. 1: 996. 1867. Small to medium-sized trees, glabrous or nearly so. Leaves pinnately-veined. Inflorescence of axillary and/or (sub)terminal shortly-pedunculate few- to manyflowered corymb-like panicles; bracts small. Flowers bisexual; pedicels short. Calyx 5-dentate, small in anthesis, much enlarged towards fruiting stage; petals 5, linearelongate, cucullate-dilated at apex, membranaceous, connate or coherent below, or free, pubescent inside. Stamens 5, slightly shorter than petals and inserted in front of them; filaments filiform, reddish or purplish, adnate to petals at base; anthers subglobose, small, sub-4-lobed, opening by longitudinal slits. Ovary free, oblong, 5ribbed lengthwise, 2-locular below, 1-locular above; style filiform, slightly shorter than petals; stigma capitate, 5-lobed. Disk absent. Drupe globose or subovoid, usually with 5–10(–20) vertical low ribs (when dried) and an equal number of projections or teeth which protrude beyond the somewhat flattened or subtruncate apex, sometimes with a row of warts in place of each rib. Fruiting calyx much accrescent, greenish to red, reddish purple, or brown, either subcoriaceous and funnelform, loosely enveloping the drupe entirely, or papery and enveloping the base of the drupe only, with the remainder wing-like, rotate and spreading horizontally. Seed globose, with a narrow furrow through which the central placenta passes.

Fig. 166. Chaunochiton angustifolium

Fig. 167. Chaunochiton loranthoides

170

O LACACEAE

Rare in Costa Rica and western Colombia, mainly Amazonian Colombia, Venezuela, Guyana, Suriname, French Guiana, and Brazil; 3 species 2 in Venezuela, both in the flora area. Key to the Species of Chaunochiton 1.

1.

Petioles 1–2(–3) mm long; leaf blades 3–5(–6) cm wide, with 6–7 pairs of lateral veins; petals ca. 0.8–1.2 cm long, free; fruiting calyx cupular around base of drupe, otherwise horizontally spreading and wing-like, 3.5-8 cm diameter; drupe ca. 0.5–0.6 cm diameter ......... C. angustifolium Petioles ca. 5 mm long; leaf blades 1.2–2.5(–3.5) cm wide, with 3 or 4 pairs of lateral veins; petals (3–)4–5(–6.5) cm long, including 1–2 cm connate portion at base; fruiting calyx funnelform, enveloping entire drupe, 2–3 cm diameter at edge; drupe ca. 1–1.8 cm diameter ............................................................................................. C. loranthoides

Chaunochiton angustifolium Sleumer, Repert. Sp. Nov. Regni Veg. 42: 261. 1937. —Copito negro. Large shrub to tree (2.5–)3.5–15 m tall; corolla greenish; fruit with 10(20) low or obscure longitudinal ribs. Shrub islands and edges of forests near savannas, granitic outcrop along rivers, 50–300(–700) m; northwestern Bolívar, northwestern and westcentral Amazonas. Apure; Amazonian and west-coastal Colombia, Brazil (Amazonas, Roraima). ◆Fig. 166.

Chaunochiton loranthoides Benth. in Hook., Icon. Pl. 11: pl. 1005. 1867. Bushy tree (2–)4–7(–12) m tall; corolla cream-colored or light yellow; fruit with 5 low longitudinal ribs. Rocky outcrops along rivers, shrub savannas, 50–200 m; central and southwestern Amazonas. Amazonian Colombia, northwestern Brazil. ◆Fig. 167.

5. DULACIA Vell., Fl. Flumin. 32. 1825 [1829]; Fl. Flum. Icon. 1: pl. 78. 1827 [1831]. Liriosma Poepp., Nov. Gen. Sp. Pl. 3: 33. 1843; 3: t. 239. 1841. Shrubs (occasionally scandent) or trees (rarely lianas), often with minute unbranched pubescence, glabrescent. Branches slender, flexuous, often yellowish when dried. Leaves entire, pinnately-veined, short-petiolate. Inflorescence an axillary simple or laxly composed (cymoid) raceme; rachis straight to zigzag; pedicels short; bracteoles subpersistent. Flowers bisexual, heterostylous, generally rather few per raceme. Calyx small, the edge obsoletely and obtusely 5-lobed or -undulate, or truncate, narrowed towards base and continuous with pedicel, i.e. forming an hypanthium with the latter, much accrescent towards fruiting stage and finally covering the fruit almost entirely; petals 6, slightly perigynous, valvate, connate ± halfway, apex inflexed. Fertile stamens 3, each inserted on the fused bases of 2 petals (alternate with corolla lobes); filaments flat, adnate to basal part of petals, usually pubescent; anthers oblong, dorsifixed, longitudinally deshicent. Staminodes 6, epiand antepetalous, spathulate, usually pubescent below, bifid and glabrous in the upper half. Disk hypogynous, cup-shaped and adnate to calyx, free only at the upper margin (on which petals are inserted), hardly noticeable and not accrescent. Ovary semi-inferior, i.e. immersed halfway in the disk, 3-locular below, 1-locular above; style subcylindric, rather slender, either short and ending at base of anthers, or

Dulacia 171

elongate and exceeding anthers somewhat; stigma subcapitate, slightly 3-lobed in the short-styled, more distinctly so in long-styled flowers. Fruit a pseudo-drupe, covered by the enlarged thin-fleshy calyx except the free umbonate top of ovary (drupe), and style-base; endocarp woody to crustaceous. Cotyledons ovate. Amazonian Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, and Bolivia; 13 species, 8 in Venezuela, all of these in the flora area. Distyly is common in this genus, and most species are known to have both longstyled and short-styled morphs. In other species, only one morph is known. Fruits are generally produced on the long-styled morph, rarely on the short-styled morph, which may be functionally staminate. Key to the Species of Dulacia 1. 1. 2(1).

2.

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

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

7.

Low shrub 0.1–0.7(–1.5) m tall; leaves 1.3–3.3 × 0.5–1.3 cm ..... D. redmondii Shrub to tree, rarely liana, (1.5–)2–20(–30) m tall; leaves much larger ................................................................................................................ 2 Leaves thick-coriaceous, ± stiff, the apex shortly attenuate to rounded, the very apex obtuse, rounded, or slightly emarginate; at elevations of (600–)1000–2000 m ................................................................................ 3 Leaves membranaceous, chartaceous, or only subcoriaceous, ± flexible, the apex acute to long-acuminate; in lowlands ca. 100–200 m elevation ................................................................................................................ 4 Leaves with 4 or 5 pairs of lateral veins; petiole 2–4 mm long ................................................................................................. D. tepuiensis Leaves with 6–10 pairs of lateral veins; petiole ca. 5 mm long ...... D. crassa Free part of ovary and the umbonate apex of drupe glabrous or nearly so; petals glabrous outside; pedicels glabrous at least distally ................ 5 Free part of ovary and the umbonate apex of drupe pubescent, often densely so; petals minutely papillate-puberulent to puberulent outside; pedicels papillate-puberulent to densely puberulent .................. 6 Leaf blades 2–2.7(–2.8) times longer than wide, generally shorter than 9 cm; staminodes pubescent below ........................................ D. inopiflora Leaf blades (2.7–)2.9–4 times longer than wide, generally longer than 9 cm; staminodes glabrous ................................................. D. macrophylla Liana or scandent shrub; fruits obovoid, blue-green ............... D. cyanocarpa Shrub or tree; fruits oblong, widely ellipsoid, ovoid, or obovoid, yellow to orange ..................................................................................................... 7 Free part of ovary conical-pyramidal or narrowly conical at anthesis, gradually attenuate to the style; fruit ellipsoid, widely ellipsoid, or slightly ovoid (i.e., widest at middle or below); leaves conspicuously glaucous-papillose beneath (usually lost in drying) ............. D. guianensis Free part of ovary hemispherical to broadly conical at anthesis, ± abruptly narrowed to the style; fruit oblong, widely ellipsoid, or slightly obovoid (i.e., widest at or above middle); leaves glaucous or not on lower surface ......................................................................... D. candida

172

O LACACEAE

Dulacia candida (Poepp.) Kuntze, Rev. Gen. Pl. 1: 111. 1891. —Liriosma candida Poepp., Nov. Gen. Sp. Pl. 3: t. 239. 1841. —Huevillo de agua. Dulacia adhaerens var. stenopoda Cuatrec., Brittonia 11: 171. 1959. Shrub to tree (0.5–)2–10 m tall; flowers whitish to light yellow; fruit yellow to orange, malodorous. Gallery forests, savannas, 50– 200(–600) m; Bolívar (Río Orinoco, Río Parguaza), scattered in western Amazonas. Anzoátegui, Táchira-Apure border; Colombia, Ecuador, eastern Peru, Brazil, Bolivia. ◆Fig. 168. A specimen from the slopes of Cerro Aracamuni (Liesner 22358, MO) differs in its smaller leaves and abaxially glabrous corolla. Dulacia crassa (Monach.) Sleumer, Fl. Neotr. 38: 120. 1984. —Liriosma crassa Monach., Brittonia 13: 114, fig. 1. 1961. Small tree to 10 m tall; inflorescences ca. 10-flowered, glabrous; petals ca. 10 mm long; fruit unknown. Not yet known from flora area, but expected in Bolívar. Guyana (southern slope of Roraima, Kaieteur savanna). Dulacia cyanocarpa Sleumer, Fl. Neotr. 38: 130. 1984. Scandent shrub or liana, ca. 3 m tall; corolla ca. 7 mm long, whitish; fruit blue-green. Gallery forest, 50–100 m; Amazonas (near Puerto Ayacucho). Apure. There is some question whether the fruit remains bluish at maturity. Dulacia guianensis (Engl.) Kuntze, Revis. Gen. Pl. 1: 111. 1891. —Liriosma guianensis Engl. in Mart., Fl. Bras. 12(2): 25. 1872. —Burro muerto. Tree (4–)6–25(–30) m tall; sap from bark or stem malodorous; leaves glaucous beneath; flowers white to pale green or cream; fruit yellow to orange, malodorous. Evergreen lowland and lower montane forests, 100–300 m; Bolívar (near El Dorado), Amazonas (Cerro Sipapo, Río Guayapo, Río Parú). Apure, Miranda; Guyana, Suriname, French Guiana, Brazil (Amazonas, Goiás/ Pará, Maranhão). ◆Fig. 171. The orange fruit of Dulacia guianensis is said to be edible.

Dulacia inopiflora (Miers) Kuntze, Revis. Gen. Pl. 1: 111. 1891. —Liriosma inopiflora Miers, Ann. Mag. Nat. Hist. ser. 3, 4: 362. 1859. Liriosma micrantha Spruce ex Engl. in Mart., Fl. Bras. 12(2): 28. 1872, nom. nudum. Shrubby tree or tree (3–)4–15(–18) m tall; corolla ca. 3-4 mm long, pale green or yellowish; fruit yellow to orange. Understory of swampy or seasonally flooded forests, gallery forests, 50–200 m; Amazonas (Río Casiquiare basin, La Esmeralda). Colombia, Peru, Brazil. ◆Fig. 169. Dulacia macrophylla (Benth.) Kuntze, Revis. Gen. Pl. 1: 111. 1891. —Olax macrophylla Benth., Proc. Linn. Soc. London 1: 89. 1841; Trans. Linn. Soc. London 18: 678. 1841. —Liriosma macrophylla (Benth.) Roem. in M.R. Schomb., Reis. Br.-Guiana 3: 1095. 1848 [1849]. —Cupi fino. Shrub or small tree 2–3 m tall; leaves usually long-acuminate; flowers white; fruit yellow to orange. Edges of gallery and seasonally flooded forest, or tall nonflooded forest, 50–200 m; Amazonas (upper Río Negro, upper Río Orinoco, Río Casiquiare). Táchira; Colombia, Peru, Brazil (Amazonas). ◆Fig. 170. Dulacia redmondii Steyerm., Phytologia 38: 217, fig. 1. 1978. Prostrate or usually erect, single-stemmed or sparsely branched, dwarf shrub 0.1–0.7 (–1.5) m tall; leaves dropping, conspicuously wrinkling when dried; flowers pale green to yellow-green; fruit orange to red. Open savannas or shrub islands, often on white sand, 50–200 m; western Amazonas (Río Atabapo, Río Orinoco, Río Ventuari). Eastern and southeastern Colombia, southern Guyana, Brazil (Amazonas). ◆Fig. 173. Dulacia tepuiensis (Steyerm.) Sleumer & Steyerm. in Sleumer, Fl. Neotr. 38: 119. 1984. —Liriosma tepuiensis Steyerm., Bol. Soc. Venez. Ci. Nat. 32: 328, fig. 11. 1976. Tree 3–8 m tall; leaves with secondary and tertiary veins deeply impressed in upper surface. Dwarf forest on tepui summits, (600–) 1300–2000 m; Bolívar (Cerro Guanacoco, Cerro Sarisariñama, Macizo del Chimantá [Apácara-tepui], Sierra Ichún). ◆Fig. 172.

Dulacia 173

Fig. 168. Dulacia candida

Fig. 169. Dulacia inopiflora

Fig. 170. Dulacia macrophylla

174

O LACACEAE

Fig. 171. Dulacia guianensis

Fig. 172. Dulacia tepuiensis

Fig.173. Dulacia redmondii

Heisteria 175

6. HEISTERIA Jacq., Enum. Syst. Pl. 4, 20. 1760, nom. cons. Rhaptostylum Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 2: 139, t. 125. 1813. Shrubs (rarely scandent) or trees (or rarely lianas) with slender branchlets, glabrous or nearly so. Leaves entire, always glabrous, the margin cartilaginous and often a little revolute; petioles markedly canaliculate; blades sometimes wrinkled and/or granular-tuberculate on one or both surfaces by idioblasts showing through the epidermis in dry leaves, usually rather densely pellucid-micropunctulate against strong light, the discolorous laticifers generally visible beneath with the naked eye, pinnately-veined, sometimes the basal 1(2) pairs strong and ascending (in the flora area only H. scandens). Inflorescence of axillary cushions or glomerules, few- to many-flowered, few- to multibracteolate. Flowers bisexual, small, sessile or pedicellate. Calyx small, 5(rarely 6)-dentate or -lobed, the lobes sometimes with a minute apical gland, usually much accrescent and brightly colored towards fruiting stage, very rarely remaining practically unaltered and pale; petals 5 (very rarely 6), free or a little coherent at base, valvate, glabrous or usually pubescent inside, whitish, pale green, to pale yellow. Stamens usually 10 (rarely 12), the 5(6) outer ones a little longer and antesepalous, the 5(6) inner ones a little shorter and antepetalous, exceptionally 5(6) by abortion and then all antesepalous (in the flora area, only H. pentandra); filaments adnate below to base of petals, free otherwise, ligulate or filiform; anthers small, subglobose, dehiscing lengthwise, basifixed. Disk completely adnate to the lower 10-sulcate part of ovary. Ovary superior, 3-locular below, 1-locular above; style short-conical; stigma shortly 3-lobed. Drupe globose, ellipsoid, oblong, (or barrel-shaped outside the flora area); pericarp thin-fleshy, whitish or yellowish to red or purplish-blackish, in the dry state smooth or tuberculate by pustular excretions of the crustaceous endocarp which show through the pericarp, subtended or included at full maturity by the generally much accrescent thin-fleshy to coriaceous, white, gray, green, yellowish, deep red to purplish, hardly to deeply 5(6)-lobed fruit-calyx. Cotyledons orbicular, foliaceous. Mainly Neotropics, 3 species in Africa; ca. 33 species, 13 in Venezuela, 11 of these in the flora area. Specimens of Heisteria can often only be named with certainty if they bear leaves and nearly mature fruits. The flowers in Heisteria, except for the number of stamens, are relatively homogeneous and offer few characters for the discrimination of species. In the key, the diameter of the fruiting calyx refers to the size when expanded or spread out. Key to the Species of Heisteria based on fruiting specimens 1.

1.

2(1).

Fruiting calyx 3–6(–7) mm diameter, 1.5–4 mm long, a small erect closefitting cup, entire or nearly so, only slightly accrescent, concealing only the base of drupe; fruiting pedicel 1–4 mm long; drupe yellow to reddish orange at maturity ................................................... H. densifrons Fruiting calyx either ca. 8 mm diameter and reflexed, or else larger and spreading, reflexed, erect, or cupular, (entire to) shallowly to deeply 5lobed, not concealing drupe or loosely but nearly completely enveloping it; fruiting pedicel 1–20 mm long; drupe white, gray, light bluegray, yellow, orange, red, dark green, or blackish at maturity ............ 2 Fruiting calyx reflexed or subhorizontally spreading, never plicate basally, not enveloping or concealing the drupe, entire to shallowly lobed

176

O LACACEAE

or undulate, 6–20 mm diameter, greenish or red; fruiting pedicel 8– 15(–20) mm long; drupe orange, red, bluish-black, or black, but never white ....................................................................................................... 3 2. Fruiting calyx erect, lobed-cupular, bowl-shaped, or shallowly bowlshaped and spreading, sometimes plicate basally, partially enveloping or loosely concealing the drupe, moderately to deeply 5-lobed, (10–)12– 70 mm diameter, red (sometimes green in H. maytenoides); fruiting pedicel 1–10(–15) mm long; drupe white, gray, pale blue-gray, yellow, orange, or red ......................................................................................... 6 3(2). Fruiting calyx green at maturity .............................................................. 4 3. Fruiting calyx red at maturity (rarely sometimes green in H. pentandra?) ................................................................................................................ 5 4(3). Liana (rarely scandent shrub); leaves often with basal pair of lateral veins prominent and strongly ascending (the leaf sometimes almost pliveined); drupe orange to red ............................................... H. scandens 4. Small to large erect tree; leaves without prominent pair of basal lateral veins; drupe blackish ................................................................. H. barbata 5(3). Erect shrub or tree; leaves with 6–10(–12) pairs of lateral veins; fruiting calyx 10–18 mm diameter; drupe blackish at maturity; widespread in Delta Amacuro, Bolívar, and Amazonas ............................... H. acuminata 5. Scandent shrub or slender treelet; leaves with 5 or 6(–9) pairs of lateral veins; fruiting calyx poorly known, ca. 8 mm diameter?; drupe red (or black?) at maturity; rare in Amazonas ................................. H. pentandra 6(2). Leaves longer than 18 cm; lateral veins 12–18 pairs, deeply impressed in upper surface of leaf; drupe whitish or light gray .................. H. insculpta 6. Leaves shorter than 15 cm and lateral veins 6–10(–12) pairs, or if some leaves to 20 cm then the lateral veins 6–9 pairs; lateral veins impressed or ± flat on upper surface of leaf; drupe whitish, gray, pale blue-gray, yellow, orange, or red ........................................................... 7 7(6). Fruiting calyx 10–20(–25) mm diameter, red or greenish; drupe yellow, orange, or red ......................................................................................... 8 7. Fruiting calyx 25–60(–70) mm diameter, red; drupe white, light gray, or pale blue-gray ........................................................................................ 9 8(7). Leaves drying with noticeably raised laticifers on lower surface, the apex shortly-, rarely more elongately-attenuate or subacuminate, only the very tip often subacute or blunt; fruiting pedicels 4–8(–15) mm long ........................................................................................................ H. ovata 8. Leaves drying with flat or poorly visible laticifers on lower surface, the apex acute to broadly obtuse; fruiting pedicels 3–6 mm long ............................................................................................. H. maytenoides 9(7). Fruiting calyx 25–30(–35) mm diameter, thin-chartaceous; leaf apex usually gradually attenuate or subacuminate ..................... H. spruceana 9. Fruiting calyx (36–)40–60(–70) mm diameter, fleshy or drying thick; leaf apex usually abruptly acuminate, the acumen 3–10 mm long .......... 10 10(9). Leaves thick-coriaceous, mostly 5–8 cm wide, drying conspicuously and equally wrinkled and tuberculate on both surfaces, the laticifers not discolorous beneath; fruiting calyx 40–60(–70) mm diameter .................................................................................................. H. maguirei

Heisteria 177

10.

Leaves subcoriaceous or thin-coriaceous, mostly 3–5 cm wide, drying smooth or tuberculate but not particularly wrinkled, often smoother above, the laticifers often conspicuously discolorous beneath; fruit calyx (35–)40–50 mm diameter .................................................... H. duckei

Heisteria acuminata (Bonpl.) Engl. in Mart., Fl. Bras. 12(2): 14. 1872. —Rhaptostylum acuminatum Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 2: 139, t. 125. 1813. —Cafecito, Limoncillo, Pasita, Pasita hoja fina. Heisteria longipes Standl., J. Wash. Acad. Sci. 17: 8. 1927. Heisteria guianensis V.M. Badillo in Pittier et al., Cat. Fl. Venez. 1: 272. 1945, “guayanensis,” non Engl. 1872. Shrub to small (rarely large) tree (1.5–)3– 10(–20) m tall, the bark gray, cracking in squarish plates on larger trees; immature drupe red, blue-black when mature. Gallery forests, lower montane forests, 100–200 (–800) m; Delta Amacuro (Caño Orocoima, Río Acure), Bolívar (lower Río Caura), Amazonas (Río Atabapo, upper Río Orinoco basin, Sierra Parima). Widespread elsewhere in Venezuela; Nicaragua, Costa Rica, Panama, Colombia, Amazonian Brazil, Bolivia. ◆Fig. 175. Heisteria barbata Cuatrec., Trop. Woods 101: 26. 1955. —Pasita hoja fina. Tree 3–15(–30) m tall. Nonflooded evergreen lowland forests, 50–300 m; Amazonas (Río Cunucunuma). Amazonian Colombia, French Guiana, Peru, Brazil. ◆Fig. 177. Heisteria densifrons Engl. in Mart., Fl. Bras. 12(2): 17, t. 5, f. 1. 1872. Heisteria microcarpa Spruce ex Engl. in Mart., Fl. Bras. 12(2): 21. 1872. Subscandent shrub to tree 3–10(–16) m tall; petals 4 or 5(6) mm long. Gallery forests and seasonally flooded forests, 50–100 m; Amazonas (basin of Río Casiquiare). Guyana, Suriname, French Guiana, Peru, Amazonian Brazil. Heisteria duckei Sleumer, Repert. Spec. Nov. Regni Veg. 38: 207. 1935. Tree 6–20(–35) m tall. Evergreen lowland to lower montane forests, 100–400 m; Amazonas (Cerro Yapacana). Colombia, Peru, Brazil, Bolivia. ◆Fig. 178.

Heisteria insculpta Sleumer, Notizbl. Bot. Gart. Berlin-Dahlem 12: 65. 1934. Shrub or usually small tree (2–)3–7 m tall, often cauliflorous; drupe white; fruiting calyx red, 4–7 cm diameter. Riparian forests and lower montane forests, 100–300 m; Amazonas (Río Mawarinuma at base of Sierra de la Neblina, Río Pasimoni basin), Peru, Brazil (Amazonas). ◆Fig. 174. Heisteria maguirei Sleumer, Fl. Neotr. 38: 68. 1984. —Corona de rey. Tree (4–)10–20(–26) m tall. Evergreen lowland forests, ca. 200 m; border of Delta Amacuro and Bolívar (Serranía de Imataca). Guyana, Brazil (Amapá). Heisteria maytenoides Spruce ex Engl. in Mart., Fl. Bras. 12(2): 15. 1872. —Palo de panema. Tree 5–9 m tall. Seasonally flooded forests along black-water rivers, 100–200 m; Amazonas (Caño Cupavén al norte de San Fernando de Atabapo, Río Cuao, Río Guainía, Río Sipapo, San Carlos de Río Negro). Colombia, Peru, Brazil. ◆Fig. 179. Heisteria ovata Benth. in Hooker’s J. Bot. Kew Gard. Misc. 3: 366. 1851. —Corona de rey, Tue-becok. Heisteria flexuosa Engl. in Mart., Fl. Bras. 12(2): 17, t. 4, fig. 3. 1872. Heisteria nitida auct. non Spruce ex Engl.: sensu V.M. Badillo in Pittier et al., Cat. Fl. Venez. 1: 272. 1945. Shrub to tree (2.5–)6–20(–25) m tall. Gallery forests, rainforests, mixed premontane forests, 100–300 m; Bolívar (Río Asa, Río Parguaza, Río Suapure), Amazonas (Cerro Sipapo, near Puerto Ayacucho). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 180. Heisteria pentandra (Benth. ex Reisseck) Engl. in Mart., Fl. Bras. 12(2): 20. 1872. —Rhaptostylum pentandrum Benth. ex Reisseck in Mart., Fl. Bras. 11(1): 77, t. 4, fig. 19. 1861.

178

O LACACEAE

Fig. 174. Heisteria insculpta

Fig. 175. Heisteria acuminata

Heisteria 179

Fig. 176. Heisteria spruceana

Fig. 177. Heisteria barbata

180

O LACACEAE

Fig. 178. Heisteria duckei

Fig. 179. Heisteria maytenoides

Fig. 180. Heisteria ovata

Minquartia 181

Heisteria acuta Engl. in Mart., Fl. Bras. 12(2): 21. 1872. Heisteria flexuosa auct. non Engl. 1892: sensu V.M. Badillo in Pittier et al., Cat. Fl. Venez. 1: 272. 1945, pro parte, non Engl. Scandent shrub or treelet ca. 2.5 m tall; stamens 5, antesepalous; mature fruit unknown. Evergreen lowland forests, 50–200 m; Amazonas (Isla Solitario in Río Orinoco, Río Casiquiare, Río Mawarinuma). Amazonian Brazil. Heisteria pentandra is the only species with five stamens in the flora area. Heisteria scandens Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 9. 1925. Heisteria fatoensis Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 137. 1930.

Liana 10–30 m, or rarely a scandent shrub 2–4 m tall. Riparian and lower montane forests, 100–800 m; Bolívar (58 km southeast of Los Pijiguaos, Río Arisa, upper Río Caroní, upper Río Paragua), Amazonas (Río Mawarinuma below Sierra de la Neblina, Río Jénita in lower Río Ocamo). Costa Rica, Panama, western and southeastern Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil, and Bolivia. Heisteria spruceana Engl. in Mart., Fl. Bras. 12(2): 15. 1872. Shrub or small tree 2–10(–15) m tall. Riparian forests, 50–200 m; Amazonas (Río Cataniapo near Las Pavas, Río Orinoco near Caño Yapacana). Colombia, Peru, Brazil. ◆Fig. 176.

7. MINQUARTIA Aubl., Hist. Pl. Guiane 4, t. 370. 1775. Trees with milky juice in bark and leaves; trunk usually deeply grooved or perforated; young parts covered with a rusty tomentum of minute branched trichomes, glabrescent (rarely initially glabrous outside of flora area). Leaves entire, usually retaining a minute branched pubescence on the veins beneath, pinnately-veined with (8–)10–14(–17) pairs of veins, the tertiary veins subparallel, with schizogenous secretory cavities with resinous contents and laticifers, often with spicular cells. Inflorescences axillary, spicate; spikes pedunculate, rather short, generally simple, rarely few- and short-branched, many-flowered. Flowers bisexual, hypogynous, small, subsessile, 2–5-fascicled along rachis, each flower subtended by a minute ovate caducous bract. Calyx 5(6)-denticulate, small, persistent, not accrescent; petals (4)5 or 6(7), valvate, fleshy, campanulate, connate in the lower half, ovate-oblong, small. Stamens generally 10 (5 antesepalous and 5 antepetalous), sometimes 15 (5 antesepalous and 10 antepetalous), alternately slightly unequal; filaments filiform, adherent to petals basally; anthers subquadrate-suborbicular, 4-locular, dehiscent lengthwise. Ovary (3)4(5)-locular below, 1-locular above; style very short; stigma sessile, shortly 3–5-lobed. Drupe with thin-fleshy finally dissolved exocarp; endocarp crustaceous, tuberculate outside, the tubercles showing through the exocarp in dry submature fruits. Nicaragua, Costa Rica, Panama, Amazonian Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 1 species. Minquartia guianensis Aubl., Hist. Pl. Guiane 4, t. 370. 1775. —Cuyuvi, Palo de cuyubi, Urana-u-yek (Arekuna). Tree 4–30 m tall; sapwood yellow; spikes solitary, simple or rarely few-branched; petals connate 1–1.5 mm, free 1–1.5 mm, pubescent within; fruit purplish. Lowland primary rainforests, premontane wet forests, lower slopes of tepuis, 100–1200 m; Bolívar (Cerro Uei, Cerro Venamo, Gran Sabana, Macizo del

Chimantá [Torono-tepui]), Amazonas (El Porvenir, Río Siapa near mouth of Caño Chimoni, near San Carlos de Río Negro, Santa Bárbara de Orinoco, 15 km southwest of Tamatama). Nicaragua, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 181. The heavy wood of Minquartia guianensis is resistant to rot and termites and is prized in construction for framing poles.

182

O LACACEAE

Fig. 181. Minquartia guianensis

8. SCHOEPFIA Schreb., Gen. Pl. 1: 129. 1789. Shrubs or low trees, glabrous or nearly so, reportedly root-parasites. Leaves entire, pinnately veined. Inflorescence axillary, of short, fascicled spikes; base of inflorescence (peduncle) with a few small imbricate persistent scale-like bracts; bract and 2 bracteoles at apex of pedicel ± united into a small acutely 3-lobed epicalyx (the pedicel nearly absent to 1.5 mm long in the flora area). Flowers bisexual, often dimorphous and heterostylous, small. Calyx inconspicuous, i.e., adnate to the cup-shaped truncate flower axis; petals (3)4 or 5(6), inserted on the edge of flower axis, fused in the lower 1/2–2/3 into a tube which often is cylindric in the short-styled form and urceolate to subcampanulate in the long-styled form, free above in the form of valvate and revolute lobes, with a tuft of trichomes inside the tube behind each anther. Stamens as many as petals, epipetalous and opposite the corolla lobes, the slender filaments adnate to corolla tube for nearly their full length, free at apex; anthers free, 2-locular, attached below the middle, sometimes smaller in shortstyled flowers. Disk epigynous, annular, fleshy. Upper half of ovary superior, included by the disk, lower half within the flower axis, 3-locular below, 1-locular above; style slender, either as long as corolla tube and with a well-developed stigma in the long-styled form, or half as long in the short-styled form, the stigma then a little below the anthers and smaller. Fruit drupaceous, subtended by the persistent epicalyx, crowned by remains of the calyx and disk; epicarp originating from the somewhat accrescent flower axis, thin, fleshy; endocarp parchment-like to crustaceous, striate lengthwise. Neotropics, Asia, Malesia; 25 species, 5 in Venezuela, 4 of these in the flora area.

Schoepfia 183

Fig. 182. Schoepfia brasiliensis

Fig. 183. Schoepfia tepuiensis

Fig. 184. Schoepfia lucida

184

O LACACEAE

Key to the Species of Schoepfia 1.

1. 2(1).

2.

3(1).

3.

Inflorescence spike-like, 5–8-flowered, with flowers scattered or in groups of 2 or 3 along rachis, 5–15 mm long from base of peduncle to end of rachis ...................................................................................................... 2 Inflorescence of 2 or 3 flowers at or near apex of short (2–10 mm long) peduncle ..................................................................................................... 3 Leaves ca. 1.2–1.5 cm wide; corolla ca. 1.8–2 mm long; bracts and bracteoles of epicalyx connate more than halfway, forming a 3-lobed or -cusped involucre; leaf blades with the lowermost 1 or 2 pairs of lateral veins not notably conspicuous and extending strongly to or past the middle of blade .............................................................................. S. clarkii Leaves (2.5–)3–5(–6) cm wide; corolla ca. 4–5 mm long; bracts and bracteoles nearly free or connate near base only; leaf blades with the lowermost 1 or 2 pairs of lateral veins very conspicuously and strongly extending to or past the middle of blade ....................................... S. lucida Leaves 1.2–2(–3) cm wide; inflorescence 2- or 3-flowered, the peduncle 2– 3(–5) mm long; corolla 5–5.5(–6) mm long; habitat elevation 1900– 2200 m ......................................................................................S. tepuiensis Leaves (1.5–)2–4(–5) cm wide; inflorescence (2)3–5-flowered, the peduncle (3–)5–10 mm long; corolla 5–6.5(–7) mm long; habitat elevation < 1500 m ................................................................................ S. brasiliensis

Schoepfia brasiliensis A. DC., Prodr. 14: 622. 1857. Large shrub or small tree 4–8 m tall; corolla pale yellow to yellow, sometimes reddish; fruit blood-red. Lower montane forests, forest-savanna ecotones, 400–1400 m; Bolívar (Altiplanicie de Nuria, Gran Sabana). Anzoáteguí, Monagas; Brazil (mainly coastal). ◆Fig. 182. Schoepfia clarkii Steyerm., Ann. Missouri Bot. Gard. 75: 1061. 1988. Thin shrub ca. 2 m tall; corolla greenish yellow; fruit unknown. Low Amazonian caatinga on white sand, ca. 100 m; rare in Amazonas (near San Carlos de Río Negro). Endemic.

Schoepfia lucida Pulle, Recueil Trav. Bot. Néerl. 9: 136. 1912. Shrub or tree 3–25 m tall; corolla pale green or greenish yellow; fruit dark red or purplish red. Lower montane forests, ca. 800 m; Amazonas (Sierra Parima). Brazil (Mato Grosso). ◆Fig. 184. Schoepfia tepuiensis Steyerm., Bol. Soc. Venez. Ci. Nat. 22: 331, fig. 12. 1976. Shrub or shrubby tree (1.5–)2–4(–8) m tall; corolla dark red, sometimes apically yellow or greenish yellow; fruit dark red. Along streams, savanna edges, gallery forest, rocky areas of tepui summits, 1900–2200 m; Bolívar (Auyán-tepui, Macizo del Chimantá). Endemic. ◆Fig. 183.

9. XIMENIA Plum. ex L., Sp. Pl. 1193. 1753. Root-parasitic shrubs or low trees; branches usually armed with axillary spines and/or ending in thorns. Leaves sometimes fascicled on short lateral twigs, entire, pinnately-veined, usually ending with a short mucro. Inflorescences axillary, or at the end of a short lateral twig (which may be reduced to a thorn), arranged in rather few-flowered pedunculate, sometimes umbel-like cymes, or in fascicles, rarely solitary; bracteoles 2–4 near base of pedicel, or ebracteolate. Flowers usually bisexual, rarely functionally unisexual (not known if plants monoecious or dioecious then). Calyx small, cupular-expanded, with (3)4(5) teeth or lobes, persistent, hardly or not

Ximenia 185

accrescent in fruit; petals 4 (rarely 5), valvate, free, linear-oblong, finally revolute about halfway, with a brush of trichomes inside. Stamens free, hypogynous, rarely 4(5) and antepetalous, or usually 8 (rarely 10) with half antepetalous and the other half antesepalous; filaments filiform; anthers oblong-linear to subovate, basifixed, deshicent lengthwise (not shedding pollen in the functionally pistillate flowers); connective thick. Disk absent. Ovary generally smaller and of slightly different shape in functionally staminate flowers, superior, sessile, (3)4-locular; style columnar, sometimes very short or even absent in functionally unisexual flowers; stigma capitate, small. Fruit drupaceous; pericarp (sarcocarp) rather thin, pulpy; endocarp crustaceous to woody. Tropics and subtropics worldwide (mostly Neotropics); 10 species, 1 in Venezuela. Ximenia americana L., Sp. Pl. 1193. 1753. Shrub to tree 1–8(–12) m tall; usually deciduous; branchlets with spines or thorns. Deciduous or semideciduous lowland forests, 100–300 m; Bolívar (islands in Lago Guri). Scattered through much of northern Venezuela; pantropical and subtropical. There are at least three varieties, var. argentinensis DeFilipps being restricted to Bolivia, Paraguay, and Argentina, and var. microphylla Welw. ex Oliv. to Africa.

Fig. 185. Ximenia americana var. americana

186

O LACACEAE

X. americana var. americana. —Limoncillo. Flowers light yellow-green or cream; petals 4(5), the inner surface conspicuously long-bearded except apex; fruits yellow-orange. Margins of forests in usually dry savannas, lajas, rocky hillsides, 50–400 m;

Bolívar (Altiplanicie de Nuria, Cerro San Borja), Amazonas (near Puerto Ayacucho). Anzoátegui, Aragua, Carabobo, Distrito Federal, Falcón, Lara, Miranda, Nueva Esparta, Sucre; pantropical and subtropical. ◆Fig. 185.

OLEACEAE by James S. Miller and Gerardo A. Aymard C. Trees, shrubs, or vines, evergreen or deciduous, the stems often prominently lenticellate and often with a well developed pith. Leaves opposite or rarely alternate, simple or pinnately compound or trifoliolate, entire, lobed, or serrate, exstipulate. Inflorescences mostly terminal, dichasial panicles or cymes, rarely solitary, bracteate. Flowers bisexual, or some or all unisexual, the plants then dioecious, actinomorphic, pedicellate, often fragrant. Calyx usually small, cupulate, 4–8-lobed, the lobes valvate. Petals connate into a tube or free (some Fraxinus) or rarely lacking, the corolla 4(–12)-lobed, the lobes imbricate or induplicate valvate. Stamens 2(4), alternate with the corolla lobes; filaments adnate to the corolla tube; anthers bithecal, back to back, dehiscing longitudinally, the connective broad, apiculate. Ovary superior, 2(4) carpellate, 2(4) locular, the placentation axile; style terminal, bifid with 2 deltoid to fusiform stigmas or simple with a capitate stigma, or rarely with the stigma sessile; ovules (1)2(3-numerous) per locule, ascending or pendulous, anatropous or amphitropous. Fruits capsular, samaroid, or drupaceous, 1–4-seeded; seeds usually with abundant, oily endosperm. Worldwide (most diverse in Asia and Malaysia); ca. 30 genera and 400 species, 1 genus and 2 species in the flora area. Many Oleaceae are valued as cultivated plants. Olea eruopea L. (Olive, Aceituno) is cultivated for its edible fruits. Species of Fraxinus (Ash, Fresno) are valued as timber or shade trees. Many species of Jasminum, Ligustrum, Osmanthus, Forsythia, Syringa, Chionanthus, and Noronhia are cultivated for showy, fragrant flowers on attractive foliage. 1. CHIONANTHUS L., Sp. Pl. 8. 1753. Mayepea Aubl., Hist. Pl. Guiane 1: 81. 1775. Thouinea Thunb. ex L. f., Suppl. Pl. 89. 1781. Linociera Schreb., Gen. Pl. 2: 784. 1791. Trees, evergreen or deciduous, the twigs often prominently lenticellate. Leaves opposite, simple, entire, often punctate below, usually petiolate, exstipulate. Inflorescences usually terminal, paniculate with opposite branching, bracteate. Flowers usually bisexual, actinomorphic, pedicellate, fragrant. Calyx cupulate, small, shortly 4-lobed. Corolla white to pale yellow, basally connate into a tube, the 4 lobes linear, usually elongate and forming more than half the length of the corolla. Stamens 2, rarely 4, alternating with the corolla lobes; filaments adnate to the corolla tube; anthers bithecal, oblong, with a broad, apiculate connective. Ovary 2-carpellate, 2-locular; style short, 2-lobed, the stigmas ventral; ovules 2 per locule, pendulous. Fruit a single-seeded drupe.

Chionanthus 187

Tropical and north-temperate areas of both hemispheres; ca. 100 species, 5 in Venezuela, 2 of these in the flora area. Most of the tropical species have more commonly been treated in the genera Mayepea and Linociera, but Stearn (Ann. Missouri Bot. Gard. 63: 355–357. 1977) has shown that these genera are synonymous with Chionanthus. Green (Kew Bull. 49: 261–286. 1994) agreed with this and updated all South American species. Key to the Species of Chionanthus 1.

1.

Inflorescences axillary at the end of branches, 5–10 cm long, open cymosepaniculate, flowers numerous; calyx lobes puberulent, especially at the margins .................................................................................. C. compactus Inflorescences axillary, 1–4 cm long, dense cymose-paniculate, flowers usually 11–15; calyx lobes densely strigose-pubescent ........ C. implicatus

Chionanthus compactus Sw., Prodr. 13. 1788. Chionanthus caribea Jacq., Collectanea 2: 110. 1788 [1789]. —Mayepea caribea (Jacq.) Kuntze, Revis. Gen. Pl. 2: 411. 1891. —Linociera caribea (Jacq.) Knobl., Bot. Centralbl. 61: 84. 1895. Tree to 10 m tall; flowers white, fragrant. Lower montane forests, granitic outcrops, 100–600 m; Bolívar (Altiplanicie de Nuria), Amazonas (Cerro Ucata southeast of Síquita). Aragua, Distrito Federal, Falcón, Nueva Esparta, Sucre, Táchira, Trujillo, Yaracuy; Lesser Antilles, Colombia. ◆Fig. 186. Chionanthus implicatus (Rusby) P.S. Green, Kew Bull. 49: 270. 1993. —Mayapea implicata Rusby, Bull. New York Bot. Gard. 4: 314. 1907. Tree to 12 m tall; flowers white. Evergreen lowland forests, 100–200 m; Amazonas (Caño Yureba, Río Sipapo, Río Ventuari). Apure; Colombia, Ecuador, Peru, Brazil, Bolivia.

Fig. 186. Chionanthus compactus

188

O NAGRACEAE

ONAGRACEAE by Elsa M. Zardini and Peter H. Raven Herbs, shrubs, lianas, or rarely trees. Leaves alternate, opposite, or rarely whorled, simple; blades entire, lobed, or pinnatifid; stipules minute or absent. Flowers actinomorphic or zygomorphic, bisexual (sometimes male-sterile in Fuchsia), 4 or 5(rarely 2-, 3-, or 6)-merous, borne in axils of usually reduced foliage leaves or in a more or less distinct inflorescence, usually a corymb or raceme. Floral tube (extension of hypanthium beyond ovary) often present (absent in Ludwigia). Sepals free or rarely connate; petals free or very rarely fused to sepals, sometimes absent. Stamens usually twice as many as sepals and in 2 whorls, or else as many as sepals or reduced to 2; anthers versatile, sometimes attached near base, dehiscing longitudinally. Ovary inferior; style 1; stigma variously lobed, discoid, elongate, or capitate. Fruit a loculicidally or more rarely irregularly dehiscent capsule, sometimes indehiscent, or a berry, 1–many-seeded; placentation axile. Seeds small, anatropous, lacking endosperm. Cosmopolitan; 17 genera and ca. 680 species, 1 genus and 17 species in the flora area. 1. LUDWIGIA L., Sp. Pl. 118. 1753. —Clavo de pozo. Jussiaea L., Sp. Pl. 388. 1753. Erect or creeping herbs rooting at the nodes, shrubs, or trees, aquatic or terrestrial. Underwater parts often swollen and spongy or bearing inflated white spongy pneumatophores. Leaves alternate, opposite, or whorled, mostly entire. Stipules absent or reduced and deltoid. Flowers borne singly, clustered in leaf axils, or in racemes or panicles; floral tube lacking; bracteoles lacking or conspicuous, then usually 2 at or near base of ovary. Sepals 3–7, persistent after anthesis; petals as many as sepals or absent, caducous, yellow or white, contorted in bud. Stamens as many as or twice the number of sepals, very rarely an intermediate number; anthers usually versatile, at times apparently basifixed by reduction; pollen shed in tetrads or singly. Summit of ovary (the disk) flat or conical, often with depressed nectaries surrounding the bases of the epipetalous stamens. Stigma hemispherical or capitate, the upper 1/2–2/3 receptive, often lobed, lobes as many as the locules. Ovary 3–7-locular (as many locules as sepals), very rarely more; ovules pluriseriate or uniseriate in each locule (in L. hyssopifolia uniseriate below, pluriseriate above), when uniseriate, the seeds sometimes embedded in powdery or woody endocarp. Capsule irregularly dehiscent, opening by flaps separating from the valve-like top or by a terminal pore, or indehiscent. Seeds rounded or elongate, the raphe usually easily visible, sometimes ± equal in length to the body of the seed. Cosmopolitan; 83 species, 19 in Venezuela, 17 of these in the flora area. Key to the Species of Ludwigia 1. 1. 2(1).

Plants free-floating, rooting at nodes along the stem .............................. 2 Plants erect or prostrate, rooted to the substrate, generally not rooting at the nodes along the stem ....................................................................... 4 Plants forming symmetrical floating rosettes with rhombic-ovate leaves .................................................................................................... L. sedoides

Ludwigia 189

2. 3(2).

3. 4(1).

4. 5(4). 5. 6(5). 6. 7(6). 7. 8(7). 8. 9(6). 9. 10(9). 10. 11(10).

11.

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

Plants not rosette-forming, with more elongate leafy stems, the leaves rounded to oblong, obovate, or oblanceolate ......................................... 3 Leaves oblong, obovate, or oblanceolate, the upper ones alternate, the lower ones verticillate (sometimes absent in herbarium material); plants without pneumatophores .............................................. L. inclinata Leaves rounded, all alternate; plants with white, erect, spindle-shaped pneumatophores at the nodes ...................................... L. helminthorrhiza Capsules woody and indehiscent, knobby around the seeds and constricted between them; seeds firmly embedded in the woody endocarp and fused to the capsule wall .................................................... L. torulosa Capsules smooth, dehiscent; seeds free from the endocarp or loosely embedded in it ............................................................................................ 5 Seeds uniseriate and embedded in endocarp in the lower part of the capsule, pluriseriate and free in the upper part; sepals 4 ....... L. hyssopifolia Seeds all uniseriate or all pluriseriate; sepals 4–6 .................................. 6 Seeds uniseriate in each locule and embedded in horseshoe-shaped pieces of endocarp ............................................................................................. 7 Seeds pluriseriate in each locule, not embedded in horseshoe-shaped pieces of endocarp .................................................................................. 9 Sepals and petals 4 ............................................................ L. quadrangularis Sepals and petals 5 or 6 ............................................................................. 8 Leaves ovate; sepals 3.5–5 × 1.5 mm; summit of ovary elevated; raphe 1/3–1/4 the width of the seed body .................................................. L. affinis Leaves lanceolate; sepals 5–8 × 2–3 mm; summit of ovary flat; raphe 1/6–1/8 the width of the seed body ............................................ L. leptocarpa Raphe equal in diameter to the body of the seed ....................... L. octovalvis Raphe ≤ 1/4 the diameter of the body of the seed .................................... 10 Capsule wall woody and tough, tardily dehiscent; petals 4–6 mm long .............................................................................................................. 11 Capsule wall not woody, dehiscent and friable; petals usually > 6 mm long .............................................................................................................. 12 Capsule cylindric, 1.5–2 mm wide, with hollow ridges between the angles; flowers usually crowded into spikes; summit of ovary flat, glabrous; stems narrowly winged .......................................... L. densiflora Capsule globose, 7–9 mm wide, not markedly ridged between the angles; flowers solitary or 2 or 3 in leaf axils; summit of ovary 2 mm high, pubescent; stems not winged .......................................................... L. latifolia Capsules oblong, sessile or subsessile; summit of ovary flat; annuals .............................................................................................................. 13 Capsules obconic, long-pedicellate; summit of ovary elevated; perennials .............................................................................................................. 14 Petals and sepals 3–6 mm long; summit of ovary usually glabrous ........................................................................................................ L. erecta Petals 10–20 mm long; sepals 7–12 mm long; summit of ovary pubescent .................................................................................................. L. decurrens Bracteoles foliaceous, 10–25 mm long, accrescent in fruit, borne on middle of capsule ........................................................... L. foliobracteolata Bracteoles not foliaceous, < 10 mm long, usually deciduous, borne on pedicel or at base of ovary ................................................................... 15

190

O NAGRACEAE

15(14). Leaves pubescent ........................................................................ L. peruviana 15. Leaves glabrous or only pubescent along veins on lower surface of leaf .............................................................................................................. 16 16(15). Leaves lanceolate-oblong, 2–8 mm wide; style 2–3 mm long; summit of ovary scarcely elevated .................................................................. L. rigida 16. Leaves elliptic, oblong-ovate, ovate, or lanceolate, 10–40 mm wide; style < 2 mm long; summit of ovary 1–2.5 mm tall ............................ L. nervosa Ludwigia affinis (DC.) Hara, J. Jap. Bot. 28: 291. 1953. —Jussiaea affinis DC., Prodr. 3: 53. 1828. Robust herb to 2.5 m tall, openly branched, entirely covered with spreading trichomes; leaves 2–11 × 1.1–4.2 cm; petals yellow, 6–8 × 4–6 mm. Moist forest edges, gallery forests, near sea level to 400(–800) m; Delta Amacuro (widespread), Bolívar (widespread), Amazonas (San Carlos de Río Negro). Mérida, Monagas, Portuguesa, Táchira, Yaracuy, Zulia; Guatemala, West Indies, southern Brazil, Bolivia and; western Africa. ◆Fig. 190. Ludwigia decurrens Walter, Fl. Carol. 89. 1788. —Jussiaea decurrens (Walter) DC., Prodr. 3: 56. 1828. Erect glabrous herb to 1 m tall, slightly or profusely branched; branches sharply 4angled and 4-winged from decurrent leaf bases; leaves 2–20 × 0.2–5 cm; pedicel 2–15 mm long; petals yellow, 10–20 × 10–18 mm. Muddy stream banks, marshy shores of lakes and ponds, swamps, ditches; near sea level to 600 m; Delta Amacuro (widespread), northern Bolívar, Amazonas (widespread). Widespread at low elevations in Venezuela; southern U.S.A. to northern Argentina, introduced in the Old World. ◆Fig. 192. Ludwigia densiflora (Micheli) Hara, J. Jap. Bot. 28: 292. 1953. —Jussiaea densiflora Micheli, Flora 57: 301. 1874. Robust, glabrous, annual herb to 1.5 m tall; stem simple or branched, erect, manyangled; leaves crowded, 1–10 × 0.3–2.5 cm; petals yellow, 4–5 mm long. Sandy river banks, near sea level to 100 m; Delta Amacuro (Caño Macareo), Bolívar (Ciudad Bolívar). Apure; Amazonian Colombia, Peru, Brazil, and Bolivia. ◆Fig. 187. Ludwigia erecta (L.) Hara, J. Jap. Bot. 28: 292. 1953. —Jussiaea erecta L., Sp. Pl. 388. 1753.

Jussiaea linifolia auct. non Vahl: sensu Hutch. & Dalziel, Fl. W. Trop. Afr. 1: 146. 1927. Erect, glabrous, annual herb to 3 m tall; flowers subsessile; leaves 2–20 × 0.2–4 cm; petals yellow, 3–6 × 2–3 mm. Pond margins, wet depressions, seasonally flooded areas, near sea level to 1100 m; Delta Amacuro (widespread), Bolívar (scattered, including Rio Parupa on the Gran Sabana), Amazonas (scattered). Widespread at low elevations in Venezuela; widespread in Neotropics, U.S.A., introduced throughout tropical Africa. Ludwigia foliobracteolata (Munz) Hara, J. Jap. Bot. 28: 292. 1953. —Jussiaea foliobracteolata Munz, Darwiniana 4: 228. 1942. Shrub to 3 m tall, densely branched, villous on very young shoots, hirsute on lower surfaces of leaves, otherwise glabrous; leaves 5–25 × 2–8 cm; pedicels 10–25 mm long; petals yellow, 10–20 mm long, orbicular. Ponds, streams, swamps, near sea level to 800 m; Delta Amacuro (Río Amacuro, Río Toro), Bolívar (Río Icabarú, Río Mato at north end of Sierra de Maigualida). Táchira; southern Mexico to southern Peru and northern Brazil. Ludwigia helminthorrhiza (Mart.) Hara, J. Jap. Bot. 28: 292. 1953. —Jussiaea helminthorrhiza Mart., Flora 22(1) Beibl. 1: 61. 1839. Jussiaea natans Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 1: 16, fig. 3B. 1808 [1805], non Ludwigia natans Elliott 1821. Floating, glabrous herb; leaves 1.2–5.6 × 1.2–4 cm; pedicels 25–50 mm long; petals white, 7–15 × 5–13 mm. In still waters, near sea level to 100 m; Delta Amacuro (Tucupita), Bolívar (Ciudad Bolívar). Widespread at low elevations in Venezuela; Neotropics from central Mexico to Paraguay. Ludwigia hyssopifolia (G. Don) Exell, Garcia de Orta 5: 471. 1957.

Ludwigia 191

—Jussiaea hyssopifolia G. Don, Gen. Hist. 2: 693. 1832. Jussiaea linifolia Vahl, Eclog. Amer. 2: 32. 1798, non Ludwigia linifolia Poir. 1813. Annual or perennial herb 5 cm to 3 m tall, glabrous or minutely puberulent; leaves 1–9 × 0.2–3 cm; petals yellow, 2–3 × 1–2 mm. Wet areas, near sea level to 200 m, widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread at low elevations in Venezuela; southern Mexico and West Indies south to Brazil and Bolivia, also widely distributed in the Old World. ◆Fig. 194. Ludwigia inclinata (L. f.) M. Gómez, Anales Soc. Esp. Hist. Nat. ser. 2, 3: 66. 1894. —Jussiaea inclinata L. f., Suppl. Pl. 235. 1781 [1782]. Aquatic, ± floating herb, sparingly or profusely branched; leaves 0.8–10 × 0.2–0.7 cm, the basal ones verticillate and linear, the upper ones alternate and oblong-obovate or oblanceolate; pedicels 2–7 mm long; petals yellow, 12–25 × 10–19(–25) mm. Ponds, swamps, wet ditches, 50–200 m; northwestern Bolívar, Amazonas (near Puerto Ayacucho). Venezuelan Llanos; southern Mexico and West Indies south to southern Brazil and Bolivia. ◆Fig. 196. Ludwigia latifolia (Benth.) Hara, J. Jap. Bot. 28: 292. 1953. —Jussiaea latifolia Benth., J. Bot. (Hooker) 2: 317. 1840. Perennial herb or shrub to 2.5 m tall, profusely branched and spreading; leaves 5–18 × 1.5–5 cm; pedicels 2–9 mm long; petals yellow, 5–6 × 6–7 mm. Wet and seasonally flooded areas, near sea level to 800 m; Delta Amacuro (widespread), Bolívar (widespread), Amazonas (Simarowochi). Nicaragua south to northern Brazil and central Bolivia. ◆Fig. 193. Ludwigia leptocarpa (Nutt.) Hara, J. Jap. Bot. 28: 292. 1953. —Jussiaea leptocarpa Nutt., Gen. N. Amer. Pl. 1: 279. 1818. Jussiaea leptocarpa var. genuina Munz, Darwiniana 4: 255. 1942. Robust herb to 3 m tall, densely pubescent to subglabrous; leaves 3.5–18 × 1–4 cm; pedicels 2–20 mm long; petals yellow, 5–11 × 4–8 mm. Wet areas, evergreen lowland forests, near sea level to 300 m; Delta Amacuro (widespread), Bolívar (Río Caura, Tumeremo

to Anacoco), Amazonas (Nericagua, Puerto Ayacucho, San Carlo de Río Negro). Apure; Southeastern U.S.A. to Argentina, Africa. ◆Fig. 188. Ludwigia nervosa (Poir.) Hara, J. Jap. Bot. 28: 293. 1953. —Jussiaea nervosa Poir. in Lam., Encycl. suppl. 3: 199. 1813. —Jussiaea nervosa var. typica Munz, Darwiniana 4: 208. 1942 Jussiaea maypurensis H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 100, pl. 531. 1823. Jussiaea nervosa var. pubescens Micheli in Mart., Fl. Bras. 13(2): 155. 1875. Shrub to 5 m tall, profusely branched, glabrous or only pubescent on lower surface of leaves; leaves 2–16 × 2–4 cm; pedicels 8–40 mm long; flowers numerous; inflorescences ± paniculate; petals yellow, 14–20 × 13–20 mm. Savannas, Mauritia palm swamps, lake margins, 50–1000 m; Bolívar (widespread), Amazonas (Puerto Ayacucho, Río Manapiare basin). Apure, Guárico, Miranda, Monagas, Sucre, Táchira, Zulia; southern Mexico, Central America, Colombia, Guianas, Ecuador, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 195. Ludwigia octovalvis (Jacq.) Raven, Kew Bull. 15: 476. 1962. —Oenothera octovalvis Jacq., Enum. Syst. Pl. 19. 1760. Jussiaea suffruticosa L., Sp. Pl. 388. 1753, non Ludwigia suffruticosa Walt. 1788. Robust herb or shrub to 4 m tall, wellbranched, subglabrous, puberulent, or densely villous; leaves 0.7–14.5 × 0.1–4 cm; pedicels 0–10 mm long; petals yellow, 3–17 × 2– 17 mm. Wet areas, near sea level to 600 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Throughout Venezuela; pantropics. ◆Fig. 191. Ludwigia peruviana (L.) Hara, J. Jap. Bot. 28: 293. 1953. —Jussiaea peruviana L., Sp. Pl. 388. 1753. —Jussiaea peruviana var. typica Munz, Darwiniana 4: 232. 1942. Jussiaea mollis H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 102. 1823. Jussiaea peruviana var. glaberrima J.D. Sm., Bot. Gaz. (Crawfordsville) 16: 6. 1891. Erect perennial herb or shrub, occasionally decumbent, to 4 m tall, villous; leaves 2– 45 × 1–10 cm; pedicels 5–65 mm long; petals yellow, 10–40 mm long, orbicular. Swampy

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savannas, palm swamps, ca. 400 m; Bolívar (Altiplanicie de Nuria). Widespread in northern half of Venezuela; southern U.S.A. and West Indies south to Chile and Argentina, naturalized at scattered localities in Asia. ◆Fig. 189.

Central America, Colombia, Guyana, Suriname, Brazil, Bolivia. Some unusually pubescent plants were collected near Kamarata and on Cerro Apacará in Bolívar state (Koyama & Agostini 7249, VEN, and Cardona 1964, VEN).

Ludwigia quadrangularis (Micheli) Hara, J. Jap. Bot. 28: 294. 1953. —Jussiaea quadrangularis Micheli, Flora 57: 302. 1874. Scandent shrub to 2 m tall, glabrous or minutely puberulent, well branched; leaves 3–11 × 1.5–3.5 cm; pedicels 4–8 mm long; petals yellow, 12–15 × 6–10 mm. Moist areas, 50–200 m; Bolívar (El León near Río Caroní, Tumeremo to Bochinche). Colombia, Ecuador, Peru, Brazil.

Ludwigia sedoides (Bonpl.) Hara, J. Jap. Bot. 28: 294. 1953. —Jussiaea sedoides Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 1: 15. 1808 [1805]. Floating perennial herb; leaves 0.5–2 × 0.5–2 cm, symmetrically and densely arranged; pedicels 10–30 mm long; petals yellow, 10–15 × 8–13 mm. Forming distinctive colonies in lakes and ponds, near sea level to 200 m; Delta Amacuro (Caño de Uricoa), northern Bolívar, Amazonas (near Puerto Ayacucho). Venezuelan Llanos; southern Mexico, Central America, West Indies, Colombia, Guyana, Ecuador, Brazil, Bolivia, Paraguay. ◆Fig. 197.

Ludwigia rigida (Miq.) Sandwith, Kew Bull. 19: 197. 1965. —Jussiaea rigida Miq., Stirp. Surinam. Select. 58. 1850 [1851]. Jussiaea lithospermifolia Micheli, Flora 57: 300. 1874. —Jussiaea lithospermifolia var. typica Munz, Darwiniana 4: 214. 1942. —Ludwigia lithospermifolia (Micheli) H. Hara, J. Jap. Bot. 28: 292. 1953. Jussiaea lithospermifolia var. pubescens Munz, Darwiniana 4: 214. 1942. Shrublet or perennial herb to 1.5 m tall, glabrous or sometimes hirsute; leaves 3–12 × 0.2–0.8 cm; pedicels 5–35 mm long; petals yellow or rarely white, 10–15 mm long, orbicular. Savannas, palm swamps, gallery forests, lake borders, 50–1000 m; widespread in Bolívar and Amazonas. Venezuelan Llanos and Coastal Cordillera; southern Mexico,

Ludwigia torulosa (Arn.) Hara, J. Jap. Bot 28: 294. 1953. —Jussiaea torulosa Arn., Ann. Sci. Nat. Bot. sér. 2, 3: 251. 1835. Perennial, erect, aquatic herb to 1.7 m tall, attached to the substrate, with a single main stem; leaves 7–17 × 0.3–2.5 cm; pedicels 2–6 mm long; petals white, 2–5 mm long, orbicular. In shallow waters at edges of lakes and swamps, near sea level to 200 m; Delta Amacuro (Laguna Alamilla along Caño Acoimito near Río Orocoima), Bolívar (south of Tumeremo). Neotropics from southern Mexico south to southern Brazil and Bolivia. ◆Fig. 198.

1.5 mm 0.5 mm 1 cm 1 cm 1 cm 0.5 mm Fig. 187. Ludwigia densiflora

Fig. 188. Ludwigia leptocarpa

Fig. 189. Ludwigia peruviana

Ludwigia 193

0.5 mm

1.5 mm

Fig. 190. Ludwigia affinis

1 cm

Fig. 191. Ludwigia octovalvis

1 cm

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Fig. 192. Ludwigia decurrens

0.8 mm

1 cm

1 cm

0.4 mm

Fig. 193. Ludwigia latifolia

Ludwigia 195

Fig. 194. Ludwigia hyssopifolia

Fig. 195. Ludwigia nervosa

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Fig. 196. Ludwigia inclinata

0.6 mm

1 cm

1 cm

0.6 mm Fig. 197. Ludwigia sedoides

Ludwigia 197

1 mm

5 mm

Fig. 198. Ludwigia torulosa

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O PILIACEAE

OPILIACEAE by James S. Miller Evergreen trees or shrubs, sometimes lianas, hemiparasitic on roots of other plants. Leaves alternate, simple, entire, estipulate. Inflorescences axillary or cauliflorus, spicate, racemose, catkin-like, paniculate, or umbellate, each flower usually subtended by a caducous bract. Flowers small, actinomorphic, bisexual or less commonly unisexual (Agonandra, Gjellerupia), the plants then dioecious. Torus (sometimes called calyx) minute, cupulate, truncate or 4- or 5-lobed or toothed. Tepals (sometimes called petals) 4 or 5, valvate, free or basally connate, lacking or caducous in pistillate flowers. Stamens 4 or 5, opposite the tepals; filaments free or adnate to the tepals, alternating with free or nearly connate nectaries; anthers bithecal, dehiscing longitudinally. Ovary superior or half-inferior and immersed in the disk, unilocular with a single pendulous, anatropous ovule or the ovule basal (Agonandra); style simple, stigma entire or shallowly lobed. Fruits drupaceous; seeds with copious oily, starchy endosperm. Widespread in tropical and subtropical regions of both hemispheres; 10 genera and 33 species, 1 genus and 2 species in the flora area. 1. AGONANDRA Miers ex Benth. in Benth, & Hook. f., Gen. Pl. 1: 349. 1862. Evergreen trees or shrubs, rarely vinelike, usually with corky bark. Leaves usually glabrous, ovate to elliptic, base of blade decurrent on the petiole. Inflorescences axillary racemes, with 1–3(4) flowers per bract, bracts caducuous. Flowers small, pedicellate, unisexual, the plants dioecious. Torus (“calyx”) minute, cupulate, 4- or 5-toothed or merely undulate. Tepals (“petals”) 4(5), green, free, valvate, rapidly caducous in pistillate flowers. Staminate flowers with 4(5) stamens; filaments opposite and adnate to the tepals, alternate with the lobes of the nectariferous disk; ovary rudimentary. Pistillate flowers with superior ovary, with one basal, anatropous ovule; style usually absent, the stigma borne directly on the apical surface of the ovary, usually 4-lobed. Drupe ellipsoid to globose. Mexico, Central America, South America south to Argentina; 10 species, 2 in Venezuela, both in the flora area. Key to the Species of Agonandra 1.

1.

Shrubs or small trees in savannas or deciduous forest; trunk with large, corky ridges; flowers and racemes densely puberulous; petioles 5–20 mm long ................................................................................. A. brasiliensis Large forest trees in evergreen lowland forest; trunk with smooth bark; flowers and racemes glabrous; petioles 2–5 mm long .............. A. silvatica

Agonandra brasiliensis Miers ex Benth. in Benth. & Hook. f., Gen. Pl. 1: 349. 1862. Shrub or trees. Savannas and deciduous forests. Panama, Colombia, Venezuela, Guyana, Brazil, Bolivia, Paraguay; 2 subspecies, both in the flora area. The seeds of Agonandra brasiliensis are used as an oil source yielding a partially hydrogenated oil, and the tree also yields a rubber substitute.

Key to the Subspecies of A. brasiliensis 1. Staminate inflorescence with 3(4) flowers per bract, pistillate inflorescence at least in lower half with 3 flowers per bract ............................... subsp. brasilensis 1. Staminate and pistillate inflorescences with 1 flower per bract ............................ ............................... subsp. racemigera

Agonandra 199

A. brasiliensis subsp. brasiliensis. —Aceituna, Ocoma (Yanomami). Shrub or tree 2–20 m tall. Savannas and deciduous forests or shrublands, 50–400 m; Delta Amacuro ( Los Castillos de Guayana), Bolívar (Cerro Marimarata, Ciudad Bolívar, El Manteco, El Palmar, La Paragua, La Urbana, Maripa, Represa Guri, Río Ariza, lower Río Suapure, 30 km east of Upata), Amazonas (environs of Puerto Ayacucho, Río Matacuni, Río Ocamo). Anzoátegui, Cojedes, Falcón, Guárico, Lara, Sucre, Zulia; Panama, Colombia, Guyana, Brazil, Bolivia, Paraguay. ◆Fig. 199.

Fig. 199. Agonandra brasiliensis subsp. brasiliensis

A. brasiliensis subsp. racemigera Hiepko, Bot. Jahrb. Syst. 117: 498. 1995. —Aceituno. Small tree (2–)4–8 m tall. Deciduous forests, near sea level to 50 m; Bolívar (San Felix to Puerto Ordaz). Portuguesa, Zulia; Colombia. Agonandra silvatica Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 41. 1922. Tree to 30 m tall. Evergreen lowland forests, 50–100 m; Amazonas (expected at base of Sierra de la Neblina and known just across border with Brazil). Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

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ORCHIDACEAE by Germán Carnevali, Ivón M. Ramírez-Morillo, Gustavo A. Romero-González, Carlos A. Vargas, and Ernesto Foldats

History of Orchid Botanical Exploration by Gustavo A. Romero-González A broad variety of people with different backgrounds and objectives have collected plants in the Venezuelan Guayana. Although much is known about the scientific collectors, such as Alexander von Humboldt and Richard Spruce, the work of professional plant collectors hired by commercial horticultural firms to collect live material remains largely unknown and difficult to trace, and collectors are often only known by their last names. Because the latter group focused most of their energy on ornamental plants, among them orchids, they will be the primary focus of this discussion. They arguably may have had the greatest impact on our knowledge of the orchid flora of the Venezuelan Guayana. Nevertheless, the accomplishments of a few of the scientific collectors aforementioned will be highlighted because of the novelty or rarity of the orchids they collected. Collectors are listed in the chronological order they visited the Venezuelan Guayana until 1980. Alexander von Humboldt and Aimé Bonpland collected few orchids relative to the volume of their other botanical collections. Only two species were described from the flora area in the Nova Genera et Species Plantarum (Humboldt et al. 1815– 1825). Habenaria angustifolia H.B.K. “Crescit in humidis, uliginosis Guayanae inter El Trapiche de fereras et urbem Santo Thomas del Angostura” [Humboldt, Bonpland & Kunth, Nov. Gen. Spec. (quarto ed.) 1: 330. 1821], is known only from the type in Paris and has not been examined by contemporary orchidologists. The identity of this species could not be clarified and furthermore its affinity with other species of Habenaria from the flora area is unknown. Cymbidium violaceum H.B.K., currently referred to the genus Cattleya, is without doubt the showiest and most conspicuous orchid species in the lowland forests of the Venezuelan Guayana. Robert H. Schomburgk collected on Roraima-tepui (Mount Roraima) and along the upper Río Orinoco, including Cerro Duida and Cerro Marahuaka (Schomburgk 1931). Several new species were later described from his collections, among them Masdevallia guayanensis Lindl., Sobralia liliastrum Lindl., Cypripedium lindleyanum R.H. Schomb.[= Phragmipedium lindleyanum], Cleistes parviflora Lindl., Diothonea imbricata Lindl. [= Hexisea imbricata], Maxillaria eburnea, and Stelis schomburgkii Fawc. & Rendle. However, his major contribution was to document the existence of two orchids of great ornamental value that would later become the target of commercial collectors: Cattleya lawrenceana Rchb. f. and Zygopetalum burkei Rchb. f. [= Galeottia burkei]. The first one he did not actually collect, believing it to be Cattleya mossiae Hook., a well-known species at that time. He did collect the then-undescribed Zygopetalum burkei Rchb. f., but his botanical specimens remained misfiled at Kew until recently. Nonetheless, drawings of both species were deposited at the British Museum and, in conjunction with Schomburgk’s notes, allowed commercial firms to accurately guide their collectors (Sander 1888, 27).

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Richard Spruce collected at least one new orchid species in the Venezuelan Guayana (Masdevallia sprucei Rchb. f.). It is noteworthy that Spruce explored such remote areas in Venezuelan territory that some of the orchids he collected were not uncovered again for more than a hundred years [e.g., Trisetella triglochin (Rchb. f.) Luer]. Richard Payer, a professional collector working for L’Horticulture Internationale of Brussels, collected along the upper 0rinoco in 1881 (Linden et al. 1894, 120). Catasetum pileatum Rchb. f. was among his findings, but this valuable ornamental remained obscured until it was rediscovered by E. Bungeroth and redescribed as Catasetum bungerothii N.E. Br. Ironically, this species was first collected by Spruce in 1854, but his specimen remained misfiled in Kew until 1889 (Rolfe 1889). David Burke, a professional collector for the British firm J. Veitch & Sons, followed R.H. Schomburgk’s path to Roraima, where he collected three new orchid species from 1881–1882 (Im Thurn 1884b; Veitch 1906, 87), including Zygopetalum burkei Rchb. f. (currently referred to Galeottia). E. Seidel (sometimes spelled Seidle or Seidl), a German professional orchid collector working for F. Sander and Company, collected in Roraima twice in 1884. His first expedition reached Roraima in April via Trinidad to Ciudad Bolívar and then to Upata and finally to El Callao (called “Callovo” by Seidel), thus becoming the first collector to reach Roraima entirely through Venezuelan territory (Seidel 1890, 77). Seidel collected a large shipment of a beautiful Cattleya, then undescribed, but the plants were somehow lost on their way to England (Im Thurn 1885a, 24). He returned in October of 1884, via Georgetown, when he encountered the Im Thurn expedition on his way to Roraima (Im Thurn 1885a, 1–2). He later left Roraima with Im Thurn, carrying a large shipment of the same Cattleya, which was later described as C. lawrenceana Rchb. f. (Reichenbach 1885; Sander 1888, 27). Everard Ferdinand Im Thurn collected ten new orchid species in Roraima during the course of his 1885 expedition, many of them on the summit, including the rare ornamental Oncidium orthostates Ridl. (Im Thurn 1885a, 1885b, 1886). Ed. Kromer, an American professional collector, organized two expeditions to Roraima, one in 1886 and the second some years later. The published account of the second trip (Kromer 1911–1912) provides only the months, September–January, not the years spent on the trip to collect Cattleya lawrenceana Rchb. f. Both expeditions were mounted via Georgetown. Erich Bungeroth, a German professional collector working for L’Horticulture Internationale (Linden et al. 1894, 120), collected along the upper Río Orinoco from 1886–1889. He introduced several ornamental novelties including Paphinia lindeniana Rchb. f., Peristeria aspersa Rolfe (“ … de la Sierra de Marawaca … “, type locality information from Rolfe 1891, 57), Zygopetalum jorisianum Rolfe (= Galeottia jorisiana), and Zygopetalum lindeniae Rolfe (= Zygosepalum lindeniae). Perhaps Bungeroth’s greatest contribution was the large collection of live Catasetum plants he shipped to Brussels. Among them was C. bungerothii N.E. Br., a species that “ … caused a flutter among the Orchidologists … At the sale in question the best plants realized fifty guineas … “ (Anonymous 1887, 139). This species was, of course, the Catasetum pileatum previously collected by Payer (see above). The German explorer Ernst H. G. Ule collected on Roraima in early 1910 (Ule 1914). At least one of his collections turned out to be a new species, Koellensteinia roraimae Schltr.

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Members of the Tyler-Duida Expedition (Gleason 1931) collected five new species described by Ames and Schweinfurth (1931), including the beautiful Zygopetalum tatei Ames & C. Schweinf. (= Zygosepalum tatei). Julian A. Steyermark undoubtedly will be remembered as the most prolific orchid collector of the Venezuelan Guayana in the 20th century. Starting in 1944, Steyermark collected a great number of new records for the orchid flora of the area, including many new species that bear his name, e.g., Aspidogyne steyermarkii Carnevali & Foldats, Brachionidium julianii Carnevali & I. Ramírez, Campylocentrum steyermarkii Foldats (here synonymized under C. hondurense), Jacquiniella steyermarkii Carnevali & Dressler, Octomeria steyermarkii Garay & Dunst., Selenipedium steyermarkii Foldats, Trichosalpinx steyermarkii Luer, and Xylobium steyermarkii Foldats (here synonymized under Bifrenaria steyermarkii). Ernesto Foldats, author of the orchid treatment for the official Flora de Venezuela project (Foldats 1969–1970), explored the basins of Río Atabapo, Río Ventuari, and upper Río Orinoco from 1960–1970. He collected several new species including Koellensteinia lilijae Foldats. In addition, Foldats participated in several collecting trips to the Gran Sabana region, including the 1968 Angel Falls expedition organized by the Explorers Club of Pittsburgh and the Pittsburgh Zoological Society (Jirak 1968). The team of Galfrid C. K. Dunsterville and Ellinor Dunsterville carefully collected, cultivated, and drew hundreds of orchid species of the Venezuelan Guayana over three decades, including many new reports and species (e.g., Dunsterville and Dunsterville 1982). At least six orchid species (Bulbophylllum dunstervillei, Catasetum ×dunstervillei, Cryptocentrum dunstervilleorum, Maxillaria ×dunstervillei, Paphinia dunstervillei, Scaphyglottis dunstervillei, and one genus (Dunstervillea Garay) from this region are named after them. Galfrid C. K. Dunsterville and Leslie A. Garay described at least 20 new orchid species of the Venezuelan Guayana for their monumental series Venezuelan Orchids Illustrated (1959–1976). These books, and their Orchids of Venezuela: An Illustrated Field Guide (Dunsterville and Garay 1979), remain classic reference works for those studying the orchids of northern South America (see Romero and Carnevali for a second, revised edition).

References Cited Ames, 0., and C. Schweinfurth. 1931. Orchidaceae. In Botanical Results of the Tyler-Duida expedition, by A. Gleason. Bull. Torrey Bot. Club 68: 345–353. Anonymous. 1887. Catasetum bungerothii. Gard. Chron. ser. 3, 1: 139–140. Dunsterville, G. C. K., and L. A. Garay. 1959–1976. Venezuelan Orchids Illustrated. 6 volumes. London: Andre Deutsch. Dunsterville, G. C. K., and L. A. Garay. 1979. Venezuelan Orchids: An Illustrated Field Guide. 3 volumes. Cambridge: Harvard University Press. Dunsterville, G. C. K., and E. Dunsterville 1982. Auyán-tepui: Reminiscences of an orchid search. In Orchid Botany: Reviews and Perspectives, edited by J. Arditti, 19–38. Ithaca: Cornell University Press. Foldats, E. 1969–1970. Orchidaceae. In Flora de Venezuela, edited by T. Lasser, volume 15, parts 1–5. Caracas: Instituto Botanico. Gleason, H. A. 1931. Botanical results of the Tyler-Duida expedition. Bull. Torrey Bot. Club 58: 277–506.

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Humboldt, A. von, A. Bonpland, and K. S. Kunth. 1815–1825. Nova Genera et Species Plantarum. 7 volumes. Paris: Sumtibus Librariae Graeco-LatinoGermanicae. Im Thurn, E. F. 1884a. Cattleya lawrenceana. Gard. Chron. n.s. 25: 168. Im Thurn, E. F. 1884b. New plants from Guiana. Timehri 3: 156–158. Im Thurn, E. F. 1885a. The first ascent of Roraima. Timehri 4: 1–48. Im Thurn, E. F. 1885b. Roraima. Timehri 4: 255–267. Im Thurn, E. F. 1886. Notes on the plants observed during the Roraima expedition of 1884. Timehri 5: 147–223. Jirak, I. L. 1968. Angel Falls Expedition 1968. Mimeo. Kromer, E. 1911–1912. An orchid collector’s travels through British Guiana to Brazil. Orchid World 2: 55–57, 88–90, 107–110, 136–139, 175–177. Linden, L., A. Cogniaux, and G. Grignan. 1894. Les Orchidées Exotiques. Gand: Eug. Vander Haeghen. Reichenbach, H. G. 1885. Cattleya lawrenceana. Gard. Chron. n.s. 23: 338. Rolfe, R. A. 1889. Catasetum bungerothii var. aureum. Gard. Chron. ser. 3, 6: 466. Rolfe, R. A. 1891. Peristeria aspersa. Lindenia 6: 57–58, t. 267. Romero-González, Gustavo A., and Germán Carnevali Fernández-Concha. 2000. Orchids of Venezuela: An Illustrated Field Guide. Second edition. Caracas, Venezuela: Armitano Editores. Sander, F. 1888. Reichenbachia. Volume 1. London: H. Sotheran. Schomburgk, R. H. 1931. Travels in Guiana and on the Orinoco. Translated by W.E. Roth. Georgetown, Guyana: Argosy. Seidel, E. 1890. Voyage aux montagnes de Roraima. Orchidophile 10: 5–7, 76–77. Ule, E. 1914. Die vegetation des Roraima. Bot. Jahrb. Syst. 52(Beibl. 115): 42–53. Veitch, J. H. 1906. Hortus Veitchii. London: Gilbert & Rivington.

Orchidaceae Family Description by Germán Carnevali and Ivón M. Ramírez-Morillo Strongly mycotrophic, lithophytic, terrestrial, or often epiphytic, sympodial or monopodial perennial herbs or rarely vines, usually green and photosynthetic, but some without chlorophyll or wholly subterranean. Roots of various forms, e.g., tuberoid, a root or stem tuber, or stolonoid, the epidermis of which most often modified into a spongy, multilayered velamen (water-absorbing layer of dead cells). Rhizomes delicate to stout, sometimes creeping or condensed, simple or branched. Stem of various forms (especially diverse in sympodial orchids), a corm, pseudobulb (variously thickened aerial stem), ramicaul (aerial slender stem in Pleurothallidinae, generally 1-foliate), or absent, sometimes proliferous. Leaves alternate or distichous, seldom opposite and whorled, sometimes all basal, or reduced to mere scales (most often in saprophytic and leafless orchids), simple, entire or rarely deeply lobed, variously convolute, duplicate, conduplicate, cylindrical, triangular, or laterally flattened, sheathing at the base or petiolate, plicate, articulate or not, parallelveined or rarely net-veined (Epistephium). Inflorescences racemose or rarely cymose, spicate, or paniculate, 1–many-flowered, erect or variously pendent, lax or dense, terminal or lateral, flowering successively or simultaneously. Flowers bisexual or sometimes unisexual (Catasetum), pedicellate or sessile, 3-merous, gener-

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ally zygomorphic, rarely actinomorphic, sometimes with a calyculus (small cup of bract-like structures outside the sepals, Epistephium), with an abscission layer between the pedicel and the peduncle, rarely with an abscission layer between the ovary and perianth (Vanilla) or between the ovary and the pedicel (e.g., Pleurothallis or subtribe Pleurothallidinae in general), epigynous, resupinate (twisted 180), hyper-resupinate (twisted 360 degrees), or not. Perianth typically 6 tepals in 2 whorls, all petaloid, or the sepals sometimes greener and more foliaceous in texture; sepals alike or not, the median one (morphologically abaxial, but in resupinate flowers appearing adaxial) or 2 (forming synsepal) or all 3 sepals variously connate; petals 3, the median one (morphologically abaxial, but in resupinate flowers appearing adaxial) strongly differentiated from the others (excepting subfamily Apostasoideae), commonly larger or differing in form and color, forming a lip (labellum), the 2 lateral petals commonly but not always ± similar to the sepals; nectaries of various sorts, sometimes forming a hollow spur extended backward or downward from the base of the lip, or at the sepal tips, sometimes a cup within the perianth atop the ovary, or embedded in the ovary wall and opening toward the summit (modified septal nectaries); extrafloral nectaries sometimes present on the pedicels, bracts, or leaf sheaths. Stamens usually 1, rarely 2 or 3, all on the morphologically abaxial (usually seemingly adaxial) side of the flower, on the opposite side of the lip, fully or partially adnate to the style, forming a column (gynostemium); pollen grains in monads or most often in tetrads, usually compacted in solid aggregates called pollinia; pollinia 2–8, sometimes subdivided into small packets or massulae, united by elastoviscin threads (sectile condition), rarely granular, sometimes the pollinia with extensions called caudicles (produced by the anther), and/or with stalks (a hamulus when produced by stigmatic tissue, a tegula when produced by column tissue), the aggregation of pollinia, viscidium, caudicles, and stalks form the pollinarium (= pollination unit transferred by the pollinators). Gynoecium 3-carpellate, united to form a compound, inferior, 1- or 3-locular ovary; style fused with the filaments in various degrees; stigmas 1–3-lobed, concave to convex, wet, the median stigma lobe modified into the rostellum, which often separates the anther from the other 2 (often connate) stigmatic lobes and commonly prevents self-pollination, the rostellum produces the viscidium (sticky structure which helps the attachment of the pollinaria to their pollinators); column often with a basal outgrowth, the column foot, to which the lip, or lateral sepals, or even the petals are united; ovules numerous and very tiny, anatropous. Fruits mostly capsular, opening by longitudinal slits, rarely fleshy and indehiscent (Vanilla). Seeds tiny, numerous (millions in some species), without endosperm, association with an appropriate fungus necessary for germination. Cosmopolitan except Antarctica (most diverse in tropics); ca. 800 genera and 20,000–25,000 species, 157 genera and 732 species plus 6 natural hybrids in the flora area.

Glossary of Orchid Morphology by Carlos A. Vargas anther cap: the operculate covering of the pollinia. callus, pl. calli: a thickening, a protuberance, or one or more crests or keels of the lip disk.

O R C H I D A C E A E 205

calyculus: a small cup or circle of bract-like structures outside of the sepals (Epistephium). caudicle: produced by the anther, it is generally a slender, mealy, or elastic extension of the pollinium, or a mealy portion at one end of the pollinium (not a tegula or a hamulus). clinandrium: the anther bed, that portion of the column under or surrounding the anther. column: the central structure of an orchid flower, composed of the style united to the filaments of 1–3 anthers. column foot: a ventral extension of the base of the column that has the lip attached at its tip. conduplicate: leaves folded along the midvein (e.g., Cattleya and Pleurothallis). corm: a bulb-like fleshy stem or base of stem. cuniculus: a tubular nectary that is concealed within the floral tube above the ovary of the flower. disk (disc): the upper surface of the central portion of the lip. epichile: the terminal part of a complex lip, which is divided into 2 or 3 distinct parts. glenion: a small area with a shiny surface that appears wet, which is on the uppermost plane of the lip where it rests beneath the column. hamulus, pl. hamuli: pollinium stalk developed from the apex of the rostellum. heteroblastic: pseudobulbs each with a single internode. homoblastic: pseudobulbs each with several internodes. hypochile: the basal part of a complex lip, which is divided into 2 or 3 distinct parts. incumbent: said of anthers that bend downward for 90–100°. labellum: see lip. lepanthiform sheaths: tubular, ribbed, imbricating sheaths with oblique, dilated, margined ostia, the ribs and margins of the ostia are grossly or microscopically ciliate or scabrous, unique to Draconanthes, Lepanthes, Lepanthopsis, and Trichosalpinx. lip: the modified third petal, often showy and distinct. massula, pl. massulae: a packet of pollen in those orchids that have the pollinium subdivided into small packets; see sectile. mentum: a chin-like extension at the base of the flower, made up of the column foot, the base of lip, and the lateral sepals. mesochile: the middle portion of a complex lip (only present when lip is divided into 3 distinct parts). ostia: the open apex of funnel-like bracts, commonly ciliate, unique to Draconanthes, Lepanthes, Lepanthopsis, and Trichosalpinx. plicate: leaves with several ± equally prominent longitudinal veins and folded between the veins (e.g., Catasetum and Stanhopea). pollinarium: the complete set of pollinia from an anther, with associated parts, viscidium, or viscidium and stipe; when there are 2 viscidia, each half of the set might be termed a pollinarium, generally being the unit transported by pollinators. pollinium, pl. pollinia: a ± compact and coherent mass of pollen. proliferous: producing roots (usually) and pseudobulbs or ramicauls from the apex of other pseudobulbs or ramicauls (e.g., Scaphyglottis). pseudobulb: a thickened stem, usually aerial.

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VEGETATIVE TERMS

(1)

(1) (2)

Heteroblastic pseudobulb

Homoblastic pseudobulb

Proliferous stems

Ostium (opening)

Stem

Calycule (bract-like structure below sepal)

Lepanthiform sheaths

(2)

(2)

(2)

FLOWER TERMS Dorsal sepal Petal

Dorsal sepal

Petal Column Ovary Pedicel

Bract

Disk (darkened area)

Anther (incumbent) Lateral Callus sepal Lip

Column

Lip

(1) Lateral sepal

(1) Flowers resupinate

Fig. 200. Drawings courtesy of: (1) Robert L. Dressler. Phylogeny and Classification of the Orchid Family. 1993. Portland, OR: Dioscorides Press. (2) G. C. K. Dunsterville and Leslie A. Garay. 1979. Orchids of Venezuela: An Illustrated Field Guide. Allston, MA: Botanical Museum of Harvard University.

O R C H I D A C E A E 207

FLOWER TERMS (continued)

Column foot

(1)

(1)

Cuniculus

(Fruit)

Mentum (chin-like part of lip) Hypochile

(1)

Lip spur Mesochile

Anther

(2) Epichile

Rostellum Stigma

Staminodium Anther 3-lobed stigma

(1)

(1)

(1)

(1)

Pollinia not sectile

Pollinia sectile (many massulae)

Viscidium Hamulus (derived from viscidium; darkened area)

Caudicle Tegula (derived from column; darkened area)

(3)

Viscidium

Tegula

(3)

(1) Viscidium

(1)

Fig. 201. Drawings courtesy of: (1) Robert L. Dressler. Phylogeny and Classification of the Orchid Family. 1993. Portland, OR: Dioscorides Press. (2) G. C. K. Dunsterville and Leslie A. Garay. 1979. Orchids of Venezuela: An Illustrated Field Guide. Allston, MA: Botanical Museum of Harvard University. (3) Finn N. Rasmussen. 1982. Opera Botanica 65. Copenhagen.

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ramicaul: an aerial, unthickened, cylindrical (rarely winged or channeled) stem with 1 terminal leaf (rarely with several distichous leaves, e.g., Frondaria Luer), found only in the subtribe Pleurothallidinae. resupination: the process of twisting (or bending) the ovary and pedicel during development to position the lip on the lower side of the flower. rostellum: a portion of the stigma that aids in gluing the pollinia to the pollinating agent; the tissue that separates the anther from the fertile stigma, sometimes beak-like. sectile: the condition in which soft, granular pollinia are subdivided into small packets, these usually connected by an elastic material. stelidia: lateral or sideways projections of the column (usually toward its apex), often also called “column wings,” most commonly in the Pleurothallidinae. stipe: either a tegula or a hamulus (any pollinium stalk composed of sterile tissue). synsepal: the blade formed by the union of the lateral sepals. tegula: a pollen stalk derived from the surface tissues of the clinandrium and column. velamen: water-absorbing layer of dead cells. viscidium, pl. viscidia: a viscid part of the rostellum that is clearly defined and removed with the pollinia as a unit and serves to attach the pollinia to an insect or other pollinating agent. Key to the Subkeys to the Genera of Orchidaceae 1.

1.

2(1).

2.

Plants climbers with ± fleshy stems lacking pseudobulbs or any differentiation in function or thickness; leaves distichous, fleshy, with convolute vernation, or absent .................................................................. 152. Vanilla Plants not climbers, stems elongated or abbreviated, bearing pseudobulbs or not; leaves various, either conduplicate with duplicative vernation (and ± fleshy) or either plicate or with herbaceous texture and with convolute vernation ............................................................................... 2 Column with 2 fertile anthers; ovary 1- or 3-locular, with axile placentation; flowers with a distinct combination of features: lip slipper-like, i.e., deeply sacciform with the outer margins folded inward forming a central opening to the cup and a basal opening with 2 exits at each side of the short column with 2 anthers; petals linear-filiform to narrowly elliptical, often with meristematic apices that keep growing after the flower opens; lateral sepals fused into a synsepal usually hidden under the slipper-like lip ...................................................... Subkey 1, page 210 Column with 1 fertile anther; ovary 1-locular, with parietal placentation; flowers without the above combination of features: lip various, but if deeply sacciform, then margins erect or folded outward, never forming a central cup and a basal 2-way exit; petals various but never apically meristematic nor growing and elongating after the flower opens; lateral sepals free or fused, usually ± conspicuous ................................... 3

O R C H I D A C E A E 209

3(2).

3.

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

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

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

9. 10(9).

10. 11(5). 11. 12(11). 12. 13(11). 13. 14(13). 14. 15(13). 15.

Plants autotrophic or sometimes saprophytic; leaves absent or inconspicuous at anthesis; secondary stem not modified into pseudobulbs or corms ....................................................................................Subkey 2, page 210 Plants always autotrophic; leaves conspicuous at anthesis or, if absent, then the secondary stem modified into pseudobulbs; secondary stem modified into pseudobulbs or corms, or not modified ........................... 4 Plants with terete, subterete, or laterally compressed (equitant) leaves .................................................................................... Subkey 3, page 211 Plants with leaves not as above ................................................................. 5 Plants with terminal inflorescences, the inflorescence arising from the apex of the last internode ...................................................................... 6 Plants with lateral inflorescences, the inflorescence not arising from the apex of the last internode, or inflorescence arising from the rhizome .............................................................................................................. 11 Stem modified into pseudobulbs or corms .................... Subkey 4, page 213 Stem not modified into pseudobulbs or corms .......................................... 7 Secondary stem absent; leaves basal and/or rosette-like, not articulate; inflorescence usually an erect raceme; pollen always soft, friable, never waxy ...........................................................................Subkey 5, page 214 Secondary stem elongate, with internodes; leaves not basal, rosette-like or not, articulate or not; inflorescence variable; pollen soft or waxy ... 8 Leaves not articulate, persisting after they wilt or falling with the sheath ....................................................................................Subkey 6, page 216 Leaves articulate, only the sheath persisting after the leaf falls ............. 9 Ovary articulate with the pedicel (the pedicel may persist after the flower drops); secondary stems 1-foliate; pollinia waxy, usually clavate ....................................................................................Subkey 7, page 217 Ovary not articulate with the pedicel; secondary stems 1-foliate to multifoliate; pollinia soft or waxy ................................................................ 10 Leaves conduplicate, without prominent ribs, folded only at the midline, roughly V-shaped in cross section; pollen soft or waxy ........................... ....................................................................................Subkey 8, page 220 Leaves plicate, with several conspicuous ribs, the leaf blade usually folded at each rib; pollen waxy, not friable .......................... Subkey 9, page 221 Stems simple, not modified into pseudobulbs or corms .......................... 12 Stems modified into pseudobulbs or corms ............................................. 13 Leaves plicate .............................................................. Subkey 10, page 221 Leaves conduplicate ..................................................... Subkey 11, page 222 Leaves plicate ........................................................................................... 14 Leaves conduplicate ................................................................................. 15 Pseudobulb homoblastic (consisting of several internodes) ....................... ................................................................................. Subkey 12, page 223 Pseudobulb heteroblastic (consisting of a single swollen internode) ................................................................................. Subkey 13, page 224 Inflorescence of only 1 flower ..................................... Subkey 14, page 227 Inflorescence of few to many flowers.......................... Subkey 15, page 227

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Subkey 1 to the Genera of Orchidaceae Characters in common: Flowers with slipper-like lip; column with 2 fertile anthers; ovary 3-locular, with axile placentation. Genera (2): Phragmipedium, Selenipedium. 1.

1.

Plants stemless, cespitose, rhizomatous, with crowded, basal leaves; leaves conduplicate; ovary 3-locular; abscission layer between ovary and pedicel present ........................................................... 107. Phragmipedium Plants with well-developed, reed-like secondary stem, with distichous, spaced leaves; leaves plicate; ovary 1-locular; abscission layer between pedicellate ovary and pedicel absent ............................ 133. Selenipedium

Subkey 2 to the Genera of Orchidaceae Characters in common: Plants saprophytic or autotrophic; secondary stem not modified into pseudobulbs; leaves absent or inconspicuous at anthesis. Genera (14): Brachystele (part), Campylocentrum (part), Cleistes (part), Duckeella (part), Habenaria (part), Lyroglossa (part), Prescottia (part), Pterichis (part), Sacoila, Sarcoglottis (part), Stellilabium (part), Uleiorchis, Vanilla (part), Wullschlaegelia. 1. 1. 2(1). 2. 3(2). 3. 4(2). 4.

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

6.

Plants climbers; bracts leaf-like, succulent ......... 152. Vanilla (V. penicillata) Plants terrestrial or epiphytic, never climbers; leaves absent or if leaf-like bracts present, then not succulent ........................................................ 2 Plants epiphytic, minute, stemless or nearly so; inflorescences shorter than to ± equal to root length; roots often photosynthetic ................... 3 Plants terrestrial, minute to relatively large; inflorescences longer than roots; roots not photosynthetic .............................................................. 4 Column basally without furcate hairs; lip with a well-developed spur .................................................................................... 19. Campylocentrum Column basally with thick, apically 2–5-furcate hairs; lip spurless .......................................................................................... 140. Stellilabium Flowers with a prominent spur or mentum noticeable from the outside ................................................................................................................ 5 Flowers without spur or spur hidden (embedded within ovarian tissue contained in the decurrence on ovary and fused bases of lateral sepals, hence not noticeable from outside) ........................................................ 9 Petals 2-lobed; lip deeply 3-lobed; anther with 2 well-separated, sessile thecae, hence column appears to have 2 anthers ................ 62. Habenaria Petals simple; lip simple or shallowly lobed; anther with thecae closely set, hence column appearing monandrous (with one anther) .............. 6 Inflorescence few-flowered; sepals and petals fused into a cup or hood open ventrally; plants basally with horizontal underground tuber and filamentous roots above it ......................................................... 151. Uleiorchis Inflorescence many-flowered; sepals and petals free or only fused basally; plants with fasciculate, fusiform roots .................................................. 7

O R C H I D A C E A E 211

7(6). 7.

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

9.

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

12(10). 12.

13(12).

13.

Plants with stems and external surface of floral segments pubescent; flowers < 5 mm long, pale cream or white, not resupinate ..... 154. Wullschlaegelia Plants with stems and external surface of floral segments glabrous; flowers > 10 mm long, usually bright red, more rarely pink or white .................................................................................................. 127. Sacoila Flowers not resupinate (i.e., with the lip uppermost and oriented toward inflorescence apex) ................................................................................. 9 Flowers resupinate (i.e., with lip lowermost and oriented toward inflorescence base) ........................................................................................... 10 Lip slipper-like, with retrorse glands at the lip base, longer than wide; petals conspicuously narrower than dorsal sepal; dorsal sepal to 2.5 mm long ....................................................................................... 116. Prescottia Lip not slipper-like, without retrorse glands at the lip base, broader than long; petals subequal to dorsal sepal; dorsal sepal 8–10 mm long ................................................................................................ 120. Pterichis Pedicellate ovary and sepals externally glabrous ................................... 11 Pedicellate ovary and sepals externally pubescent ................................ 12 Plants drying black; column < 1/2 the length of lip, the apex biauriculate, the base not attenuate; pollinia 4; flowers bright yellow ..... 44. Duckeella Plants drying dark brown or brownish; column about 1/2 as long or longer than lip length, apically entire or lobed, not auriculate, attenuate basally; pollinia 2; flowers pink, red, or purple ............................ 27. Cleistes Flowers minute; dorsal sepal 2–4 mm long; lip 2–3.5 mm long, widest toward base; column very short, widened toward apex ....... 14. Brachystele Flowers small; dorsal sepal 6–12 mm long; lip 6–17 mm long, widest toward the middle; column not elongate, slightly or not widened toward the apex ................................................................................................ 13 Pedicellate ovary ± equal to dorsal sepal length; lateral sepals not decurrent on ovary; lip not deflexed apically, constricted about the middle, basally cuneate; column footless .......................................... 82. Lyroglossa Pedicellate ovary much longer than dorsal sepal length; lateral sepals decurrent on ovary; lip apically deflexed, constricted below middle and again at the apical 1/4, basally long-sagittate; column with a long foot ........................................................................................... 127. Sarcoglottis

Subkey 3 to the Genera of Orchidaceae Characters in common: Plants with leaves terete, subterete, or laterally compressed. Genera (22): Brassavola, Cryptocentrum, Dunstervillea, Eloyella, Erycina, Ionopsis (part), Jacquiniella, Lockhartia, Macroclinium, Maxillaria (part), Octomeria (part), Ornithocephalus, Plectrophora, Pleurothallis (part), Polyotidium, Quekettia, Reichenbachanthus, Scuticaria, Stictophyllorchis, Trichocentrum (part), Trisetella (part), Trizeuxis. 1. 1.

Leaves laterally compressed ...................................................................... 2 Leaves terete or subterete ....................................................................... 12

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O RCHIDACEAE

2(1). 2. 3(2). 3. 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(1). 12. 13(12). 13. 14(13). 14. 15(14). 15. 16(15). 16. 17(16). 17. 18(12). 18. 19(18).

Inflorescence terminal ............................... 70. Jacquiniella (J. steyermarkii) Inflorescence axillary or lateral ................................................................. 3 Column very elongate ......................................................... 84. Macroclinium Column abbreviate ..................................................................................... 4 Plants without pseudobulbs ....................................................................... 5 Plants with pseudobulbs ............................................................................ 6 Flowers not in crowded terminal racemes; leaves conspicuously spotted with red; anther erect; column with 2 introrse, obtuse stigmatic arms ................................................................................... 142. Stictophyllorchis Flowers produced successively in crowded terminal racemes; leaves not spotted with red; anther partially incumbent; column without 2 introrse, obtuse stigmatic arms ................................................ 150. Trizeuxis Rear of the flower with a spur-like structure formed by the base of the lip and the free lateral sepals ............................................... 110. Plectrophora Rear of the flower without a spur-like structure ...................................... 7 Plants elongate, not fan-shaped ................................................................ 8 Plants short, fan-shaped ............................................................................ 9 Petals and sepals fleshy; lip purple; pollinia 4 .. 87. Maxillaria (M. equitans) Petals and sepals membranaceous to subfleshy; lip yellow or white; pollinia 2 .................................................................................... 79. Lockhartia Pollinia 2; flowers yellow .............................................................. 53. Erycina Pollinia 4; flowers white, cream, or pinkish ............................................ 10 Flowers subtended by large, conspicuous bracts; lip with a triangular spur from the base .................................................................... 45. Dunstervillea Flowers subtended by inconspicuous bracts; lip without a spur ............ 11 Column winged on each side at the base, with a curled ligule at the junction with the lip ......................................................................... 47. Eloyella Column wingless, without a curled ligule at the junction with the lip ................................................................................... 100. Ornithocephalus Inflorescences terminal ............................................................................ 13 Inflorescence lateral ................................................................................. 18 Pedicellate ovary ≥ 4 cm long; petals and sepals ≥ 2.5 cm long .............................................................................................. 16. Brassavola Pedicellate ovary < 2 cm; petals and sepals < 2 cm ................................ 14 Secondary stems distichously foliate ..................................... 70. Jacquiniella Secondary stems apically 1-foliate .......................................................... 15 Secondary stems proliferous, pendulous; pollinia 8 .. 122. Reichenbachanthus Secondary stems neither proliferous nor pendulous; pollinia 2 or 8 ...... 16 Flowers fasciculate on short peduncles; pollinia 8 .................. 94. Octomeria Flowers either racemose or solitary on long peduncles; pollinia 2 ........ 17 Synsepal without subapical, filiform tails; flowering in each raceme occurs almost simultaneously (gregariously) ......... 111. Pleurothallis (P. galeata) Synsepal with 2 subapical, filiform tails in about the apical 1/4; flowering occurs a single flower at a time (successively) ..................... 149. Trisetella Petals and sepals > 2.5 cm long, yellow with purple blotches; inflorescenses (1)2- or 3-flowered ................................................... 132. Scuticaria Petals and sepals < 1.5 cm, variously colored; inflorescences a single flower or a few- to many-flowered raceme .......................................... 19 Inflorescence a single flower .................................................................... 20

O R C H I D A C E A E 213

19. Inflorescence a few- to many-flowered raceme or panicle ...................... 21 20(19). Peduncle much longer than pedicellate ovary; flowers spurred ................ ........................................................................................ 34. Cryptocentrum 20. Peduncle subequal to shorter than pedicellate ovary; flowers not spurred ........................................................................... 87. Maxillaria (M. uncata) 21(19). Flowers medium-sized to large; perianth segments well expanded .......... ................................................................. 144. Trichocentrum (T. cebolleta) 21. Flowers minute; perianth segments parallel, scarcely open .................. 22 22(21). Column with 2 pairs of ear-like projections; flowers orange ...................... .......................................................................................... 113. Polyotidium 22. Column without 2 pairs of ear-like projections; flowers pink or greenish yellow ................................................................................................... 23 23(22). Lip simple ............................................................. 68. Ionopsis (I. satyrioides) 23. Lip 3-lobed ................................................................................. 121. Quekettia

Subkey 4 to the Genera of Orchidaceae Characters in common: Plants with pseudobulbs; inflorescence terminal. Genera (16): Cattleya, Caularthron, Dimerandra (part), Encyclia, Epidendrum (part), Galeandra (part), Hexisea (part), Liparis, Malaxis, Nidema, Orleanesia (part), Pinelianthe, Polystachya (part), Prosthechea (part), Scaphyglottis (part), Schomburgkia. 1. 1. 2(1). 2. 3(2). 3. 4(1). 4. 5(4). 5. 6(5). 6. 7(4). 7. 8(7).

Leaves plicate ............................................................................................. 2 Leaves conduplicate ................................................................................... 4 Leaves narrow, coriaceous; lip with a distinct spur; pollinia 2, attached by a conspicuous tegula ............................................................. 57. Galeandra Leaves broad, membranaceous; lip without a distinct spur; pollinia 2 or 4, not attached by a tegula ........................................................................ 3 Inflorescence an umbel; column usually < 2 mm long ................. 85. Malaxis Inflorescence a raceme; column usually ≥ 2 mm long .................. 78. Liparis Column with a conspicuous foot ................................................................ 5 Column without a foot ............................................................................... 7 Pseudobulbs short, ovoid, not reduced to a stalk at the base; pollinia spherical to subspherical .................................................. 114. Polystachya Pseudobulbs short or elongate, fusiform, reduced to a stalk at the base; pollinia usually laterally flattened ........................................................ 6 Shoots not prolific (stems not arising from the apex of other stems); flowering pseudobulbs with 4–several leaves ................................ 99. Orleanesia Shoots prolific; flowering stems with 1 or 2 leaves ........... 129. Scaphyglottis Plants very small, the leaves < 1 cm long and the inflorescence elongate ........................................................................................... 108. Pinelianthe Plants large, the leaves > 1 cm long or, if less, then the inflorescence sessile ..................................................................................................... 8 Inflorescence long pedunculate with a short rachis bearing a sub-umbel of several to many (6–25) simultaneously opening flowers; sepals strongly undulate; pollinia 8 ...................................................... 131. Schomburgkia

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8. 9(8). 9. 10(9). 10. 11(9). 11. 12(11). 12. 13(12). 13. 14(13). 14. 15(14).

15.

16(14). 16.

Inflorescence various but never as above; sepals not undulate; pollinia 4 or 6 ....................................................................................................... 9 Shoots prolific (producing apically one or more new pseudobulbs instead of these only produced from the rhizome) .......................................... 10 Shoots not prolific (pseudobulbs borne only from the rhizome) ............. 11 Flowers reddish orange; lip abruptly bent at junction with the column .................................................................................................... 64. Hexisea Flowers greenish, yellowish, or lavender; lip not bent at junction with the column ............................................................................ 129. Scaphyglottis Pseudobulbs hollow, often housing ant colonies; lip with 2 prominent, horn-like, hollow, basal calli .............................................. 22. Caularthron Pseudobulbs rarely hollow, seldom housing ant colonies; lip without 2 prominent, horn-like, hollow, basal calli .......................................... 12 Column with 2 prominent externally keeled apical lobes; anther completely hidden within these 2 lobes ................................... 40. Dimerandra Column without 2 keeled lobes; anther not hidden ................................ 13 Column connate with the lip its entire length ...................... 50. Epidendrum Column completely free (or fused only at its basal 1/3 to the ventral face of the column) .......................................................................................... 14 Lip with calli or keels ............................................................................... 15 Lip without keels or calli ......................................................................... 16 Flowers resupinate; lip totally free from the ventral face of the column; inflorescences lacking a spathe, often paniculate; central lobe of the clinandrium not conspicuously different from the other two; fruits ellipsoid, not 3-winged .................................................................... 49. Encyclia Flowers not resupinate (in all flora area species); lip fused to the ventral face of the column in the basal 1/3–1/2; inflorescences emerging from a spathe, racemose or 1-flowered (teratologically paniculate in robust specimens of some taxa); midtooth appendage (central lobe of the clinandrium) fleshy, knob-like, obtuse or truncate; fruit 3-winged or -angled ............................................................................... 117. Prosthechea Flowers large (dorsal sepal > 5 cm long) ...................................... 21. Cattleya Flowers small (dorsal sepal to 1.5 cm long) ................................. 92. Nidema

Subkey 5 to the Genera of Orchidaceae Characters in common: Secondary stem absent or inconspicuous; leaves basal and/or rosette-like, not articulate; inflorescence usually an erect raceme; pollen always soft, friable, never waxy. Genera (19): Aracamunia (part), Baskervilla, Brachystele (part), Cranichis, Cyclopogon, Discyphus, Duckeella (part), Eltroplectris, Eurystyles, Gomphichis, Habenaria (part), Helonoma (part), Lyroglossa (part), Mesadenella, Pelexia, Ponthieva, Prescottia (part), Pterichis (part), Sarcoglottis (part). 1.

Plants epiphytic, minute (leaves to 4 × 1 cm), margins of leaves and floral and inflorescence bracts ciliate or ciliolate; inflorescences shortpedunculate, pendent, capitate ............................................ 56. Eurystyles

O R C H I D A C E A E 215

1.

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

4(3). 4. 5(4). 5. 6(5). 6. 7(5). 7. 8(7). 8. 9(8). 9. 10(4). 10. 11(10). 11. 12(11). 12. 13(12). 13. 14(13). 14. 15(14). 15. 16(15).

16.

Plants usually terrestrial (very rarely humus epiphytes), medium- to large-sized (leaves normally > 4 × 1 cm; if leaves within that range, then plants terrestrial with an erect, long-pedunculate inflorescence); margins of leaves and floral or inflorescence bracts entire or at most glandular, rarely coarsely pilose; inflorescences usually long-pedunculate, erect racemes or spikes ................................................................. 2 Anthers incumbent, bending down to become ± operculate on the apex of the column; flowers bright yellow ......................................... 44. Duckeella Anthers remaining erect, not operculate on the apex of the column; flowers usually white of greenish, rarely greenish yellow .......................... 3 Anther projecting beyond the rostellum; viscidium, if present, usually at the base of the pollinia; plants often with root tubers ........ 62. Habenaria Anther subequal to the rostellum; viscidium at the apex of the anther and attached to the apex or the middle of the pollinia or caudicle; plants without root tubers ................................................................................ 4 Flowers not resupinate .............................................................................. 5 Flowers resupinate. .................................................................................. 10 Petals and lip adnate to the column to a distance from the base ............. 6 Petals and lip free from the column .......................................................... 7 Lip saccate at its base, with replicate lateral lobes .................. 8. Baskervilla Lip not saccate, without replicate lateral lobes ...................... 115. Ponthieva Lip margin conspicuously glandular ......................................... 120. Pterichis Lip margin not glandular .......................................................................... 8 Column bent ............................................................................ 59. Gomphichis Column straight ......................................................................................... 9 Sepals free to base, flat to cochleate, usually with reticulate, colored veins ................................................................................................ 32. Cranichis Sepals connate in the lower 1/3–1/4 to form a tube that encloses the lower half of the lip, the lip calceolate .......................................... 116. Prescottia Stigmas 2, cup-shaped; plant with only a single, sessile, cordate-subrotund leaf ......................................................................................... 42. Discyphus Stigma 1; plant not with a single, sessile cordate-subrotund leaf ......... 11 Column ballooned out in front due to inflated clinandrium ..... 63. Helonoma Column never ballooned .......................................................................... 12 Leaf sheaths with 1(2) erect, strap-shaped structures covered by multicellular, short-pedunculate, globose hairs ............................... 4. Aracamunia Leaf sheaths without strap-shaped structures ....................................... 13 Rostellum very short ............................................................... 14. Brachystele Rostellum prominent ............................................................................... 14 Rostellum strap-like, soft ......................................................................... 15 Rostellum subulate, rigid ......................................................................... 17 Column foot not prominent...................................................... 35. Cyclopogon Column foot prominent, attached to the side of the ovary ..................... 16 Column foot embedded internally in the ovary with the connate lateral sepals forming an internal nectary without a clear line of adnation on the outside ......................................................................... 127. Sarcoglottis Column foot attached to the outside of the ovary with a clear line of adnation, lateral sepals basally connate into a ventricose, saccate, or tubular spur ............................................................................................ 105. Pelexia

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17(14). 17. 18(17). 18.

Lip conspicuously pandurate (fiddle-shaped) .......................... 82. Lyroglossa Lip not pandurate .................................................................................... 18 Flowers with a spur; viscidium at the apex of the pollinia .. 48. Eltroplectris Flowers without a spur; viscidium at the middle of the pollinia ................ ............................................................................................ 88. Mesadenella Subkey 6 to the Genera of Orchidaceae

Characters in common: Plants without pseudobulbs; secondary stem elongate; leaves not articulate; inflorescence terminal; plants terrestrial; pollen soft. Genera (13): Aracamunia (part), Aspidogyne, Cleistes (part), Duckeella (part), Epistephium, Erythrodes, Habenaria (part), Helonoma (part), Ligeophila, Palmorchis, Psilochilus, Triphora, Xerorchis. 1. 1. 2(1). 2. 3(2).

3.

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

5.

6(5).

6.

Stems woody or thickened and cane-like, frequently branching above the base; roots usually wiry ......................................................................... 2 Stems herbaceous (rarely somewhat thickened basally); roots herbaceous ................................................................................................................ 4 Leaves conspicuously plicate (with 1–3 lateral veins parallel to the midvein) ............................................................................. 103. Palmorchis Leaves not plicate, either grass-like or ± fleshy ....................................... 3 Stems straight or at least not zigzag; plants drying black; leaves subfleshy, 2–4 times as long as wide, conspicuously reticulate-veined; flowers large and conspicuous (sepals at least 2 cm long, usually at least 3.5–6 cm long), purple or lilac, rarely white, with a conspicuous calyculus beneath the perianth ........................................ 51. Epistephium Stems conspicuously zigzag in the upper half; plants dying not black, most often dark brown or greenish brown; leaves grass-like, 4.5–10 times longer than wide, not reticulate-veined (if somewhat reticulate, as in Helonoma bifida and H. chiropterae, then leaves < 4 cm long); flowers small and inconspicuous, sepals to 11 mm long, dull-yellow, greenish, or white, lacking a calyculus beneath the perianth ............ 155. Xerorchis Flowers with conspicuous spurs produced from the base of the lip ......... 5 Flowers lacking spurs at the base of the lip .............................................. 8 Plants without a basal creeping portion, bearing 1 or 2 corms or tubers (“stem tuberoids”) underground; lip and petals usually deeply 3-lobed at base; anther erect, basally firmly united with the column, not deciduous, the locules bulging out on an elongated channel ......... 62. Habenaria Plants with a conspicuous basal creeping portion, producing roots along its length, lacking tubers or corms (but roots somewhat fleshy in some genera); lip simple or 3-lobed above middle; anther versatile, eventually caducous, the locules not bulging out on an elongated channel ... 6 Lip with a pair of fleshy plates or ridges at base leading to the entrance to the spur; rostellum articulate, triggered, porrect in natural position when pollinia are attached, becoming erect after removal of pollinia; stigmas confluent into one, transversely elliptic body ........ 77. Ligeophila Lip without a pair of fleshy plates or ridges at base leading to the entrance to the spur; rostellum neither articulate nor triggered; stigmas

O R C H I D A C E A E 217

2, distinct, approximate ......................................................................... 7 Rostellum entire; column elongate, with a distinct, ± cylindrical stalk ................................................................................................ 6. Aspidogyne 7. Rostellum deeply bipartite; column short, sessile to subsessile, without a distinct stalk ........................................................................ 54. Erythrodes 8(4). Leaf sheaths with 1(2) erect, strap-shaped structures covered by multicellular, short-stalked, globose hairs; flowers greenish white, the sepals fused into a tube for 3/4 of their length ................................ 4. Aracamunia 8. Leaf sheaths without strap-shaped structures covered by multicellular hairs; flowers variously colored, the sepals free or fused below the basal 1/2 ................................................................................................... 9 9(8). Plants small with leaves mostly basal; petioles conspicuous; lip with basal retrorse lobules; column ballooned out in front due to an inflated clinandrium ............................................................................ 63. Helonoma 9. Plants small or large, with leaves distributed mostly along the stem (or if somewhat basal and rosulate, then larger than 5 cm long); petioles absent or inconspicuous; lip without basal retrorse lobules; column never ballooned due to an inflated clinandrium ........................................... 10 10(9). Plants mostly from the forest floor; rhizomes ± well developed, creeping; anthers erect on top of the column; flowering gregarious (i.e., all or most members of the population produce fugacious flowers on the same days, then rest for several others) ....................................................... 11 10. Plants mostly from open, sunny places like savannas, tepui-scrubs, or bogs; rhizomes lacking; anthers incumbent, bending down to become ± operculate on the apex of the column; flowering sequential, (i.e., at flowering time most members of the population have buds and flowers at anthesis which open ± randomly as they mature on the plant) .... 12 11(10). Pollinia 2; column winged ....................................................... 118. Psilochilus 11. Pollinia 4; column terete ............................................................ 148. Triphora 12(10). Plants drying black; column < 1/2 the length of lip, the apex biauriculate, the base not attenuate; pollinia 4; flowers bright yellow ..... 44. Duckeella 12. Plants drying dark brown or brownish; column about 1/2 as long or longerthan lip length, apically entire or lobed, not auriculate, attenuate basally; pollinia 2; flowers pink, red, or purple, rarely cream or pale yellow (if cream or pale yellow, then leaves inconspicuous or reduced to bract-like structures) ................................................................. 27. Cleistes 7(6).

Subkey 7 to the Genera of Orchidaceae Characters in common: Ovary articulate with the pedicel; special secondary stems (ramicauls) 1-foliate; leaves articulate; inflorescences 1-flowered or racemose; ramicauls sometimes proliferous. Genera (16): Barbosella, Brachionidium, Dryadella, Lepanthes, Lepanthopsis, Masdevallia, Myoxanthus, Octomeria (part), Ophidion, Platystele, Pleurothallis (part), Restrepiopsis, Scaphosepalum, Stelis, Trichosalpinx, Trisetella (part). [See comments on generic taxa recognized after the generic description of Pleurothallis, p. 504.]

218

O RCHIDACEAE

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

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

13.

Ramicauls clothed by lepanthiform sheaths (see figure 200) ................... 2 Ramicauls naked or, if clothed by sheaths, these not lepanthiform ........ 7 Inflorescence shorter than or ± equal to the subtending leaf .................. 3 Inflorescence longer than the subtending leaf .......................................... 4 Petals broader than long; anther apical or dorsal; stigma apical; flowers usually opening 1 or 2 at a time ............................................ 74. Lepanthes Petals longer than broad; anther apical or subapical; stigma ventral; flowers usually opening several or all simultaneously ....... 146. Trichosalpinx All 3 sepals connate basally to 2/3 their length .................. 146. Trichosalpinx Sepals free or, if connate, only the lateral ones so .................................... 5 Pedicel shorter than to 2 times as long as ovary and floral bract; column wider than long .................................................................75. Lepanthopsis Pedicel at least 3 times as long as ovary and floral bract; column longer than wide ................................................................................................ 6 Petals and lip at least 2 times as long as wide; column longer than broad ........................................................................................ 146. Trichosalpinx Petals and lip wider than long to as long as wide, or rarely to 1.5 times longer than wide; column cylindric ....................................... 74. Lepanthes Plants Peperomia-like, i.e., with small (5–15 mm long) orbicular to broadly elliptic leaves, longer than the ramicaul and appressed to the substrate; rhizome creeping, appressed to the substrate ..................... 8 Plants not Peperomia-like, with leaves not appressed to the subtrate; rhizome abbreviated to elongated ............................................................ 12 Column wider than long, apically broadly winged; lip simple ... 109. Platystele Column longer than wide, apically wingless or with small, subapical auricles; lip 3-lobed or simple .................................................................... 9 Leaves verruculose on both surfaces; lateral sepals almost totally connate into a deeply concave synsepal; pollinia 6 .. 111. Pleurothallis (P. hexandra) Leaves smooth; lateral sepals free or connate only to 3/4 of their length; pollinia 2, 4, or 8 .................................................................................. 10 Pollinia 4; lip articulate to the column by a ball-and-socket mechanism; dorsal sepal 1/4–1/3 as wide as the synsepal ............................ 7. Barbosella Pollinia 2 or 8; lip merely hinged to the column; dorsal sepal to 1/2 as wide as synsepal or lateral sepals free ........................................................ 11 Flowers white with red in column; petals subequal to sepals; lip < 2 times as long as wide; pollinia 8 ..................................................... 94. Octomeria Flowers purple or wine red; petals distinctly shorter than the sepals; lip ≥ 3 times as long as wide; pollinia 2 ............................... 111. Pleurothallis Ramicauls triquetrous (3-winged) or sharply compressed at least in the upper 1/2 ........................................................................... 111. Pleurothallis Ramicauls terete or slightly compressed ................................................ 13 Inflorescences originating from the base of a well-developed ramicaul or from the rhizome, 1–3-flowered, shorter than ramicaul ........................ ............................................................... 111. Pleurothallis (P. lappiformis) Inflorescences originating from the apex or near the apex of the ramicaul, if inflorescences ± basal, then many-flowered and longer than ramicaul (ramicauls sometimes very short and the inflorescences look basal) .............................................................................................................. 14

O R C H I D A C E A E 219

14(13). Tails from the lateral sepals originating not from the morphological apex of the sepals but laterally in the apical 1/4 ........................... 149. Trisetella 14. Tails from the lateral sepals absent or, if present, originating from the apex of sepals ....................................................................................... 15 15(14). Inflorescences many-flowered, sometimes flowering successively and appearing 1-flowered but then the rachis with flower scars .................. 16 15. Inflorescences 1-flowered ......................................................................... 21 16(15). Lateral sepals with a callose, cushion-like pad at the apex ....................... ..................................................................................... 128. Scaphosepalum 16. Lateral sepals ecallose at the apex .......................................................... 17 17(16). All 3 sepals connate at their apices to form a laterally 2-windowed synsepal; column elongate, arched, basally terete, winged in the apical 1/3 ................................................................................................. 98. Ophidion 17. Sepals free or variously connate but never forming a laterally 2-windowed synsepal; column various ..................................................................... 18 18(17). Petals transverse, i.e., much wider than long ................................ 139. Stelis 18. Petals longer than wide or, at most, as long as wide .............................. 19 19(18). Column short, wider than long, ± membranaceous; stigma 2-lobed, transverse .............................................................................. 109. Platystele 19. Column ± elongate, longer than wide; stigma 1- or 2-lobed ................... 20 20(19). Petals longitudinally and marginally callose, the lower margin developed basally into a ± retrorse tooth ........................................... 86. Masdevallia 20. Petals not callose, without a basally retrorse tooth ........... 111. Pleurothallis 21(15). Ramicauls covered with pilose or scabrous sheaths, especially toward base ....................................................................................................... 22 21. Ramicauls almost naked to covered with smooth, glabrous sheaths ..... 23 22(21). Pollinia 2; sepals externally pilose ........................................ 91. Myoxanthus 22. Pollinia 4; sepals externally glabrous ................................. 123. Restrepiopsis 23(21). Leaves deeply cordate basally ............................................ 111. Pleurothallis 23. Leaves basally acute, obtuse, or rounded, but never cordate ................. 24 24(23). Lateral sepals essentially free ................................................................. 25 24. Lateral sepals connate 1/2 or more of their length .................................. 26 25(24). Petals with an obliquely truncate apex, subquadrate in shape, dissimilar to sepals, originating from the column foot; lip basally clawed and above expanded into a basally sagittate blade; pollinia 2; lateral sepals with a basal, transverse callose fold ..................................... 43. Dryadella 25. Petals with rounded, obtuse, acute, or acuminate apex, not subquadrate in shape, usually similar to the sepals in shape and size, originating from column base; lip sessile; pollinia (6–)8; lateral sepals ecallose ............................................................................................... 94. Octomeria 26(24). Petals about the same size and shape as dorsal sepal; inflorescence with a conspicuous bract at approximately its middle ........... 13. Brachionidium 26. Petals smaller than and dissimilar to dorsal sepal; inflorescence without a conspicuous bract at approximately its middle .................................. 27 27(26). Ramicauls shorter than leaves; petals callose along the lower margin; lateral sepals apically extended into short to long tails ....... 86. Masdevallia 27. Ramicauls longer than to ± equal to leaves; petals ecallose; lateral sepals not extended into tails or tails short ................................................... 28

220

O RCHIDACEAE

28(27). Pedicels, bracts, and outer face of sepals covered with mounds of hair ............................................................................................ 91. Myoxanthus 28. Pedicels, bracts, and outer face of sepals glabrous or glabrescent ......................................................................................... 111. Pleurothallis

Subkey 8 to the Genera of Orchidaceae Characters in common: Plants with secondary stems not conspicuously thickened into a pseudobulb or corm, with an elongate stem and internodes; ovary not articulate with the pedicel; leaves conduplicate; pollen waxy, not friable. Genera (8): Dimerandra (part), Epidendrum (part), Hexisea (part), Isochilus, Orleanesia (part), Polystachya (part), Prosthechea (part), Scaphyglottis (part). 1. 1. 2(1).

2.

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

4.

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

6.

Stems superposed, forming chains (new ones on top of the older ones); leaves only at the tip of the individual stems ....................................... 2 Stems not superposed; leaves placed either along or only at the tip of the stems. ..................................................................................................... 3 Lip fused to the base of the column and then abruptly bent up against the column and down again; flowers bright red or orange; pollinia always 4 ...................................................................................... 64. Hexisea Lip hinged to a ± well-developed column foot, not bent at base; flowers usually greenish or yellowish, often tinged with purple or maroon; pollinia 4 or 6 ................................................................. 129. Scaphyglottis Lip firmly fused to the column at the apex or nearly so; pollinia with distinct semiliquid viscidium .............................................50. Epidendrum Lip free or only fused at the base of the column; pollinia naked or with caudicles, always without a distinct semiliquid viscidium. ................. 4 Flowers pink or pale purple, widely opening and appearing flat; callus with several to many small, shallow, transverse keels; column with conspicuous vertical, parallel lobes at apex . .................................. 40. Dimerandra Flowers of various colors, usually greenish or yellowish; callus various but never composed of shallow, transverse keels; column without vertical, parallel lobes at apex . .................................................................... 5 Leaves (4)5–25, distichously distributed along the upper 1/2 of the stem; flowers always resupinate ..................................................................... 6 Leaves 1–4, placed only on top of the stems; flowers usually not resupinate ................................................................................................................ 7 Leaves < 0.5 cm wide, coriaceous; plants from elevations above 800 m; column at least 3 times as long as wide, with an inconspicuous foot at base; lateral sepals fused in the lower 1/2; flowers tubular, red-carmine or red-pink, secund, several flowering simultaneously; lip acute, flat or shallowly concave ..................................................................... 69. Isochilus Leaves > 1 cm wide, fleshy-coriaceous; plants from elevations below 300 m; column as long as to about 2 times as long as wide, with a definite foot; lateral sepals free; flowers widely opening, greenish or yellowish,

O R C H I D A C E A E 221

7(5).

7.

flowering successively in a distichous or subspiral fashion; lip obtuse, sharply convex ................................................................. 99. Orleanesia Petals and sepals > 8 mm long; column without a foot; callus of the lip naked or hairy, never with loose, mealy pseudopollen ............................... .................................................................. 117. Prosthechea (P. crassilabia) Petals and sepals < 7 mm long; column with a well-developed column foot; callus of the lip covered with loose, mealy pseudopollen ........................................................................................... 114. Polystachya

Subkey 9 to the Genera of Orchidaceae Characters in common: Leaves plicate; pollen soft or waxy. Genera (3): Elleanthus, Galeandra (part), Sobralia. 1. 1. 2(1). 2.

Column with a distinct foot; pollinia 2, waxy, attached to a conspicuous tegula ..................................................................................... 57. Galeandra Column without a distinct foot; pollinia 8, soft, granular to subceraceous, not attached to a tegula ......................................................................... 2 Dorsal sepal < 2 cm long; base of the lip saccate to subsaccate, with 2 prominent calli .................................................................. 46. Elleanthus Dorsal sepal > 2 cm long; base of the lip not saccate or subsaccate, without calli ......................................................................................... 136. Sobralia

Subkey 10 to the Genera of Orchidaceae Characters in common: Plants without pseudobulbs; leaves plicate, fan-like; inflorescence lateral, 1-flowered. Genera (5): Bollea, Chaubardiella, Cochleanthes, Huntleya, Stenia. 1. 1. 2(1). 2. 3(1). 3. 4(3).

4.

Lip continuous with the column, without an articulation ........................ 2 Lip articulate to the column ...................................................................... 3 Column short, cochleate; pollinia pear-shaped, ± equal ................. 12. Bollea Column long, narrow; pollinia linear-oblong, one pair longer than the other ........................................................................................... 141. Stenia Lip divided into an erect, crested hypochile and an arched epichile ................................................................................................. 66. Huntleya Lip without distinct divisions .................................................................... 4 Dorsal sepal usually < 2 cm long; lip entire, ± triangular in outline, the margins not strongly involute-undulate; column foot not forming a conspicuous mentum; rostellum simple, arched or aristate; tegula and viscidium hooked ........................................................... 24. Chaubardiella Dorsal sepal ≥ 3 cm long; lip slightly 3-lobed, ± square in outline, the margins strongly involute-undulate; column foot forming an obtuse conspicuous mentum; rostellum 3-parted, the median tooth linearlanceolate, porrect; tegula and viscidium flat .................. 29. Cochleanthes

222

O RCHIDACEAE

Subkey 11 to the Genera of Orchidaceae Characters in common: Plants without pseudobulbs; leaves conduplicate; inflorescence lateral. Genera (9): Campylocentrum (part), Cryptarrhena, Dichaea, Epidendrum (part), Maxillaria (part), Pleurothallis (part), Stellilabium (part), Vanilla (part), Vargasiella (part). 1. 1.

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

5(4).

5. 6(3). 6. 7(6).

7.

8(7).

8.

Flowers with a definite, external free spur from the base of the lip .................................................................................... 19. Campylocentrum Flowers without an external free spur, or, if spur present, then subglobular and firmly included in and attached to the pedicellate ovary ................................................................................................................ 2 Leaves solitary on tip of the stems, these along a well-developed, creeping rhizome; fruit burr-like, ≥ 1.5 cm long ... 111. Pleurothallis (P. lappiformis) Leaves several, distichously placed along the stem, rhizome usually abbreviate; fruit smooth, or when burr-like, < 1 cm long. ....................... 3 Inflorescences many-flowered, with a definite peduncle and rachis, longer than subtending leaves .......................................................................... 4 Inflorescences usually 1-flowered, sometimes 2- or 3-flowered, with reduced peduncles, much shorter than subtending leaves . .................... 6 Stems elongated, longer than leaves; leaves plicate; plants subterrestrial; flowers purple with perianth segments > 20 mm long ..... 153. Vargasiella Stems abbreviate, shorter than leaves; leaves conduplicate; plants strictly epiphytic; flowers greenish of yellowish with perianth segments < 10 mm ................................................................................................................ 5 Lip long-clawed, 3- or 4-lobed on its apical third; column glabrous; rostellum rounded, not hooked; inflorescences arching or pendent .......................................................................................... 33. Cryptarrhena Lip sessile, 3-lobed at the base; column flanked by apically furcate hairs; rostellum linear, hooked; inflorescences erect ................ 140. Stellilabium Lip solidly fused to the column at the columnar apex .......... 50. Epidendrum Lip free from the column or only basally fused ........................................ 7 Perianth segments ≥ 4 cm long; plants vines with fleshy leaves with convolute vernation; inflorescences successively few-flowered racemes ................................................................................................... 152. Vanilla Perianth segments < 3 cm long; plants erect or creeping, not vining, leaves conduplicate with duplicate vernation; inflorescences strictly 1-flowered ............................................................................................... 8 Lip clawed, anchor-like, with lobes produced in the apical third; the inflorescence arising opposite the leaf axil; column generally with a ligule beneath the stigma ................................................................... 39. Dichaea Lip simple or lobed about middle, not anchor-like; the inflorescence arising from the leaf axil; column without a ligule beneath the stigma ............................................................................................... 87. Maxillaria

O R C H I D A C E A E 223

Subkey 12 to the Genera of Orchidaceae Characters in common: Plants with homoblastic pseudobulbs (with several internodes); leaves plicate; inflorescence lateral. Genera (10): Bletia (part), Catasetum, Clowesia, Cycnoches, Cyrtopodium, Eulophia, Koellensteinia (part), Mormodes, Otostylis, Vargasiella (part). 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).

Pollinia 4 or 8 ............................................................................................. 2 Pollinia 2 ..................................................................................................... 5 Pollinia 8, with underground corms ................................................. 11. Bletia Pollinia 4, with pseudobulbs ...................................................................... 3 Plant rhizomatous, the small pseudobulbs separated by a long rhizome, the rhizome erect to prostrate ........................................... 153. Vargasiella Plant cespitose, the pseudobulbs aggregate .............................................. 4 Lip conspicuously 3-lobed, with a 2- or 3-lobed basal callus . 72. Koellensteinia Lip subentire, with a horseshoe-like, verrucose basal callus ...... 102. Otostylis Flowers always bisexual; column forming a foot; pollinia semiterete in cross section, attached to a flat, inconspicuous tegula; fresh pollinia also attached to each other by an adhesive tissue ................................ 6 Flowers functionally unisexual or protandric (function as pollen donors first and then, after the pollinarium is removed, as pollen receptors); column footless, or if forming a foot then pollinia round in cross section or dorsoventrally compressed, always attached to a tubular or semitubular, conspicuous tegula; pollinia when fresh, never attached to each other ............................................................................................... 7 Pseudobulb short; inflorescence with linear-lanceolate, inconspicuous floral bracts; lip shallowly 3-lobed, the base saccate ............. 55. Eulophia Pseudobulb elongate, fusiform; inflorescence with broadly ovate to elliptic, conspicuous, and often showy floral bracts; lip deeply 3-lobed, the base not saccate ................................................................ 37. Cyrtopodium Inflorescence basal ..................................................................................... 8 Inflorescence not basal ............................................................................... 9 Plants with fusiform pseudobulbs and dark green or grayish green leaves; flower unisexual, staminate flowers with a functional pollinarium and a broad, semicircular pseudostigma; column not produced into a foot ............................................................................................... 20. Catasetum Plants with ovoid pseudobulbs and very light green leaves; flowers protandric, with a functional pollinarium and a narrow, transverse stigma; column forming a conspicuous foot ............................ 28. Clowesia Inflorescence produced from the base of the uppermost leaves; flowers unisexual; column in both staminate and pistillate flowers terete, not obliquely twisted, produced into a conspicuous foot; staminate flowers with an elongate column and a semicircular pseudostigma, pistillate flowers with a short, stout column and a narrow, transverse stigma ................................................................................................36. Cycnoches

224

9.

O RCHIDACEAE

Inflorescence produced usually from the base of the middle leaves; flowers protandric or staminate, both with a broad, elongate stigma, the stigma nonfunctional in staminate flowers; column semiterete, obliquely twisted, neither elongate nor produced into a conspicuous foot ............................................................................................... 90. Mormodes

Subkey 13 to the Genera of Orchidaceae Characters in common: Plants with heteroblastic pseudobulbs or corms (only one internode), inconspicuous in Chaubardia and some Koellensteinia; leaves plicate; inflorescence lateral. Genera (29): Acacallis, Acineta, Aganisia, Batemannia, Bifrenaria, Bletia (part), Braemia, Chaubardia, Cheiradenia, Coryanthes, Galeottia, Gongora, Guanchezia, Houlletia, Hylaeorchis, Kegeliella, Koellensteinia (part), Lueddemannia, Lycaste, Paphinia, Peristeria, Polycycnis, Rudolfiella, Sievekingia, Stanhopea, Teuscheria, Vargasiella (part), Xylobium, Zygosepalum. 1. 1. 2(1). 2. 3(2). 3. 4(3).

4.

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

Pollinia 2 ..................................................................................................... 2 Pollinia 4 or 8 ........................................................................................... 13 Lip base with a prominent tooth-like callus, pilose, darker than rest of the lip ................................................................................... 80. Lueddemannia Lip base without a prominent tooth-like callus ........................................ 3 Petals much narrower than the sepals, attached to the base of the column ................................................................................................................ 4 Petals from a little narrower to wider than the sepals, not attached to the base of the column ................................................................................. 5 Inflorescence usually with < 5 flowers, column with 2 liquid-producing glands, epichile deeply saccate, bucket-like, when fresh and undisturbed usually filled with a viscous fluid, margins of petals undulate.. .............................................................................................. 31. Coryanthes Inflorescence with 10–30 flowers, column without liquid-producing glands, epichile not deeply saccate, when fresh never filled with fluid, margins of petals not undulate .............................................................. 60. Gongora Pseudobulbs 1-foliate ................................................................................. 6 Pseudobulbs with 2 or more leaves (if without leaves, check leaf articulations on top of pseudobulbs) .................................................................. 9 Inflorescence an elongate, erect or arched raceme, with usually > (5–) 10(–15) flowers ....................................................................................... 7 Inflorescence a short, pendent raceme, usually with < 5(6) flowers ........ 8 Plant epiphytic, usually below 500 m; leaf with a short, conduplicate pseudopetiole ........................................................................... 15. Braemia Plant terrestrial, usually above 1000 m; leaf with an elongate terete petiole ...................................................................................... 65. Houlletia Dorsal sepal and lip < 3 cm long; lip base with a fimbriate callus; pollinarium with a linear-oblong, hooked viscidium .............. 134. Sievekingia

O R C H I D A C E A E 225

8.

9(5). 9. 10(9). 10. 11(9). 11. 12(11). 12. 13(1).

13.

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

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

17.

Dorsal sepal and lip ≥ 3 cm long; lip base without a fimbriate callus; pollinarium with a horseshoe- or triangular-shaped, flat viscidium ............................................................................................. 138. Stanhopea Pseudobulbs usually with (2)3(4) leaves; flowers fleshy, globose or subglobose ............................................................................................ 10 Pseudobulbs always with 2 leaves; flowers membranaceous, neither globose nor subglobose ............................................................................. 11 Pseudobulb first (basal-most) internode conspicuous, upper internodes inconspicuous; lip epichile not articulate .................................... 2. Acineta Pseudobulb first and second internodes conspicuous, upper internodes inconspicuous; lip epichile articulate to the hypochile .......... 106. Peristeria Petals < 1 mm wide; lip ca. 1 mm wide; column thin, terete .....112. Polycycnis Petals and lip > 1 mm wide; column semiterete ..................................... 12 Rachis and pedicel hairy; dorsal sepal < 2 cm long; labelum and column < 1 cm long ............................................................................. 71. Kegeliella Rachis and pedicel glabrous; dorsal sepal > 4 cm long; lip and column > 1.5 cm long ......................................................................... 104. Paphinia Pollinarium without conspicuous tegula or viscidium, pollinia 8, laterally flattened; plants terrestrial with pseudobulbs partially or totally buried in the soil; pseudobulbs dorsoventrally flattened; flowers pink ..... 11. Bletia Pollinarium with conspicuous tegula and viscidium, pollinia 4, dorsoventrally flattened; plants epiphytic or more rarely terrestrial, when terrestrial, the pseudobulbs not buried nor dorsoventrally flattened; flowers variable in color but rarely pink .................................................... 14 Pollinia attached to a U-shaped stipe ...................................................... 15 Pollinia attached to a single, unbranched tegula ................................... 18 Pseudobulbs suborbiculate, laterally compressed; lip with a prominent transverse callus, strongly 3-lobed, the middle lobe clawed ..... 125. Rudolfiella Pseudobulbs abruptly 4-angled or pear-shaped-angled; lip without a prominent callus, usually with a long axial, low callus, entire, if 3lobed, then the middle lobe not clawed ............................................... 16 Pseudobulbs abruptly 4-angled; rhizome elongate with the pseudobulbs spaced or more rarely the pseudobulbs aggregate .............. 10. Bifrenaria Pseudobulbs pear-shaped; rhizome abbreviated ..................................... 17 Plants terrestrial in tepui bogs; pseudobulbs partially buried in the substrate; petioles about as long or longer than the foliar blade; inflorescences with long peduncles, several times longer than the petioles, 1- or 2(3)-flowered; flowers showy with perianth segments at least 5 cm long; lip basally produced into a conical, apically slightly recurved spur, which is continuous with the column foot .................................... 61. Guanchezia Plants epiphytic in rain forests; pseudobulbs shortly creeping on the bark of the host; petioles variable in length but always shorter than the foliar blades; inflorescences short, always shorter than the petiole, first 1-flowered, then with an elongating rachis, usually totally enclosed within the papery sheaths enveloping the pseudobulbs; flowers small with perianth segments 8–10 mm long; lip with a conspicuous column foot not produced into a spur ........................................................... 67. Hylaeorchis

226

O RCHIDACEAE

18(14). Plants rhizomatous, the pseudobulbs separated by a rhizome .............. 19 18. Plants cespitose, the pseudobulbs aggregate, not separated by a rhizome .............................................................................................................. 22 19(18). Pseudobulbs inconspicuous, hidden in sheaths; pollinia not attached to a well-developed tegula ........................................................ 153. Vargasiella 19. Pseudobulbs well developed, hidden in sheaths or not; pollinia attached to a well-developed tegula ....................................................................... 20 20(19). Pseudobulb suborbicular to pear-shaped, strongly compressed; anther with a long, fleshy, apically hooked process ................... 157. Zygosepalum 20. Pseudobulbs elongate, cylindric, only lightly compressed; anther without a long, fleshy, apically hooked process ................................................ 21 21(20). Lip epichile deeply concave, the margins undulate to fimbriate; column produced into a long slender foot; pollinia attached to a short, subquadrate tegula ......................................................................... 1. Acacallis 21. Lip epichile shallowly concave, the margins entire; column produced into a short, broad foot; pollinia attached to a long, oblong-linear tegula .................................................................................................... 3. Aganisia 22(18). Inflorescence elongate, bearing a single flower ...................................... 23 22. Inflorescence short or elongate, bearing few to many flowers, one at a time or simultaneously from the same rachis (the point of attachment of old flowers sometimes hidden in sheaths) ...................................... 25 23(22). Inflorescence pendent; lip with a conspicuous spur .............. 143. Teuscheria 23. Inflorescence erect; lip without a spur .................................................... 24 24(23). Pseudobulbs inconspicuous, mostly hidden under sheaths; dorsal sepal < 2 cm long; lip with a basal, crested callus ...................... 23. Chaubardia 24. Pseudobulbs conspicuous; dorsal sepal > 4 cm long; lip with 2 long, axial keels in the basal 1/2 ................................................................... 81. Lycaste 25(22). Inflorescence an arched to pendent raceme, usually much longer than the leaves .................................................................................................... 26 23. Inflorescence an erect raceme, at least 1/2 as long as the leaves to longer than the leaves ..................................................................................... 28 26(25). Inflorescence bearing successive flowers, one at a time; lip smooth, with a very low callus ..................................................................... 67. Hylaeorchis 26. Inflorescence bearing few to several flowers; lip with a transverse, conspicuous callus or axial keels .............................................................. 27 27(26). Leaves with a conspicuous petiole, terete or conduplicate; lip entire, or if 3-lobed then only in its apical 1/2 ......................................... 156. Xylobium 27. Leaves without a conspicuous petiole; lip 3-lobed in its basal 1/2 .............................................................................................. 9. Batemannia 28(25). Pseudobulbs small, inconspicuous, hidden by sheaths; lip base with a long axial callus or a 2- or 3-lobed callus .................................................... 29 28. Pseudobulbs large; lip base with a flabellate, crested callus .................. 30 29(28). Dorsal sepal < 0.5 cm; lip with a low, long, axial callus ........ 25. Cheiradenia 29. Dorsal sepal ≥ 1 cm; lip with a basal, 2-lobed or 3-lobed retrose callus ......................................................................................... 72. Koellensteinia 30(28). Petals decurrent on column foot; lateral sepals adnate to column foot near

O R C H I D A C E A E 227

30.

apex; anther apex without a long, apically hooked process .. 58. Galeottia Petals adnate to base of column; lateral sepals adnate to sides of ± developed column foot; anther apex with a long, fleshy, hooked process ......................................................................................... 157. Zygosepalum

Subkey 14 to the Genera of Orchidaceae Characters in common: Plants with pseudobulbs; leaves conduplicate; inflorescences lateral, strictly 1-flowered. Genera (2): Maxillaria (part), Trigonidium. 1.

1.

Petals and sepals not forming a funnel-like structure nor abruptly reflexed at apex, sometimes subparallel and forming a campanulate flower; petals normally unicolored and never with an eye-like spot on the apical 1/3 ...................................................................................... 87. Maxillaria Petals and sepals basally parallel and forming a funnel-like structure, then abruptly reflexed above midlength; petals with a dark, eye-like, purpleblue spot on the apical 1/3 ................................................. 147. Trigonidium

Subkey 15 to the Genera of Orchidaceae Note: The generic limits between Otoglossum, Odontoglossum, and Oncidium are not clear. This key might have some problems when dealing with these genera, and a careful comparison is required. Characters in common: Plants with pseudobulbs; leaves conduplicate; inflorescences few- to many-flowered. Genera (26): Aspasia, Brassia, Bulbophyllum, Chelyorhcis, Comparettia, Diadenium, Dipteranthus, Epidendrum (part), Eriopsis, Ionopsis (part), Leochilus, Leucohyle, Macradenia, Miltoniopsis, Notylia, Odontoglossum, Oeceoclades, Oncidium (part), Otoglossum, Psychopsis, Rodriguezia, Scelochilus, Sigmatostalix, Solenidium, Trichocentrum (part), Trichopilia. 1. 1. 2(1). 2. 3(1). 3. 4(3). 4. 5(4). 5. 6(5).

Pollinia naked or with caudicles, without a tegula ................................... 2 Pollinia with caudicles (often reduced) and a tegula ................................ 3 Pollinia naked, without caudicles; lip free from column ... 18. Bulbophyllum Pollinia with caudicles; lip united to column to its apex ......50. Epidendrum Pollinia 4; rostellum long, exceeding the column, ribbon-like ................... ........................................................................................... 41. Dipteranthus Pollinia 2; rostellum short, not ribbon-like ............................................... 4 Pseudobulbs with 2 or 3 internodes ............................................. 52. Eriopsis Pseudobulbs of one internode .................................................................... 5 Leaves dark green spotted with white ................................... 96. Oeceoclades Leaves never spotted with white ............................................................... 6 Flowers with a spur-like structure or a well-developed gibbous cavity in

228

O RCHIDACEAE

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

12. 13(12). 13. 14(13). 14. 15(13). 15. 16(15). 16. 17(15).

17.

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

19.

the rear ................................................................................................... 7 Flowers without a spur-like or gibbous structure at the rear ................ 12 Spur-like structure or gibbous cavity without horns produced by the lip and column base .................................................................................... 8 Spur-like structure or gibbous cavity with 1 or 2 horns produced by the lip and column base .................................................................................... 9 Rear of the flower with a gibbous cavity formed by the bases of the fused lateral sepals ............................................................................. 68. Ionopsis Rear of the flower with a spur-like structure formed by the base of lip; the lateral sepals free ......................................................... 144. Trichocentrum Spur-like structure formed by the full length of the lateral sepals; column or stigmatic arms present; anther dorsal ....................... 124. Rodriguezia Spur-like structure formed by only the bases of the lateral sepals; column without appendages; anther terminal ................................................ 10 Spur-like structure with 1 horn ............................................... 38. Diadenium Spur-like structure with 2 horns ............................................................. 11 Lip much larger than the other perianth parts; spur-like structure equal to or longer than the column ................................................. 30. Comparettia Lip equal to or only slightly larger than the other perianth parts; spur-like structure shorter than the column .................................... 130. Scelochilus Petals and dorsal sepal all alike and very narrow; lateral sepals broad and petal-like; leaves rough with a sand-paper quality to their surface ............................................................................................. 119. Psychopsis Petals and dorsal sepal usually dissimilar; all sepals similar; leaves smooth .............................................................................................................. 13 Stigmatic cavity a slit longitudinally oriented on the column ............... 14 Stigmatic cavity oval or round ................................................................. 15 Column with paired appendages ............................................ 83. Macradenia Column without paired appendages .............................................. 93. Notylia Column apex with a dorsal hood or fringe extending beyond the anther cap .............................................................................................................. 16 Column apex without a dorsal hood ........................................................ 17 Leaves narrow; column hood bent ventrally and covering a large portion of the anther cap ........................................................................ 76. Leucohyle Leaves not narrow; column hood projecting straight forward ................... ............................................................................................ 145. Trichopilia Column without appendages and the lip fused to the column for 1/3–1/2 its length, appearing to emerge above the point of attachment of the other perianth parts ............................................................................. 5. Aspasia Column generally with prominent appendages and the lip free or fused to the column for 1/4 or less its length, appearing from the side to emerge from the same point of attachment of the other perianth parts ........ 18 Lip base with a cavity, often formed by a pair of fleshy, raised lamellae .. 19 Lip base without a cavity ......................................................................... 20 Column winged at its base, without well-developed, stigmatic arms perpendicular to the column; petals and sepals much longer than lip .................................................................................................... 17. Brassia Column not winged at its base, with well-developed, stigmatic arms perpendicular to the column; petals and sepals shorter than lip ..... 73. Leochilus

Acacallis 229

20(18). Inflorescences with several bracts per flower-bearing node; flowers generally borne in coordinated succession; column generally elongate-arched, swan-neck-shaped .......................................................... 135. Sigmatostalix 20. Inflorescences with only 1 bract per flower-bearing node; flowers rarely borne in coordinated succession; column when elongate not arched, never swan-neck-shaped ..................................................................... 21 21(20). Foliage gray-green; lip broad and flat with 2 acute basal projections, the column long ........................................................................ 89. Miltoniopsis 21. Foliage green; lip generally not broad, if broad, then with a short column .............................................................................................................. 22 22(21). Flowers with the column with only 2 short, obtuse stigmatic arms and the lip with 2 prominent raised lamellae that extend to the midpoint of the lip ................................................................................ 137. Solenidium 22. Flowers with the column generally without stigmatic arms, if with arms, then the callus not composed of erect lamellae .................................. 23 23(22). Basal portion of the lip parallel to (and often adnate to) the basal portion of the column for > 1/2 the length of the column .............. 95. Odontoglossum 23. Basal portion of the lip spreading from the base of the column, parallel for < 1/4 the length of the column .............................................................. 24 24(23). Column with a well-developed clinandrium and wings; basal portion of the lip paralleling the column for a few mm before spreading .............. ........................................................................................... 101. Otoglossum 24. Column usually without a clinandrium; basal portion of the lip spreading from the column ................................................................................... 25 25(24). Pseudobulbs with a thick epidermis and cuticle, often transversly striated with red or maroon lines, strongly flattened and appressed to the host’s bark; sheaths enveloping the pseudobulbs lacking foliar blades; lateral lobes of the lip obsolete ....................................................... 26. Chelyorchis 25. Pseudobulbs with thinner epidermis and cuticle, smooth or rarely streaked in red or maroon; laterally compressed or subcylindrical, not appressed to the host’s bark; sheaths enveloping the pseudobulbs with foliar blades (or if these absent, then rhizome long-creeping); lateral lobes of the lip large or small, but always conspicuous ........ 97. Oncidium 1. ACACALLIS Lindl., Fol. Orchid., Acacallis 1. 1853. [Subtribe Zygopetalinae]. by Gustavo A. Romero-González Rhizomatous, creeping to climbing, epiphytes. Stem modified into narrowly ovate to cylindrical pseudobulbs of 1 internode, when young entirely concealed by distichous, scarious sheaths. Leaf 1, membranous, narrowly ovate to narrowly obovate, acute to acuminate, plicate, petiolate, articulate. Inflorescences lateral, arising from the base of the pseudobulbs, arching, racemose, few- to many-flowered, < 1/2 as long as to longer than the leaf. Flowers resupinate, showy; floral bracts concave, appressed, narrowly ovate to ovate, shorter than the pedicellate ovary. Sepals and petals membranous, spreading, elliptic, narrowly ovate to obovate, apiculate, cuneate at the base. Lip clawed, 3-lobed, basally concave, the lateral lobes erect-spreading, dentiform, midlobe concave, flabellate to semicircular, the margins dentate or fimbriate; disk with a prominent callus. Column erect, slightly arched, semiterete, winged in the apex, produced into a long, slender foot; anther terminal; pollinia 4,

230

O RCHIDACEAE

yellow, cartilaginous, dorsoventrally flattened, subequal, obovate, attached in 2 pairs to a short, subquadrate tegula. Colombia, Venezuela, Guyana, Suriname, Peru, Brazil; 3 species, 2 in the flora area. Acacallis is closely related to Aganisia but is distinguished by a long, slender column foot and a short, subquadrate tegula. Key to the Species of Acacallis 1. 1.

Inflorescence usually as long as or longer than the leaf; dorsal sepal generally > 3 cm long; lip entire or finely dentate ................................ A. cyanea Inflorescence commonly shorter than the leaf; dorsal sepal generally < 2 cm long; lip deeply fimbriate ............................................. A. fimbriata

Acacallis cyanea Lindl., Fol. Orchid., Acacallis 1. 1853. —Aganisia cyanea (Lindl.) Rchb. f., Beitr. Orchid.-K. C. Amer. 13, t. 4. 1866 [1867]. Climbing epiphyte (rarely terrestrial); pseudobulbs 1-foliate; inflorescence arching, to 10-flowered; flowers showy; sepals and petals blue-lavender; lip brown to purple; column reddish. Wet forests, 100–500 m; Amazonas (basin of Brazo Casiquiare, La Esmeralda, basin of Río Atabapo, Río Sipapo, Río Ventuari, Tamatama, Yavita). Colombia, Brazil (Acre, Amazonas, Rondônia).

Acacallis fimbriata (Rchb. f.) Schltr., Orchis 12: 16. 1918. —Aganisia fimbriata Rchb. f., Gard. Chron. n.s. 2: 452. 1874. Climbing epiphyte; pseudobulbs 1-foliate; inflorescence arching, to 9-flowered; flowers showy; sepals and petals white; lip blue, conspicuously fimbriate. Wet forests, ca. 100 m; Amazonas (Caño Ucata southeast of Síquita, near Maroa and San Carlos de Río Negro). Colombia, Guyana, Suriname, Peru, Brazil (Amazonas). ◆Fig. 202.

2. ACINETA Lindl., Edwards’s Bot. Reg. 29: misc. 67. 1843. [Subtribe Stanhopeinae]. Neippergia Morren, Ann. Soc. Roy. Agric. Gand 5: 375, t. 282. 1849. by Gustavo A. Romero-González Cespitose, robust herbs, epiphytic or rarely terrestrial. Stem modified into ovoid to subcylindric pseudobulbs of 1 internode, often compressed and/or sulcate. Leaves 1–4, most often 3, narrowly ovate, acute, plicate, articulate, subpetiolate. Inflorescences 1–several, lateral, arising from the base of the pseudobulbs, pendent, racemose, many-flowered. Flowers globose, fleshy, waxy in texture, resupinate, showy; floral bracts concave, ovate, shorter than the pedicellate ovary. Sepals fleshy, concave, spreading, widely elliptic to very broadly ovate, the lateral sepals partially connate or not; petals spatulate, generally smaller than the sepals. Lip fleshy, rigid, subsessile to conspicuously clawed, 3-lobed, the claw concave to subsacciform, the lateral lobes large, erect, hatchet-shaped, triangular, transversely elliptic or subreniform, the midlobe subrhombic, flat or apically shallowly concave; disk with fleshy calli. Column short, erect, semiterete, glabrous or slightly pubescent toward the base, apically winged to subauriculate, rarely produced into a short, broad foot; anther terminal, operculate, incumbent, 1-locular or imperfectly 2-locular; pollinarium with 2 yellow pollinia, narrowly obovate, cleft, cartilaginous, and dorsoventrally flattened, the tegula long, oblong-linear, the viscidium subtriangular or bifurcate.

Acineta 231

Fig. 202. Acacallis fimbriata

Fig. 203. Acineta alticola

Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil; 7 species, 3 in Venezuela, 1 of these in the flora area. Acineta alticola C. Schweinf., Fieldiana, Bot. 28: 192. 1951. Epiphyte or sometimes terrestrial or lithophytic plant; inflorescences pendent to suberect, to 12-flowered; flowers yellow or pale green with pale pink spots. Sandstone bluffs or open forests on tepui summits, low-

land forests, (100–)500–2500 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Cerro Sarisariñama, Macizo del Chimantá [Toronótepui], Ptari-tepui, Río Uiri-yuk in Río Cuyuní basin), Amazonas (upper Río Orinoco, Cerro Sipapo). Brazil (Roraima). ◆Fig. 203.

232

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3. AGANISIA Lindl., Edwards’s Bot. Reg. 25: misc. 45. 1839. [Subtribe Zygopetalinae]. by Gustavo A. Romero-González Epiphytes. Rhizomes elongate, creeping to climbing. Stem modified into cylindrical pseudobulbs of 1 internode, entirely concealed by distichous, scarious sheaths. Leaf 1, membranous, narrowly ovate, acute, plicate, petiolate, articulate. Inflorescences lateral, arising from the base of the pseudobulbs, arching, racemose, few- to many-flowered, < 1/2 as long as the leaf. Flowers resupinate, showy; floral bracts concave, appressed, ovate, shorter than the pedicellate ovary. Sepals and petals membranous, spreading, oblong to oblong-obovate, acute. Lip subsessile, 3-lobed, basally concave, lateral lobes erect-spreading, obtuse, midlobe shallowly concave, rhombiform, the margins entire; disk with a fleshy, cristate callus. Column erect, slightly arched, semiterete, winged in the apex, produced into a short, broad foot; anther terminal; pollinia 4, yellow, cartilaginous, dorsoventrally flattened, subequal, obovate, attached in 2 pairs to a long, oblong-linear tegula. Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil; 1 species, in Venezuela restricted to the flora area. Aganisia is closely related to Acacallis Lindl. It is distinguished by a short, broad column foot and a long, oblong-linear tegula. Aganisia pulchella Lindl., Edwards’s Bot. Reg. 25: misc. 45. 1839. Aganisia brachypoda Schltr., Beih. Bot. Centralbl. 42(2): 126. 1925. Epiphyte or terrestrial climbing herb; pseudobulbs 1-foliate; inflorescence erect, to 8-flowered; flowers showy; sepals and petals white; lip yellowish white, reddish at the base. Locally abundant in wet forests, 400– 1000 m; Bolívar (Altiplanicie de Nuria, Gran Sabana, Macizo del Chimantá [Acopántepui], Río Churún near Guarimba). Colom-

bia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil (Amazonas, Pará). ◆Fig. 204. A plant with an old inflorescence, fitting the description of this species, was collected in the state of Amazonas, Caño Ucata, a black-water tributary of the Orinoco just north of its confluence with Río Atabapo. This collection constituted the first record of this genus for Amazonas, but the plant never flowered in cultivation and no herbarium specimen was ever made.

4. ARACAMUNIA Carnevali & I. Ramírez, Ann. Missouri Bot. Gard. 76: 962. 1989. [Subtribe Spiranthinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Small, terrestrial or muscicolous, erect herbs. Roots 3–6, cylindric, fleshy, coarse, pubescent. Stems terete. Leaves 4 or 5 at flowering; sheaths imbricating and enclosing the stem, ovate to ovate-elliptic, with 1(2) erect, strap-shaped structures connected at the center of the base and protruding from it, rigid, ± expanded apically, where covered by multicellular, short-pedunculate, globose hairs; blades not articulate, elliptic or ovate-elliptic, flat or concave, glabrous, thinly subfleshy, marcescent (remaining on the plants and eventually disintegrating there), basally contracted into a channeled pseudopetiole that merges into the sheath, margins slightly undulate, apex acute or shortly acuminate. Inflorescence a terminal, 3–6flowered spike; peduncle terete with few articulations, covered with sparse, cylindric to subcapitate, unicellular hairs; peduncle bracts glabrous, lanceolate, long-

Aracamunia 233

Fig. 204. Aganisia pulchella

Fig. 205. Aracamunia liesneri

acuminate, basally enveloping and apically spreading; rachis straight or slightly zigzag. Flowers tubular, resupinate, white; floral bracts similar to peduncle bracts but somewhat smaller; pedicellate ovary subcylindrical, much shorter than the bracts. Sepals fused in the basal 3/4, forming a cylindric tube around the column, petals, and lip, their free parts concave, triangular-oblong, rounded apically, minutely glandular-papillose outside; dorsal sepal partially adnate to back of column in the basal 1/3. Petals linear-spatulate, membranous, connivent with the dorsal sepal along their whole length, 1-veined. Lip narrowly oblong in general outline, membranaceous, 3veined, base sagittate with 2 retrorse, linear-oblong, apically rounded and thickened glands, lip apex obtuse, thickened and finely papillose-ciliate, claw linear-oblong and adnate to the sepal tube. Column erect, elongate, subcylindric; pollinia 4, linearclavate, viscidium globular, small, at the apex of the rostellum, rostellum oblong, Ushaped in transverse section, apically truncate. Endemic to the southern Venezuelan Guayana Shield; 1 species.

234

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Aracamunia liesneri Carnevali & I. Ramírez, Ann. Missouri Bot. Gard. 76: 962, fig. 13. 1989. Plant 4–7 cm tall; leaves 5–9 × 3–4 mm; inflorescence 3.4–4.5 cm long; peduncle 2.1– 2.8 cm long, 3-articulate; rachis 1.2–1.7 cm long; sepals forming a cylindric, 5–5.5 mm long tube; petals ca. 5 × 1 mm; lip 5.5–6 × 1

mm. Moss-covered streambanks in tepui-summit forests, 1500–1600 m; Amazonas (Cerro Aracamuni). Endemic. ◆Fig. 205. The unusual club-like fleshy projections in the bases of the petioles are unique in the subtribe. Further research of these structures is needed since they may have a glandular function.

5. ASPASIA Lindl., Gen. Sp. Orchid. Pl. 139. 1832. —Odontoglossum sect. Aspasia (Lindl.) Rchb. f. in Walp., Ann. Bot. Syst. 6: 849. 1864. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphyte or lithophyte, erect, cespitose. Rhizomes short. Pseudobulb a single internode (heteroblastic), compressed, stalked, with several imbricate sheaths, the innermost 1–5 developing leaf blades. Leaves 1 or 2(–4), terminal, conduplicate, thinly coriaceous, oblong-elliptic to elliptic, acute to shortly acuminate, basally attenuated into a short channeled pseudopetiole. Inflorescences 1–3, lateral, racemose, 1–8-flowered, shorter than leaves and subequal to the pseudobulb. Flowers showy, opening successively or 2 or 3 simultaneously, floral bracts concave, conspicuous but shorter than pedicellate ovary. Perianth segments spreading, similar in length. Sepals and petals yellow-green, usually with purple, maroon, or brown markings, subfleshy, subequal or the petals somewhat broader and shorter, oblongelliptic or obovate; dorsal sepal and petals connivent, inserted on column above the lateral sepals. Lip white, often with markings as in other perianth segments, shortly and narrowly clawed, fused with basal 1/2 of the column, forming a nectary-like tube, reflexed, simple or inconspicuously 3-lobed, pandurate to subquadrate, suborbicular or obovate, margins somewhat incurved or flattened; disk with 2–many raised longitudinal keels. Column erect, semiterete or somewhat recurved, sometimes apically bidentate, footless; anther terminal, incumbent, operculate, 2-locular; pollinia 2, ovoid or pear-shaped, with a short tegula and a small viscidium; stigma ventral. Central America, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 5 species, 1 in Venezuela. Aspasia variegata Lindl., Edwards’s Bot. Reg. 22: t. 1907. 1836. Herb 10–40 cm tall, usually growing low on trees; dorsal sepal 16–25 mm long. Evergreen lowland forests, 100–400 m; Delta Amacuro (Río Amacuro), Bolívar (Altiplanicie de Nu-

ria, Río Caroní, Río Caura, Río Cuyuní, Río Parguaza), Amazonas (Cerro Yutajé, Río Cataniapo, Río Sipapo). Aragua, Barinas, Miranda, Sucre; Colombia, Trinidad-Tobago, Guyana, eastern Ecuador, Amazonian Brazil, Bolivia. ◆Fig. 206.

6. ASPIDOGYNE Garay, Bradea 2: 200. 1977. [Subtribe Goodyerinae]. Erythrodes Blume, Bijdr. 410, t. 72. 1825, pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial, creeping herbs, shade-loving. Rhizomes short to long, decumbent, terete, the apical part erect and foliose. Leaves spirally arranged; sheaths widely spaced or imbricating; blade mostly elliptic or ovate, acute or acuminate, basally attenuated into a channeled pseudopetiole and then into a sheathing base. Inflores-

Aspidogyne 235

Fig. 206. Aspasia variegata

Fig. 207. Aspidogyne steyermarkii

cence terminal, a short- or long-pedunculate, erect raceme, internodes with tubular sheaths. Flowers resupinate. Floral bracts small to relatively large; ovary shortly pedicellate. Sepals free, ± similar, usually elliptic, oblong-elliptic, or narrowly obovate; petals connivent with dorsal sepal, mostly linear-obovate, 1-veined. Lip basally produced into a short to long spur, differentiated into a hypochile and an epichile; hypochile usually concave to cymbiform, marginally adnate to column, epichile recurved, ovate or elliptic to transversely oblong or reniform and with widely divaricating lateral arms and/or lobes. Column erect, elongate; rostellum large in proportion to column, simple, apex entire, rarely emarginate to 2-denticulate, margins somewhat revolute; stigmas 2, approximate, horizontal; anther erect; pollinia 4, sectile, with caudicles and a large viscidium. Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil, Paraguay, northern Argentina; 26 species, 8 in Venezuela, 4 of these in the flora area. Key to the Species of Aspidogyne 1. 1. 2(1).

Inflorescence much longer than leaves; leaf blades variegated ............... 2 Inflorescence shorter than the leaves; leaf blades not variegated ........... 3 Leaves longitudinally variegated, apical lobe of lip transversely reniform, much wider than lip blade ........................................................... A. pumila

236

2. 3(1). 3.

O RCHIDACEAE

Leaves reticulately variegated; apical lobe of lip flabellate or transversely rhombic, narrower than lip blade ....................................... A. steyermarkii Leaves 7–11 × 2–3 cm; sepals ca. 7 mm long, spur to 14 mm long ... A. confusa Leaves 14–27 × 5–7 cm; sepals 15–17 mm long, spur 23–26 mm long ..................................................................................................... A. robusta

Aspidogyne confusa (C. Schweinf.) Garay, Bradea 2: 201. 1977. —Erythrodes confusa C. Schweinf., Fieldiana, Bot. 28: 174. 1951. Herb to 40 cm tall in flower; lower surface of leaves silvery or plain green; sepals and petals green or purplish; lip white. Rain forests or cloud forests, 300–1100 m; Bolívar (Río Caroní basin), Amazonas (Cerro Duida). French Guiana. Aspidogyne pumila (Cogn.) Garay, Bradea 2: 204. 1977. —Physurus pumilus Cogn. in Mart., Fl. Bras. 3(6): 545. 1906. —Erythrodes pumilus (Cogn.) Pabst, Orquídea (Niteroi) 18: 215. 1957. Herb to 30 cm tall in flower; foliage silvery and purplish; flowers green and white. Rain forests, 200–300 m; Bolívar (Río Caura basin). Amazonian Brazil.

Aspidogyne robusta (C. Schweinf.) Garay, Bradea 2: 203. 1977. —Erythrodes robusta C. Schweinf., Bot. Mus. Leafl. 20: 8. 1962. Herb 40–100 cm tall in flower, rather large for the genus; flowers white. Cloud forests, tepui slopes, 900–1400 m; southeastern Bolívar. Endemic. Aspidogyne steyermarkii Carnevali & Foldats, Ann. Missouri. Bot. Gard. 76: 596. 1989. —Erythrodes steyermarkii (Carnevali & Foldats) Carnevali & Dodson, Lindleyana 8: 97. 1993. Herb to 35 cm tall in flower; stems apically 2–6-foliate; leaves deep olive brown with red streaks on both sides of midrib and similarly colored spots near margin above; peduncle densely pubescent; perianth segments white. Rain forests, ca. 500 m; Bolívar (Sierra de Lema). Endemic. ◆Fig. 207.

7. BARBOSELLA Schltr., Repert. Spec. Nov. Regni Veg. 15: 259. 1918. [Subtribe Pleurothallidinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Mostly epiphytes, very rarely lithophytes, usually creeping, rarely cespitose, minute to small. Rhizomes short to long but always well developed, enclosed by sheaths. Ramicauls short, without an annulus (annular thickening encircling ramicaul at the level of emergence of the inflorescence), erect to prostrate, 1-foliate, enclosed by sheaths. Leaves fleshy, conduplicate, somewhat concave to flat, usually oblanceolate but sometimes broadly ovate to suborbicular. Inflorescences 1-flowered, originating near the apex of ramicaul, longer to shorter than leaves, when long, with a sheathed articulation below the middle, when very short, 2- or 3sheathed. Flowers resupinate, yellow to purple, usually with widely spreading segments, rarely subparallel to column and divergent only at apex; dorsal sepal narrow, tapering to finger-like apex; lateral sepals often totally connate into a ± concave to flat synsepal which is subequal to the dorsal sepal but sometimes longer and wider; petals narrow, tapering to a finger-like apex, usually similar to the dorsal sepal but often narrower. Lip fleshier than other perianth segments, simple to 3-lobed, flat to concave, with a concavity at its base that articulates with the bulbous apex of the stout column foot like a ball-and-socket joint. Column well developed; anther ventral, operculate, incumbent; pollinia 4, waxy, clavate. Central America, Colombia, Venezuela, Ecuador, Peru, Brazil, northern Argentina; 20 species; 3 in Venezuela, 1 of these in the flora area.

Batemannia 237

Barbosella orbicularis Luer, Selbyana 3: 10. 1976. Minute epiphyte, repent and mat-forming; leaves 3–7 mm long, suborbicular, fleshy, borne flat against the bark of the host tree; inflorescence with a relatively large flower on short and erect peduncles; sepals 6–7 mm long, red or dark wine with cream margins; petals creamy, papillose; lip brilliant dark redmaroon; column yellow with a red foot. Lower montane to montane cloud forests, 700–1300 m; Bolívar (Cerro Jaua, Cerro Sarisariñama,

Río Icabarú basin, Serranía Marutaní). Panama, Ecuador, Costa Rica. ◆Fig. 208.

Fig. 208. Barbosella orbicularis

8. BASKERVILLA Lindl., Gen. Sp. Orchid. Pl. 505. 1840. [Subtribe Cranichidinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial, humicolous to subepiphytic herbs, shade-loving. Roots fleshy, fasciculate, frequently horizontal, variously pubescent. Stems almost absent to somewhat elongate. Leaves basal, rosulate, distinctly petiolate; blades elliptic, narrowovate, narrow-obovate, or narrow-elliptic, acute to short-acuminate. Inflorescence terminal, spicate, erect; peduncle remotely sheathed, sheaths sometimes with foliar blades; rachis lax to dense and many-flowered; floral bracts narrow-ovate, acute to acuminate. Flowers nonresupinate, mostly white, green or greenish white, often with widely spreading segments or ± campanulate; ovary cylindric, sessile, usually glabrous; dorsal sepal free, narrower than lateral sepals, elliptic, narrow-ovate, or narrow-elliptic, obtuse or acuminate; lateral sepals free or rarely slightly connate at base, oblique, ovate to oblong-elliptic, acute; petals clawed, dorsally united to the column, then abruptly bent back, usually obliquely lobed to auriculate. Lip fused to column either near or above the column base, base saccate, forming a blunt spur or chin, lateral lobes replicate, between column and rest of lip, midlobe scoop-like, fleshier than other perianth segments. Column club-shaped, pointed, footless; pollinia 4, unequal, brittle, club-shaped, with caudicles, viscidium with a small hamulus (pollinium stalk developed from rostellum). Central America, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia; 8 species, 3 in Venezuela, 1 of these in the flora area. Baskervilla colombiana Garay, Svensk. Bot. Tidskr. 47: 196, fig. 1. 1953. Terrestrial herb to 50 cm tall; leaves silvery on lower surface; inflorescence densely flowered; flowers greenish white. Montane cloud forests on upper tepui slopes and summits, 1900–2000 m; Bolívar (Macizo del

Chimantá). Costa Rica, Colombia (Cauca). ◆Fig. 209. The type of this species comes from the Western Cordillera of Colombian Andes, and the Venezuelan material may represent an undescribed species.

9. BATEMANNIA Lindl., Edwards’s Bot. Reg. 20: t. 1714. 1834. [Subtribe Zygopetalinae]. by Gustavo A. Romero-González Cespitose epiphytes. Stem modified into oblong-ovoid, compressed, furrowed pseudobulbs of 1 internode, basally concealed by distichous, scarious sheaths. Leaves 1–3, coriaceous, broadly ovate, acute to acuminate, plicate, articulate. Inflo-

238

O RCHIDACEAE

Fig. 209. Baskervilla colombiana

Fig. 210. Batemannia lepida

rescences lateral, arising from the base of the pseudobulbs, arching to pendent, racemose, few-flowered, < 1/2 as long as the leaves. Flowers resupinate, showy; floral bracts concave, appressed, ovate, < 1/2 as long as the pedicellate ovary. Sepals and petals membranous; dorsal sepal erect, oblong-elliptic, concave; lateral sepals spreading, narrowly oblong-ovate to oblong-elliptic, oblique, adnate to the apex of the column foot; petals similar to dorsal sepal, adnate to basal portion of the column foot. Lip sessile to clawed, 3-lobed, the lateral lobes erect, midlobe recurved; disk with a laminar, transverse, antrorse callus. Column elongate, slightly arched, semiterete, produced into a conspicuous foot; anther terminal; pollinia 4, yellow, cartilaginous, dorsoventrally flattened, subequal, clavate, attached in 2 pairs to a trullate tegula.

Bifrenaria 239

Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 5 species, 2 in Venezuela, both restricted to the flora area. Batemannia is closely related to Galeottia A. Rich., but is easily distinguished by a nonauriculate column and sepals not basally saccate. Key to the Species of Batemannia 1. 1.

Lip sessile (without a basal claw), the midlobe not separated from the lateral lobes by a conspicuous isthmus ........................................ B. colleyi Lip with a conspicuous basal claw, the midlobe contracted at the base into a conspicuous isthmus that separates it from the lateral lobes .. B. lepida

Batemannia colleyi Lindl., Edwards’s Bot. Reg. 20: t. 1714. 1834. Pseudobulbs 1- or 2-foliate; inflorescence arching to pendent, to 10-flowered; sepals and petals light green, with some brown or maroon at the apex; lip whitish cream inside, purple in the base, yellowish cream outside. Wet forests, 50–400 m; Bolívar (Gran Sabana, Río Parguaza basin), Amazonas (basin of Río Cataniapo, basin of Río Sipapo). Colombia, Trinidad-Tobago, Guyana, Ecuador, Peru, Brazil (Acre, Amazonas), Bolivia.

Batemannia lepida Rchb. f., Gard. Chron. n.s. 9: 588. 1878. Pseudobulbs 1-foliate; inflorescence arching, to 3-flowered; sepals and petals light greenish brown; lip midlobe white, the lateral lobes, disk, and callus light pink. Wet forests, 100–200 m; Amazonas (Brazo Casiquiare, Río Siapa, San Carlos de Río Negro). Brazil (Amazonas). ◆Fig. 210.

10. BIFRENARIA Lindl., Gen. Sp. Orchid. Pl. 152. 1833. [Subtribe Lycastinae]. Stenocoryne Lindl., Edwards’s Bot. Reg. 29: misc. 53. 1843. Adipe Raf., Fl. Tellur. 2: 101. 1836 [1837]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, lithophytes, or subterrestrial herbs, variable in size, shade- or sunloving, cespitose to creeping, usually erect. Rhizomes usually well developed, thick or thin. Pseudobulbs a single internode (heteroblastic), tetragonous or very rarely pear-shaped, 1-foliate or rarely 2-foliate. Leaves erect to spreading, plicate, coriaceous, rigid-coriaceous to subfleshy, mostly elliptic or ovate-elliptic, acute or obtuse, petiolate or sessile; petiole terete or channeled. Inflorescences axillary or basal, from lower nodes or very base of pseudobulb, relatively short, 1-flowered or few- to manyflowered dense raceme, erect to arching; floral bracts relatively small. Flowers resupinate, small to rather large and showy, variously colored, often long-lived; sepals subequal or laterals longer; dorsal sepal erect, free, concave; lateral sepals decurrent on column foot, oblique, free or basally connate and forming with column foot a variously developed spur or a conspicuous mentum; petals subequal to dorsal sepals or smaller and narrower. Lip articulate to column foot, clawed or subsessile, often 3lobed, lateral lobes erect and often enfolding the column; central lobe flat or apically concave or recurved, rounded, truncate, or emarginate at apex; disk callose, glabrous or variously pubescent. Column erect, relatively thick, semiterete, with a variously developed foot; anther terminal, operculate, incumbent; pollinia 4, cartilagi-

240

O RCHIDACEAE

Fig. 211. Bifrenaria longicornis

Fig. 212. Bifrenaria venezuelana

nous, with tegula U-shaped, split into 2 lobes; viscidium small, cordate to semilunate; stigma ventral. Capsule oblong or ellipsoid. Panama, Colombia, Venezuela, Trinidad-Tobago, Ecuador, Peru, southern Brazil; ca. 35 species, 3 in Venezuela, all in the flora area. Key to the Species of Bifrenaria 1. 1. 2(1). 2.

Plants with a short rhizome, the rhizome sometimes shortly creeping but pseudobulbs set on the rhizome for < 1/4 their length ........ B. steyermarkii Plants with a long-creeping rhizome, the pseudobulbs set on the rhizome for 1/4 or more their length ..................................................................... 2 Inflorescence a many-flowered raceme; lip with a well-developed claw ................................................................................................B. longicornis Inflorescences 1- or 2-flowered; lip without a claw ................. B. venezuelana

Bifrenaria longicornis Lindl., Edwards’s Bot. Reg. 24: misc. 93. 1838. —Stenocoryne longicornis (Lindl.) Lindl., Edwards’s Bot. Reg. 29: misc. 53. 1843. —Adipe longicornis (Lindl.) M. Wolfe, Orchideen 41(2): 36. 1990. Herb 20–25 cm tall; rhizome sometimes branched; inflorescence 6–10-flowered; flow-

ers cream or yellowish with small purple or reddish spots. Rain forests, 50–600 m; Bolívar (upper Río Caroní, Río Parguaza), Amazonas (Río Atabapo, Río Casiquiare, Río Negro, Río Sipapo). Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 211.

Bollea 241

Bifrenaria steyermarkii (Foldats) Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 56. 1976. —Xylobium steyermarkii Foldats, Noved. Ci., Ser. Bot. 35: 1, fig. 1. 1970. Epiphyte or terrestrial herb to 6 m tall; pseudobulbs sometimes 2-foliate; leaves petiolate; flowers with very narrow perianth segments, brownish. Cloud forests, 900– 1200 m; Bolívar (La Escalera to Cerro Venamo region). Endemic.

Bifrenaria venezuelana C. Schweinf., Amer. Orchid Soc. Bull. 34: 38. 1965. Epiphyte or lithophyte to 20 cm tall; leaves short-petiolate; inflorescences shorter than pseudobulbs, slightly ascending to pendulous; flowers brownish purple with pale green margins. Rain and cloud forests, 100– 1400 m; Bolívar (Cerro Guaiquinima, Cerro Ichún, Macizo del Chimantá), Amazonas (Cerro Aracamuni, Río Negro, Sierra de la Neblina, road between Yavita and Maroa). Táchira. ◆Fig. 212.

11. BLETIA Ruiz & Pav., Fl. Peruv. Prodr. 119, t. 26. 1794. [Subtribe Bletiinae]. Gyas Salisb., Trans. Hort. Soc. London 1: 299. 1812. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial to sublithophytic herbs, cespitose to rhizomatous, sun-loving, small to medium-sized. Stem a Gladiolus-like corm, underground or on the ground. Leaves plicate, articulate, deciduous, borne on apical internodes of the corm, long-petiolate; petioles connivent into an erect, ± well-developed pseudostem; blades usually elliptic or ovate-elliptic, acute or long-acuminate. Inflorescences originating from corm internodes, racemose or secondarily paniculate, ± equal to or longer than leaves, erect; peduncle elongate, terete. Flowers resupinate, showy, usually pink to purple, sometimes white, campanulate, fugacious, often cleistogamous; floral bracts small to large and relatively conspicuous. Sepals subequal, free or the lateral ones basally connate and forming a small mentum; petals similar or broader than sepals. Lip free or united with column base or foot, unlobed or more often 3-lobed; lateral lobes usually large and enfolding the column; central lobe erect or recurved, usually truncate, emarginate or 2-lobed at apex; disk lamellate or ± pubescent. Column elongate, semiterete, straight, curved, or S-shaped, foot present or absent, winged, often 2-auriculate; anther terminal, operculate, incumbent; clinandrium 3-lobed; pollinia 8, waxy, with caudicles; rostellum transverse; stigma ventral. Capsule cylindric to ellipsoid. U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, northern Argentina; ca. 30 species, 4–6 in Venezuela, 1 of these in the flora area. Bletia stenophylla Schltr., Repert. Spec. Nov. Regni Veg. Beih. 6: 42. 1919. Cespitose herb to 60 cm tall; inflorescences erect, successively few- to many-flowered; flowers campanulate, sometimes cleistogamous; perianth segments pink or purple; dorsal sepal 2–3 cm long; lip with deep purple

keels. Locally common, sandstone outcrops, rarely granitic outcrops, 800–1900 m; Bolívar (Auyán-tepui, Cerro Murú near Aravinquén, Gran Sabana, Macizo del Chimantá), Amazonas (Sierra Parima). Aragua, Distrito Federal, Lara, Miranda, Trujillo, Yaracuy; Colombia, Ecuador. ◆Fig. 213.

12. BOLLEA Rchb. f., Bot. Zeitung (Berlin) 10: 667. 1852. [Subtribe Zygopetalinae]. by Gustavo A. Romero-González Cespitose epiphytes. Stem abbreviate, not modified into pseudobulbs. Leaves numerous, distichous, in a flabelliform cluster, coriaceous, narrowly ovate to oblongobovate, acute to acuminate, plicate, articulate. Inflorescences lateral, arising from the axil of the leaf sheaths, erect to arching, 1-flowered, shorter than the leaves.

242

O RCHIDACEAE

Fig. 213. Bletia stenophylla

Fig. 214. Bollea hemixantha

Flowers resupinate, showy; floral bract concave, appressed, ovate-triangular to tubular-funnel-shaped, shorter than the pedicellate ovary. Sepals and petals fleshy, erect-spreading to spreading; sepals ovate to elliptic-ovate; petals ovate to narrowovate. Lip clawed, adnate to the apex of the column foot, ovate to obovate, basally cordate, apically rounded in outline when spread, the margins strongly recurved;

Brachionidium 243

disk with a conspicuous, thickened, keeled, transverse callus. Column short, fleshy, semiterete, basally deeply concave, produced into a short foot; anther terminal; pollinia 4, yellow, cartilaginous, dorsoventrally flattened, subequal, obovate, attached in 2 pairs to a short, oblong tegula. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 7 species, 1 in Venezuela. Bollea hemixantha Rchb. f., Gard. Chron. ser. 3, 4: 206. 1888. Bollea cardonae Schnee, Revista Fac. Agron. (Maracay) 1: 203. 1953. Plant fan-shaped, lacking pseudobulbs; sepals and petals yellowish white; lip light yellow with bright yellow callus, column ex-

ceedingly enlarged, hood-like, white with some purple markings, projecting over the entire keeled callus on the lip base. Wet forests, 100–1300 m; Bolívar (Río Icabaru, Río Kukenán, Santa María de Erebato), Amazonas (Río Siapa, Sierra de la Neblina, Simarawochi). Colombia, Guyana. ◆Fig. 214.

13. BRACHIONIDIUM Lindl., Fol. Orchid., Brachionidium 1. 1859. [Subtribe Pleurothallidinae]. Yolanda Hoehne, Arq. Mus. Nac. Rio de Janeiro 22: 72. 1919. —Brachionidium sect. Yolanda (Hoehne) Pabst, Bradea 1: 269. 1972. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes or terrestrial herbs, cespitose to long-creeping, sometimes matforming, usually shade-loving, erect to pendulous. Rhizomes erect to creeping, branching, elongate between ramicauls, enclosed by a few tubular, glabrous to scurfy sheaths, usually mucronate; roots few to many near the base, or singly from nodes on the rhizome, slender to coarse or fleshy. Ramicauls ascending to erect, shorter than the leaf, 1-foliate, usually enclosed by 2 sheaths similar to those of the rhizome, the inflorescence emerging laterally without an annulus (annular thickening encircling ramicaul at the level of emergence of the inflorescence) from near the apex (the leaf-stem abscisssion layer). Leaf erect in relation to the ramicaul, thin- to thick-coriaceous, smooth to rugose-verrucose, green to dark green, sometimes suffused with purple, elliptic to narrow-elliptic, the apex acute to obtuse, shallowly notched with a mucro in the sinus, the base cuneate or narrow-cuneate in a short petiole, sometimes with 1–3 veins at each side of the central vein, longer to shorter than the subtending leaf. Inflorescence 1-flowered. Flowers widely opening, usually not resupinate, borne on a peduncle longer or shorter than the leaf, with a bract near the middle and another near the base, mostly lasting only 1 day; floral bract cucullate, inflated, acuminate, enclosing the abbreviate pedicel and often much of the ovary; pedicel stout, with a filament much longer than the pedicel itself; ovary smooth, 3-valved. Perianth segments membranous, spreading widely, green or yellowish tinged with red or purple, sometimes wholly red-violet or purple, frequently with ciliolate margins; dorsal sepal essentially free, usually the lowermost acute to obtuse, the apex often acuminate in a slender tail; lateral sepals connate into a flat or concave uppermost synsepal, essentially free, acute to obtuse, the apex often acuminate into a slender, sometimes bifid tail; petals similar to sepals, giving appearance of a 4-tepaled flower, often minutely ciliate, the apex usually acuminate in a slender tail. Lip hinged to the apex to the column foot, smaller than other perianth segments, very fleshy, usually transverse, apex acute, obtuse to rounded, the margin smooth to fimbriate, sometimes bimarginate, with or without an apiculum, the disk ± carinate or excavate in the anterior part, with a flat, ovoid, ± pubescent callus in

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the center above the base that fits under the column in the natural position. Column stout, short, the anterior margin of the column bidentate, or smooth; anther apical, the base of the column forming a thick foot with the apex of the ovary; pollinia 6 or 8, of unequal sizes, in 2 packets, clavate, often free or attached to a small viscidium; rostellum transverse; stigma apical, transverse, 2-lobed, sometimes apparently 2 but continuous beneath the rostellum flap with the receptive surfaces protruding to either side of the flap, sometimes on short, acute lobes or “arms.” Capsule oblong to oblong-ovoid, obtuse, trigonous. Guatemala, Costa Rica, Panama, West Indies, Colombia, Venezuela, Guyana, Ecuador, Peru, southern Brazil, Bolivia; 64 species, 8 in Venezuela, 5 of these in the flora area. The recently described Brachionidium renzii Luer, from the Guyanan side of Roraima-tepui almost certainly occurs in the flora area. It would key to B. julianii but has flowers at least twice as large and the lip is much broader. Key to the Species of Brachionidium 1. 1. 2(1). 2. 3(2). 3. 4(2). 4.

Rhizome short; ramicauls closely spaced ....................................... B. julianii Rhizome relatively elongate to scandent; ramicauls widely spaced ........ 2 Synsepal elongated into a filiform tail ...................................................... 3 Synsepal shortly acuminate ...................................................................... 4 Lip geniculate; growing above 2000 m ............................... B. longicaudatum Lip flat or ± convex; longer than subtending leaf; growing at 800–1800 m ..................................................................................................... B. parvum Flowers greenish or pale brownish pink; petals ciliate; lip sagittate-triangular, ciliate .................................................................... B. brevicaudatum Flowers red-violet; petals eciliate; lip obovate, basally attenuate when flattened, eciliate .......................................................................... B. neblinense

Brachionidium brevicaudatum Rolfe, Trans. Linn. Soc. London, Bot. 6: 59. 1901. Erect herb to 10 cm tall; leaf 15–30 mm long; sepals 6-10 mm long; petals 6–8 mm long, lip 4 mm long; perianth brownish pink; lip centrally light maroon, the remainder pale brownish pink. Cloud forests, 1000– 2300 m; Bolívar (Cerro Venamo, Roraimatepui, Sororopán-tepui). Guyana. ◆Fig. 215. Brachionidium julianii Carnevali & I. Ramírez, Ernstia 39: 6. 1986, “juliani.” Herb to 2 cm tall; leaf 10–12 mm long; sepals and petals 16–18 mm long; lip 2 mm long; flowers pale green throughout. Cloud forests on tepui summits, 2500–2600 m; Amazonas (Cerro Marahuaka). Endemic. ◆Fig. 217. Brachionidium longicaudatum Ames & C. Schweinf., Bull. Torrey Bot. Club 58: 348. 1931.

Fig. 215. Brachionidium brevicaudatum

Brachionidium 245

Fig. 216. Brachionidium parvum

Fig. 217. Brachionidium julianii

Lip

Lip

Fig. 218. Brachionidium neblinense

Herb to 10 cm tall; leaf 10–11 mm long; sepals and petals 11–12 mm long; lip 1.5 mm long; perianth cream or hyaline. Cloud forests on tepui summits, 1900–2700 m; Bolívar (Kukenán-tepui), Amazonas (Cerro Duida, Cerro Marahuaka). Endemic. ◆Fig. 219. Brachionidium neblinense Carnevali & I. Ramírez, Ernstia 39: 9. 1986. Herb to 5 cm tall; leaf 15–20 mm long; sepals and petals 11–12 mm long; lip 2 mm long; perianth red-violet throughout. Among bryophytes and mosses in cloud forests, ca.

Fig. 219. Brachionidium longicaudatum

2000 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 218. Brachionidium parvum Cogn., Repert. Spec. Nov. Regni. Veg. 6: 307. 1909. Herb 4–8 cm tall; sepals and petals 5–8.5 mm long; lip 1–1.5 mm long; flowers apparently always cleistogamous (Venezuelan populations); perianth light greenish to purple. Rain and cloud forests, 500–1800 m; Bolívar (near El Paují), Amazonas (Cerro Yutajé, Sierra Tapirapecó). Haiti, Puerto Rico, Guadeloupe, Grenada, Dominica, Ecuador. ◆Fig. 216.

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14. BRACHYSTELE Schltr., Beih. Bot. Centralbl. 37(2): 370. 1920. [Subtribe Spiranthinae]. Spiranthes Rich., De Orchid. Eur. 20, 28, 36. Aug.–Sep. 1817, nom. cons., pro parte, not as to type. by Germán Carnevali, Ivón M. Ramírez-Morillo, and Carlos A. Vargas Terrestrial to muscicolous herbs, usually sun-loving, often in burned-over areas, minute to small. Roots fasciculate, fleshy, fusiform, stipitate. Leaves basal, convolute, subfleshy, petiolate, few, commonly absent at flowering (hysteranthous). Inflorescence terminal; peduncle covered by several large loose bracts; rachis pilose; spike dense, cylindric, often many-flowered. Flowers resupinate, subglobose, pubescent, minute to small; perianth green and with a white lip; floral bracts often narrowly ovate, acuminate; ovary generally pubescent, ± cylindric-arched, ± twisted. Sepals free to base, subparallel; lateral sepals obliquely decurrent over the short columnar foot, forming a ± well-developed obtuse mentum; petals oblanceolate to spatulate, fused (at least basally) to dorsal sepal (forming a galea), apically oblique. Lip sessile, conduplicate, arched with a recurved apex, basally often with 2 conspicuous basal glands; blade suborbicular or subpandurate, joined in middle to sides of column. Column very short, widened toward apex, basally produced into a short, incurved foot decurrent to the ovary; anther dorsal, short, concave, rotund; pollinia 4, short, clavate, with a small, roundish viscidium that fits tightly into a pit in the rostellum; rostellum soft, discernible only as a thin, emarginate or deeply incised truncate border above the edges of the stigmas; stigma bilobed, connivent. Southern Mexico, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Peru, southern Brazil, Chile, Paraguay, northern Argentina, Uruguay; 15 species, 1 in Venezuela. Brachystele guayanensis (Lindl.) Schltr., Beih. Bot. Centralb. 37(2): 273. 1920. —Goodyera guayanensis Lindl., Gen. Sp. Orchid. Pl. 494. 1840. Herb 7–22 cm tall; sepals 2–4 mm long; petals 2–3.5 mm long; lip 2–3.5 mm long. Open savannas, especially with Trachypogon, 50–1200 m; Bolívar (Río Caroní basin, near Río Parguaza), Amazonas (near Samariapo). Apure, Lara, Mérida; southern Mexico, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, southern Brazil. ◆Fig. 220.

Fig. 220. Brachystele guayanensis

Braemia 247

15. BRAEMIA Jenny, Orchidee (Hamburg) 36: 36. 1985. [Subtribe Stanhopeinae]. by Gustavo A. Romero-González Cespitose, robust epiphytes, often in ant gardens. Stem modified into ovoid, shallowly sulcate pseudobulbs of 1 internode, when young entirely concealed by distichous, scarious sheaths, eventually naked. Leaf 1, large (to 75 cm long in the flora area), narrow to broadly ovate, acute, plicate, articulate, short-petiolate. Inflorescences 1–several, lateral, arising from the base of the pseudobulbs, robust, erect, racemose, many-flowered. Flowers resupinate, showy; floral bracts concave, narrowly ovate, acuminate, shorter than the pedicellate ovary. Sepals and petals membranous, narrowly ovate; lateral sepals slightly oblique; petals twisted at the base, shorter and narrower than the sepals. Lip conspicuously clawed, 3-lobed, the claw linear, with a small, porrect, linear-conical, pubescent callus at the base, the lateral lobes short, shallowly concave, ovate, porrect, the midlobe subsessile, flat, rhombicovate, with a low, central-axial keel; disk with a fleshy keel terminating in an upcurved horn-like callus between the lateral lobes. Column elongate, semiterete, slightly clavate, glabrous, apically winged, footless; anther terminal, operculate, incumbent, 1-locular, apiculate; pollinarium with 2 yellow pollinia, narrowly obovate, cleft, cartilaginous, and dorsoventrally flattened, the tegula long, oblong-linear, the viscidium subtriangular. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; 1 species. Braemia vittata (Lindl.) Jenny, Orchidee (Hamburg) 36: 38. 1985. —Houlletia vittata Lindl., Edwards’s Bot. Reg. 27: misc. 47, t. 69. 1841. —Polycycnis vittata (Lindl.) Rchb. f. in Walp., Ann. Bot. Syst. 6: 618. 1863. Epiphyte, with fine root projections; inflorescence erect; flowers showy, purple with yellow markings. Wet forests, 50–500 m; Bolívar (Kilómetro 88, Macizo del Chimantá [Chimantá-tepui], Río Chicanán, Río Paragua), Amazonas (Río Atabapo, Río Cataniapo, Río Cunucunuma, Río Guainía, Río Sipapo, Río Yatúa). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Amazonas, Pará). ◆Fig. 221.

Fig. 221. Braemia vittata

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16. BRASSAVOLA R. Br. in W.T. Aiton, Hortus Kew ed. 2, 5: 216. 1813, nom. cons. [Subtribe Laeliinae]. Lysimnia Raf., Fl. Tellur. 4: 43. 1836 [1838]. Tulexis Raf., Fl. Tellur. 4: 42. 1836 [1838]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes or lithophytes, erect to pendulous. Stems short, thin, only weakly thickened, apex 1- or 2-foliate. Leaves ± terete, very fleshy to near cylindrical. Racemes terminal or emerging directly from rhizome, generally shorter than leaves, 1–many-flowered. Flowers generally large and showy; pedicellate ovary sometimes with a long neck. Sepals and petals similar, spreading, linear to linear-lanceolate, sometimes setaceous-acuminate, free. Lip sessile at the base of the column, then ± abruptly expanded into an acute or acuminate, often large, ± flat blade, the margins entire or fimbriate. Column erect, generally shorter than lip claw, usually with 2 wings, footless; anther operculate, incumbent, 2-locular, each locule with longitudinal, inconspicuous divisions; clinandrium with 3 prominent, ± equal lobes; pollinia 8, laterally compressed; rostellum transverse; stigma ventral. Capsules ellipsoid. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, northern Argentina; ca. 15 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Brassavola 1.

1.

Inflorescences 1–few-flowered; peduncle 1–3 cm long; pedicellate ovary 15–25 cm long; dorsal sepal 5–12.5 cm long; lip cordate-ovate to suborbicular-ovate, long caudate-acuminate, lateral margins conspicuously fimbriate ................................................................................... B. cucullata Inflorescences many-flowered; peduncle 4–15 cm long; pedicellate ovary 4.5–10 cm long; dorsal sepal 2.5–5.5 cm long; lip ovate, acute, shortly acuminate, margins slightly fimbriate .................................... B. martiana

Brassavola cucullata (L.) R. Br. in W.T. Aiton, Hortus Kew ed. 2, 5: 216. 1813. —Epidendrum cucullatum L., Sp. Pl. ed. 2, 2: 1350. 1763. Herb 25–35 cm tall; perianth widely spreading to reflexed, entirely white, greenish white, or yellowish; lip sometimes with purple spots. Dry to deciduous forests, 50– 400 m; Bolívar (lower Río Caroní, Río Caura, Río Cuyuní, Serranía de Imataca), Amazonas (upper Río Negro). Apure, Aragua, Cojedes, Distrito Federal, Falcón, Guárico, Miranda, Monagas, Sucre, Táchira, Yaracuy; Mexico, Central America, Colombia. ◆Fig. 222. Brassavola martiana Lindl., Edwards’s Bot. Reg. 22: sub t. 1914. 1836.

Brassavola angustata Lindl., Edwards’s Bot. Reg. 24: misc. 41. 1838. Herb 20–25 cm tall, pendulous, sun-loving; flowers with perianth segments spreading; sepals and petals yellowish or cream; lip white. Rain forests, often close to rivers, 50–400 m; Bolívar (basins of Río Caroní, Río Caura, Río Cuyuní, and Río Parguaza), Amazonas (widespread). Apure; Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 223. There is considerable variation in lip width within what we currently treat as Brassavola martiana. Brassavola angustata could be a different species characterized by smaller flowers with narrower lip. Brassavola paraensis Huber could be a synonym of B. martiana judging by its distribution.

Brassia 249

Fig. 222. Brassavola cucullata

Fig. 223. Brassavola martiana

17. BRASSIA R. Br. in W.T. Aiton, Hortus Kew ed. 2, 5: 215. 1813. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, lithophytes, terrestrial or subterrestrial herbs, cespitose to longcreeping, erect to subpendulous, shade- or sun-loving. Rhizomes almost absent to rather stout and long, covered with imbricating, scarious sheaths; pseudobulbs 1–3 (–5)-leaved, usually oblong, ovoid, or ellipsoid, sometimes subcylindric, often sharply flattened, rarely elongate and cauliform, covered with several imbricating sheaths, the innermost 1–3 may have foliar blades. Leaves conduplicate, articulate with their sheaths, oblong, oblong-elliptic, or oblong-obovate, very rarely linear, acute to obtuse, ± cuneate below, thinly coriaceous to subfleshy, sessile to subpetiolate. Inflorescences racemose, rarely paniculate, originating from the base of the pseudobulbs, 1– 3 simultaneously, lax to dense, few- to many-flowered; peduncle stout, remotely fewsheathed. Flowers resupinate, usually very showy; sepals long-lasting, with widely spreading perianth segments, mostly in shades of green, yellow, or brown, spotted or blotched with brown, purple, or maroon, often fragrant; floral bracts very small to large; pedicellate ovary straight to slightly curved, usually smooth. Sepals free, straight or variously curved, subequal or laterals longer than dorsal, narrow-elliptic

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or ovate-elliptic, linear, acute, acuminate, or with an elongate, caudate apex, flat or concave; petals similar to sepals but often shorter. Lip sessile, spreading from base of column, unlobed, ± 3-lobed, or pandurate, mostly shorter and wider than the other perianth segments, keeled or with complex callosities, flat, concave, or convex, surface smooth or verruculose, apex frequently caudate or acuminate. Column short, stout, footless, wingless; anther terminal, incumbent, operculate, 1-locular or imperfectly 2-locular; pollinia 2, cartilaginous, obovoid, attached to a short, broad tegula and a small viscidium; stigma ventral, transverse, narrow. U.S.A. (Florida), southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 50 species, 14–16 in Venezuela, 8 of these in the flora area. Key to the Species of Brassia 1. 1. 2(1). 2. 3(2).

3.

4(3).

4.

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

6.

7(5). 7.

Petals 1/5–1/4 as long as the lateral sepals ...................................... B. caudata Petals 1/2–2 times as long as the lateral sepals ......................................... 2 Lip 3.1–5 times as long as wide ......................................................B. angusta Lip < 2.5 times as long as wide .................................................................. 3 Lip callosity composed of a pair of glabrous, raised basal ridges or keels and a pair of frontal, isolated, ± well-developed, erect teeth; petals almost straight; ventral face of lip rugulose or verruculose ................... 4 Lip callosity composed of a pair of dense-pubescent ridges or keels and a pair of conspicuous or vestigial frontal teeth; petals variously curved or crossed over dorsal sepal; ventral face of lip smooth ............................ 5 Plants mainly terrestrial or lithophytic; frontal teeth of callosity obtuse or rounded; ventral face of lip basally slightly rugulose; inflorescence erect; plants with 2 or 3 well-developed leaf-bearing sheaths enveloping pseudobulbs ................................................................................... B. bidens Plants epiphytic; frontal teeth of lip acute, ventral face of lip basally conspicuously verruculose; inflorescence arching; plants without leaf-bearing sheaths enveloping pseudobulbs, or only one and this relatively small ........................................................................................... B. neglecta Lip pandurate or elliptic, wider at middle or just above middle; lateral sepals 9–10 times as long as wide ............................................................ 6 Lip without a constriction, slightly wider in the apical 1/4 or 1/3, obovate or oblanceolate; lateral sepals 5–7 times as long as wide ......................... 7 Lip abruptly dilated near middle forming a transversely elliptic central blade, (then abruptly constricted into a narrow-triangular, apical lobe); ridged platform of callosities from lip with an evident cavity apically ............................................................................................. B. lawrenceana Lip gradually dilated near middle and gradually attenuated toward the apical portion; ridged platform of callosities from lip without any evident cavity apically ................................................................... B. macrostachya Lateral sepals sharply incurved; pubescent ridges of callus not separated, almost touching each other, without a cavity in front ........... B. forgetiana Lateral sepals not sharply incurved; pubescent ridges of callus separated by a deep, longitudinal concavity, with a cavity in front ......... B. lanceana

Brassia 251

Brassia angusta Lindl., Edwards’s Bot. Reg. 30: misc. 5. 1844. Epiphyte, cespitose to subcreeping; inflorescences arching to erect; sepals and petals greenish yellow or pale brown with dark brown spots at base; lip white or pale yellowish cream with some small purple or brown spots at base, callus yellow with orange tinge. Rain forests, often near streams, 100– 1300 m; Bolívar (upper Río Paragua basin), Amazonas (Río Casiquiare, Río Siapa, Sierra Parima). Guyana, Brazil. Brassia bidens Lindl., Edwards’s Bot. Reg. 30: misc. 6. 1844. Lithophyte, subterrestrial, or rarely epiphyte, creeping; inflorescence elongate, erect; perianth greenish or yellowish with brown or maroon dots. Rain forest edges, open rocky places, or palm associations, usually growing on sandstone or granitic outcrops, always in exposed positions; 100–1400 m; Bolívar (Río Caroní, Río Parguaza), Amazonas (basins of Río Casiquiare, Río Cataniapo, Río Negro, Río Sipapo, and Río Ventuari). Guyana, Brazil. ◆Fig. 227. This typical Guayanan species has been reported from Peru, but its occurrence there is doubtful. Brassia caudata (L.) Lindl., Bot. Reg. 10: t. 832. 1824. —Epidendrum caudatum L., Sp. Pl. ed. 2, 2: 1349. 1763. Epiphyte, cespitose, shade-loving; flowers showy; perianth green, yellowish, or dull orange, spotted with purple or maroon; lip basally whitish or yellowish, apically with the color of the perianth segments. Rain forests, 50–600 m; Bolívar (Río Cuyuní basin), Amazonas (middle Río Orinoco). Aragua, Barinas, Distrito Federal, Miranda, Sucre, Táchira; U.S.A. (Florida), Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, northern Brazil, Bolivia. ◆Fig. 226. Brassia forgetiana hort. ex C. Schweinf., Bot. Mus. Leafl. 12: 196. 1946. Epiphyte, erect to spreading, ± creeping; flowers with recurved lateral sepals; petals crossed over dorsal sepal; sepals and petals orange-yellow with strong dark brown marking basally; lip white with brown basal spots,

callus orange and white. Rain forests, 300– 600 m; Bolívar (upper Río Caroní, Río Paragua), Amazonas (Río Padamo, Río Siapa). Peru, Amazonian Brazil. ◆Fig. 229. Brassia lanceana Lindl., Edwards’s Bot. Reg. 21: t. 1784. 1836. Epiphyte, cespitose to shortly creeping; lateral sepals straight; petals crossing over dorsal sepal. Rain forests, 100–300 m; Bolívar (Río Caura, Río Paragua), Amazonas (basins of Río Atacavi, Río Siapa, and Río Temi). Amazonian Colombia, Trinidad-Tobago, Guyana, Suriname, Ecuador, Brazil. ◆Fig. 225. Brassia lawrenceana Lindl., Edwards’s Bot. Reg. 27: misc. 2. 1841. Epiphyte, erect, subcreeping; inflorescences arching to nodding; flowers with twisted lateral sepals and petals crossed over dorsal sepal, sepals and petals orange-yellow with purple-brown spots; lip creamy yellow with purple-maroon at apex, callus yellow with maroon-purple spots. Rain forests, often growing on river edges, 400–600 m; Bolívar (Río Caroní, Río Paragua). Guyana, Suriname, Ecuador, Brazil. ◆Fig. 224. Brassia macrostachya Lindl., Sert. Orchid. pl. 6. 1838. Epiphyte, creeping; inflorescence nodding to pendulous, many-flowered; sepals and petals light greenish yellow or orange-yellow with purplish or chocolate-colored marks; lip creamy yellow, yellow, or whitish with a few small, brown spots at base, callus white with orange. Rain forests, 100–500 m; Bolívar (Río Caroní basin), southern Amazonas. Guyana. Brassia neglecta Rchb. f., Allg. Gartenzeitung 24: 322. 1856. Epiphyte, creeping, erect; inflorescences suberect to arching, 3–10-flowered; perianth segments widely spreading; sepals and petals greenish yellow or yellow with brown or purple spots; lip of same color with some purple dots. Rain forests, near sea level to 200 m; Delta Amacuro (Caño Araguao, Ibaruma near Río Cuyubini), Bolívar (Río Paragua basin). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 228.

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Fig. 224. Brassia lawrenceana

Fig. 227. Brassia bidens

Fig. 225. Brassia lanceana

Fig. 228. Brassia neglecta

Fig. 226. Brassia caudata

Fig. 229. Brassia forgetiana

Bulbophyllum 253

18. BULBOPHYLLUM Thouars, Hist. Orchid. 93, t. esp. 3. 1822, nom. cons. [Subtribe Bulbophyllinae]. Bolbophyllaria Rchb. f., Bot. Zeitung (Berlin) 10: 919. 1852. Didactyle Lindl., Fol. Orchid., Didactyle 2. 1852. by Germán Carnevali and Ivón M. Ramírez-Morillo Mostly epiphytes, occasionally lithophytes or subterrestrial herbs, few cm to several m long in some paleotropical creeping species, erect, pendulous, cespitose to creeping. Rhizomes short to more frequently elongate, thin to thick; pseudobulbs heteroblastic (only one internode), variously shaped and sized, usually 4- or 5-ridged or fusiform and ridged, apically 1- or 2(3)-foliate. Leaves conduplicate, articulated with their sheaths, perennial, usually oblong or elliptic, more rarely linear to suborbicular, basally often attenuated into a pseudopetiole. Inflorescences lateral racemes, originating from the pseudobulb base or rarely from the internodes of the rhizomes, 1–many-flowered; rachis sometimes thickened or flattened, elongated or short and then originating a subumbellate inflorescence. Flowers mostly resupinate, inconspicuous, sessile to short-pedicellate, frequently foul-smelling; floral bracts sometimes conspicuous; pedicellate ovary frequently with a pair of lateral bracteoles. Sepals fleshy to membranous, the lateral ones frequently fused into a variously shaped synsepal, which is sometimes the most conspicuous and showy part of the flower, the apices of the sepals frequently setose-elongate, curled or straight, variously colored and ornamented; petals usually smaller than sepals, margins usually ciliate, barbate or with feather-like projections that move with the breeze. Lip hinged to column foot by a narrow claw; blade lobed or unlobed, variously arched, ciliate, barbate, fimbriate, callose-thickened. Column short, erect, basally produced into a foot, apically ornamented or not; anther incumbent, operculate, terminal or subdorsal; clinandrium with 1 or 2 pairs of setose or short projections; pollinia 4, waxy, unappendaged, viscidium absent or rudimentary; rostellum transverse; stigma ventral. Capsules globose, ovoid, or ellipsoid. U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, northern Argentina, Paleotropics (most diverse in Asia and Africa); ca. 2000 species, 14 in Venezuela, 8 of these in the flora area. The recently described Bulbophyllum kegelii Hamer & Garay, known from Suriname, Trinidad-Tobago, and Amazonian Brazil, is expected to occur in the flora area. The other species that will probably eventually be collected within the flora area are B. aristatum (Rchb. f.) Hemsl. and B. bracteolatum Lindl. (type from Guyana). Specimens of the latter species may exist in Venezuelan herbaria but misidentified as B. pachyrhachys. Bulbophyllum bracteolatum is very similar to B. oerstedii (B. pachyrhachis of authors in the Venezuelan Guayana) but has the upper margin of the stelidia entire and extending to the clinandrium. Key to the Species of Bulbophyllum 1. 1.

Pseudobulbs 2-foliate; rachis conspicuously thickened (sepals shorter or as long as rachis thickness) ............................................................ B. oerstedii Pseudobulbs 1-foliate; rachis only slightly thickened or not at all .......... 2

254

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

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

O RCHIDACEAE

Rhizome short; plants minute, mature pseudobulbs < 10 mm long ......................................................................................... B. meristorhachis Rhizome elongate, creeping; plants larger than above ............................. 3 Lip not auriculate; mature leaves usually < 7 cm long; pseudobulbs rounded, pear-shaped, or fusiform ........................................................ 4 Lip basally auriculate, with a conspicuous central contraction; mature leaves usually > 8 cm long; pseudobulbs tetragonal ............................ 5 Plants purple-tinged, especially the pseudobulbs; flowers opening only through lateral slits between lateral and dorsal sepal; petals broader than long or about as broad as long ................................... B. dunstervillei Plants not purple-tinged; flowers totally opened; petals longer than broad .................................................................................................. B. setigerum Rachis conspicuously geniculate; lip barbate; petals shorter than column ................................................................................................. B. meridense Rachis not geniculate or only inconspicuously so; lip glabrous to finely ciliolate; petals longer than column ...................................................... 6 Central lobe of lip glabrous ....................................................... B. roraimense Central lobe of lip ciliolate ......................................................................... 7 Apex of central lobe entire; petals ciliolate ................................ B. exaltatum Apex of central lobe dentate; petals glabrous ............................... B. geraense

Bulbophyllum dunstervillei Garay in Dunst. & Garay, Venez. Orchid. Ill. 6: 70. 1976. Epiphyte, erect, creeping; pseudobulbs rounded to pear-shaped, frequently tinged with red or red-purple; racemes erect, 3–5flowered; lip maroon and white; dorsal sepal 5–6 mm long. Cloud forests, 1200–1400 m; Bolívar (La Escalera to Cerro Venamo region). Endemic. ◆Fig. 230. Bulbophyllum exaltatum Lindl., Ann. Mag. Nat. Hist. 10: 186. 1842. —Didactyle exaltata (Lindl.) Lindl., Fol. Orchid., Didactyle 2. 1852. Epiphyte or lithophyte, sun-loving; raceme elongate; sepals 1–1.5 cm long; sepals and petals greenish or yellowish with maroon or purple blotches or stripes; lip purple. Forests or scrub, 800–1500(–1800) m; Bolívar (basins of Río Caroní, Río Caura, and Río Cuyuní), Amazonas (Río Cunucunuma). Táchira; Guyana, Ecuador, Peru. ◆ Fig. 232. Bulbophyllum geraense Rchb. f. in Walp., Ann. Bot. Syst. 6: 928. 1864. Epiphyte; inflorescence erect; flowers reddish. Cloud forests, ca. 1800 m; Bolívar (Roraima-tepui). Southern Brazil.

Bulbophyllum geraense is very similar to B. exaltatum. It is known from a single, possibly misidentified specimen from the flora area. It is known elsewhere only from southern Brazil. Bulbophyllum meridense Rchb. f., Linnaea 22: 836. 1850. —Didactyle meridensis (Rchb. f.) Lindl., Fol. Orchid., Didactyle 3. 1852. Epiphyte or lithophyte, sun-loving; pseudobulbs 4-ridged; inflorescence erect; flowers with widely spreading sepals and minute petals; sepals to 10 mm long, greenish or yellowish with maroon or purple blotches or stripes; lip reddish or purple. Open scrubland or sandstone outcrops, 900–1400 m; Bolívar (Río Caroní basin), Amazonas (Cerro Duida). Lara, Mérida; Ecuador. Bulbophyllum meristorhachis Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 72. 1976. Twig epiphyte, erect; pseudobulbs ovoid; inflorescence erect, 1- or 2-flowered; flowers minute, campanulate; sepals ca. 8 mm long; perianth greenish with purple longitudinal stripes; lip dark maroon-purple, with orange at base. Rain forests, 500–600 m; Bolívar (Río Churún near Guarimba). Endemic.

Bulbophyllum 255

Bulbophyllum oerstedii (Rchb. f.) Hemsl., Biol. Cent.-Amer., Bot. 3(16): 213. 1884. —Bolbophyllaria oerstedii Rchb. f., Bonplandia (Hanover) 3: 223. 1855. Bulbophyllum pachyrrhachis auct. non (A. Rich.) Griseb. 1864, sensu: Dunst. & Garay, Venez. Orchid. Ill. 1: 65. 1959. Epiphyte, erect; pseudobulbs 4-ridged, 2foliate, (rarely 1-foliate); inflorescence erect to pendulous; flowers almost sessile, purplish, with parallel sepals at anthesis; sepals ca. 5 mm long. Rain forests, rarely low open scrub on sandy soil, 50–500(–1200) m; Bolívar (upper Río Caura basin, Uaipán-tepui), Amazonas (basins of Río Orinoco and upper Río Sipapo). Apure, Aragua, Barinas, Falcón, Miranda; U.S.A. (Florida), Mexico, Central America, West Indies, Colombia, Ecuador.

Bulbophyllum setigerum Lindl., Edwards’s Bot. Reg. 24: misc. 21. 1838. Bulbophyllum correae Pabst, Bradea 1: 180. 1972. Epiphyte, erect; pseudobulbs fusiform, grooved; inflorescence erect with a geniculate rachis; flowers minute; sepals ca. 5 mm long; sepals and petals brown with dark maroon veins; lip brown suffused with maroon. Rain forests, low open scrub or sabanas, 100–200 m; Amazonas (Río Atabapo, Río Casiquiare, Río Negro, Río Sipapo). Guyana, Amazonian Brazil. ◆Fig. 233.

Bulbophyllum roraimense Rolfe, Trans. Linn. Soc. London, Bot. 6: 61. 1901. Epiphyte, erect; pseudobulbs 4-ridged; inflorescence an erect raceme; flowers with perianth segments opening widely; dorsal sepal ca. 15 mm long; sepals and petals greenish, with purple or maroon blotches; lip purple. Open scrub to dense, high forests, 900–2700 m; Bolívar (southern Gran Sabana, Roraima-tepui). Endemic. ◆Fig. 231. Bulbophyllum roraimense is very similar to B. exaltatum and possibly only a subspecific variant of it.

Fig. 230. Bulbophyllum dunstervillei

Fig. 231. Bulbophyllum roraimense

256

O RCHIDACEAE

Fig. 232. Bulbophyllum exaltatum

Fig. 233. Bulbophyllum setigerum

19. CAMPYLOCENTRUM Benth., J. Linn. Soc., Bot. 18: 337. 1881. [Subtribe Angraecinae]. Todaroa Rich., Ann. Sci. Nat. Bot. sér 3, 3: 28. 1845, non Parl. 1843. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, rarely lithophytes, monopodial. Roots frequently rather large, terete or flattened, fleshy, thick, photosynthetic, originating from stem base and from the internodes of the stem. Stems terete or flattened with evident edges, branching or not, elongate to short or essentially absent (plants acaulescent). Leaves conduplicate, terete or reduced to scale-like vestiges or totally absent, articulate with their sheaths, distichous, when present usually elliptic or oblong, subfleshy. Inflorescences originating from stem internodes or subapically when stem is absent, racemes, rarely spikes, usually shorter than leaves, few- to many-flowered, generally dense and usually with flowers arranged to one side of the inflorescence; peduncles terete, frequently verruculose. Flowers small to minute, frequently distichous, usually white but frequently yellowish, greenish, or purplish, short-lived, campanulate, rarely widely opening; floral bracts small, fleshy, persistent; pedicellate ovary usually much shorter than sepals, terete, ridged, frequently pubescent or verruculose. Perianth fleshy, segments usually elliptic or oblong-elliptic, sometimes linear and acute, rarely ovate and obtuse; sepals free or basally connate, subequal or the laterals somewhat oblique, 2–6 mm long; petals usually shorter and narrower than sepals. Lip sessile, fleshy, unlobed or 3-lobed, flat or ± concave, basally produced into a short to very long spur, varied in shape. Column short to very short, exauriculate, footless; anther apical, operculate, incumbent, 2-locular; pollinia 2,

Campylocentrum 257

waxy, ± spherical, tegula usually elongate, occasionally almost absent, viscidium very small; rostellum transverse; stigmatic surface ventral to almost apical. Capsules small, obliquely ellipsoid or ovate-ellipsoid. U.S.A. (Florida), Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina; ca. 30 species, 11 in Venezuela, 9 of these in the flora area. Key to the Species of Campylocentrum 1. 1. 2(1). 2. 3(2). 3. 4(1). 4.

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

Leaves absent or reduced to scale-like projections ................................... 2 Leaves always well developed ................................................................... 4 Plants caulescent; inflorescences originating from sides of well-developed stems ......................................................................................... C. poeppigii Plants acaulescent; inflorescences originating from the center of a bunch of roots .................................................................................................... 3 Roots terete; spur clavate, broader toward apex ........................... C. fasciola Roots dorsoventrally flattened; spur cylindric ..................... C. pachyrrhizum Spur ca. 4 times as long as pedicellate ovary; lip blade sharply 3-lobed with the central lobe ca. 2 times as long than the lateral lobes ...... C. huebneri Spur 1–2 times as long as pedicellate ovary; lip unlobed or inconspicuously 3-lobed with the central lobe at least 3–3.5 times longer than the lateral lobes ............................................................................................ 5 Stems < 1 cm long ...................................................................................... 6 Stems > 3 cm long ...................................................................................... 7 Inflorescences glabrous or glabrescent; leaves longer than stem .............. ............................................................................................... C. hondurense Inflorescences densely pubescent; leaves shorter than the elongate stem .................................................................................................... C. nataliae Leaves 8–11 cm long, 5–6 times as long as wide; spur sharply curving at midlength making a 90° angle to the perianth segments ... C. panamense Leaves 3–5(–8 cm)long, 2–4 times longer than wide; spur parallel to the perianth segments and making an angle < 45° ................................. 8 Spur ± equal to to 2 times as long as the pedicellate ovary; lip lobed near middle ...................................................................................... C. lansbergii Spur ca. 2.5 times as long as pedicellate ovary; lip lobed in basal 1/3 ............................................................................................. C. micranthum

Campylocentrum fasciola (Lindl.) Cogn. in Mart., Fl. Bras. 3(6): 520, t. 106, fig. 1. 1906. —Angraecum fasciola Lindl., Edwards’s Bot. Reg. 26: sub t. 68. 1840. Often twig epiphyte; roots terete; peduncle and rachis scarcely puberulent; flowers white or pale yellow; perianth segments erect and spreading; lip with long spur. Rain forests, 100–400 m; Delta Amacuro (Río Toro), Bolívar (Altiplanicie de Nuria), Amazonas (Río Ventuari, San Carlos de Río Negro). Miranda; Central America, Mexico,

Colombia, Guyana, Suriname, Peru, Brazil, Bolivia. ◆Fig. 234.

Ecuador,

Campylocentrum hondurense Ames, Sched. Orchid. 5: 37. 1923. Campylocentrum steyermarkii Foldats, Acta Bot. Venez. 3: 316, fig. 4. 1968. Epiphyte, erect to subpendulous; leaves arched, apically obliquely lobed; racemes 3– 15-flowered; flowers white, with widely spreading segments, apparently self-fertilized. Rain forests, usually on branches over-

258

O RCHIDACEAE

hanging water, ca. 200 m; Amazonas (Río Mavaca). Apure, Táchira; Mexico, Belize, Honduras, Peru. ◆Fig. 235. Campylocentrum huebneri Mansf., Notizbl. Bot. Gart. Berlin-Dahlem 10: 382. 1928. Campylocentrum uroplectron Pabst, Arq. Bot. Estado São Paulo 3: 135. 1955. Epiphyte or lithophyte; leaves distichous; spikes many-flowered; flowers distichous, yellowish, sometimes with diminutive gold spots; lip spur 2.5–6 times longer than lip blade. Rain forests, 100–200 m; Amazonas (Río Baría, Río Cataniapo, San Carlos de Río Negro, Tamatama). Colombia, Ecuador, Brazil. Campylocentrum lansbergii (Rchb. f.) Schltr., Repert. Spec. Nov. Regni Veg. 6: 100. 1919. —Angraecum lansbergii Rchb. f., Ned. Kruidk. Arch. 4: 316. 1859. Campylocentrum micranthum auct. non (Lindl.) Rolfe 1903: sensu Foldats in Lasser, Fl. Venez. 15(5): 427. 1970, pro parte. Epiphyte or lithophyte, the stem long; leaves coriaceous; inflorescences frequently < 1/2 the leaf length; flowers white; perianth segments not widely opening. Rain forests, 50–900 m; Delta Amacuro (Caño Atoiba north of Caño Araguao, Caño Güiniquina, Río Amacuro), Bolívar (Icabarú, La Escalera). Aragua, Carabobo, Cojedes, Miranda; Brazil. Campylocentrum micranthum (Lindl.) Rolfe, Orchid Rev. 11: 245. 1903. —Angraecum micranthum Lindl., Edwards’s Bot. Reg. 21: t. 1772. 1836. Epiphyte, sometimes a lithophyte; leaves coriaceous; inflorescences usually > 1/2 the leaf length, many-flowered; flowers conspicuously spurred, white, cream, green, lilac, or pink; perianth segments not widely opening. Rain forests, tepui summits and slopes, 50– 1100 m; Delta Amacuro (Caño Capure, Río Acure), Bolívar (Altiplanicie de Nuria, Peraitepui, Río Caura). Aragua, Barinas, Cojedes, Distrito Federal, Miranda, Monagas, Sucre, Portuguesa; Mexico, Central America, West Indies, Suriname, Peru, Brazil. Campylocentrum nataliae Carnevali & I. Ramírez, Biollania Edición Especial 6: 282. 1997.

Epiphyte ca. 4–8 cm tall; flowers whitish, spur cylindric, thickened toward the apex; dorsal sepal 1.4–1.6 mm long; fruit orange. Rain or cloud forests, 300–1000 m; Bolívar (Altiplanicie de Nuria, La Escalera to Cerro Venamo region). Endemic. ◆Fig. 236. Campylocentrum pachyrrhizum (Rchb. f.) Rolfe, Orchid Rev. 11: 246. 1903. —Aeranthus pachyrrhizus Rchb. f., Flora 48: 279. 1865. Epiphyte, acaulescent, leafless; roots long, flattened; racemes 5–10-flowered; flowers white or cream-colored; perianth segments not widely opening. Rain forests, always in deep shade in very humid places, 200–900 m; Bolívar (Auyán-tepui, Santa María de Erebato, Serranía de Imataca). Aragua, Barinas, Falcón; U.S.A. (Florida), Mexico, Central America, West Indies, Guyana, Suriname, French Guiana, Ecuador. ◆Fig. 237. Campylocentrum panamense Ames, Orchidaceae 7: 88. 1922. Campylocentrum micranthum auct. non (Lindl.) Rolfe 1903: sensu Foldats in Lasser, Fl. Venez. 15(5): 427. 1970, pro parte. Epiphyte, shade-loving; leaves coriaceous; inflorescences relatively short, 18–25 mm long; flowers white, lip ca. 3 mm long; spur 3 mm long. Rain forests, on the edges of rivers and waterfalls, 100–600 m; Bolívar (Chalimana rapids on upper Río Paragua), Amazonas (Río Cuntinamo, upper Río Orinoco opposite Caño Pato, Río Padamo at Raudal Caballete). Costa Rica, Panama, Colombia. This rather distinctive taxon has passed as a large-leaved form of the widespread (and possible polyphyletic) Campylocentrum micranthum. The flowers are different, however, particularly in the shape of the spur. The short, thin inflorescences are also distinctive. The species is strikingly disjunct in the flora area from the rest of its range in the area of southern Central America through the Chocó of Colombia, but this might be a reflection of taxonomic lumping and collection artifact. Campylocentrum poeppigii (Rchb. f.) Rolfe, Orchid Rev. 11: 246. 1903. —Angraecum poeppigii Rchb. f., Linnaea 22: 858. 1849. Elongate epiphyte; roots photosynthetic; flowers shortly spurred, white; spikes short.

Campylocentrum 259

Low to high canopies in rain forests, 100–300 m; Bolívar (Río Caura), Amazonas (upper Río Fig. 234. Campylocentrum fasciola

Fig. 236. Campylocentrum nataliae

Orinoco). Monagas, Yaracuy; Mexico, West Indies, Guyana, Ecuador. Fig. 235. Campylocentrum hondurense

Fig. 237. Campylocentrum pachyrrhizum

260

O RCHIDACEAE

20. CATASETUM Rich. ex Kunth, Syn. Pl. 1: 330. 1822. [Subtribe Catasetinae]. Monachanthus Lindl., Edwards’s Bot. Reg. 18: sub t. 1538. 1832. Myanthus Lindl., Edwards’s Bot. Reg. 18: sub t. 1538. 1832. Cuculina Raf., Fl. Tellur. 4: 49. 1836 [1838]. by Gustavo A. Romero-González Epiphytes, often on rotten tree trunks, lithophytic, or sometimes terrestrial herbs. Stems modified into pseudobulbs of several internodes, slender-ovoid or fusiform, when young entirely concealed by distichous, scarious sheaths, eventually naked. Leaves several, distichous, convolute, plicate, articulate, deciduous at the end of the growing season. Inflorescences 1–several, lateral, arising from the base of the pseudobulbs, erect, arching, or pendent, racemose, few- to many-flowered. Flowers unisexual, staminate and pistillate ones often sexually dimorphic, rarely also with polymorphic intermediate ones; floral bracts lanceolate, funnel-shaped. Staminate flowers resupinate or not, green or of various colors, spotted or not, sometimes showy; sepals and petals membranous, the lateral sepals strongly reflexed or not; lip hooded or not, fleshy-thickened; column semiterete, elongate, with 2 filiform processes (antennae) arising from the rostellum, the antennae short or long, bilaterally symmetrical or asymmetrical, one bent behind the other; anther ventral, operculate, unilocular, membranous or fleshy, apiculate or not; clinandrium short to elongate, apiculate; pollinarium with 2 yellow, ovate, cleft, cartilaginous, dorsoventrally flattened pollinia, a tubular, narrowly oblong tegula, and a large, adhesive, circular or elliptic viscidium. Pistillate flowers always nonresupinate, green or yellow-green; sepals and petals fleshy, strongly reflexed; lip helmet-shaped, fleshy-thickened; column semiterete, short and thick, without antennae; pollinarium and anther poorly developed, deciduous at anthesis; stigma a narrow, transverse cleft near the apex. Intermediate flowers usually sterile, color and morphology a continuum between the pistillate and staminate flowers, often combining features of both types of flowers, rarely bilaterally 1/2-pistillate and 1/2-staminate. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina; ca. 120 species and several natural hybrids, 26 in Venezuela, 21 of these plus 4 hybrids in the flora area. Only characters of the staminate flowers (those with fully developed antennae and/or pollinaria) are considered in this key and the species descriptions, as pistillate flowers are sometimes indistinguishable between species. Key to the Species of Catasetum 1. 1. 2(1). 2. 3(2). 3.

Antennae poorly developed, < 3 mm long, thickened, truncate ............... 2 Antennae well developed, ≥ 3 mm long, filiform, bilaterally symmetrical or not ...................................................................................................... 4 Plant pendent, leaves < 3.5 cm wide and usually > 50 cm long ............................................................................................... C. longifolium Plants erect, or leaves > 4 cm wide and usually < 40 cm long ................. 3 Plant lithophytic or terrestrial; inflorescence erect; lip spherical, not dorsoventrally flattened, the margins serrate .................................... C. discolor Plant epiphytic or lithophytic; inflorescence erect or pendent; lip dorsoventrally flattened, the lateral lobes with long fimbriae ...... C. ×roseo-album

Catasetum 261

4(1). 4. 5(4). 5. 6(5). 6. 7(6). 7. 8(5). 8. 9(4). 9. 10(9). 10. 11(10). 11. 12(11). 12. 13(12). 13. 14(11). 14. 15(14).

15.

16(15). 16. 17(9). 17. 18(17).

Antennae bilaterally asymmetrical, one strongly bent behind the other; pollinia < 2 times as long as wide .......................................................... 5 Antennae bilaterally symmetrical, convergent or divergent; pollinia > 2 times as long as wide ....................................................................... 9 Flowers nonresupinate; lip deeply saccate, hood-like, the hood enclosing most of the length of the antennae ....................................................... 6 Flowers resupinate; lip sacciform, cup-shaped, not enclosing the antennae ................................................................................................................ 8 Lip entire at the apical margin, margins reflexed, finely denticulate ....................................................................................................... C. collare Lip lobed at the apical margin, margins not reflexed ............................... 7 Lip apical lobe a subconical tooth ......................................... C. macrocarpum Lip apical lobe a flat or channeled, truncate, obtuse, or retuse flap ............................................................................................... C. ×tapiriceps Lip sack without a transverse callus ............................................ C. pileatum Lip sack with a transverse callus ............................................. C. ×tapiriceps Lip flat to shallowly convex, if with a small sack near the base, then with a single callus at the base or the upper side closely crested .............. 10 Lip deeply saccate, semiglobose, or sacciform with 2 calli at the base, enclosing the antennae ............................................................................ 17 Lip sacciform, fleshy-thickened, without a horn-like basal callus, upper and inner surface closely crested ..................................... C. microglossum Lip flat to shallowly concave, not fleshy-thickened, if sacciform, then with a conspicuous basal callus ................................................................... 11 Lip surface and/or margins barbate or papillose .................................... 12 Lip surface smooth or at most verrucose, the margins smooth, finely dentate or shortly fimbriate ...................................................................... 14 Lip surface papillose .................................................................... C. cristatum Lip surface and/or margins barbate ........................................................ 13 Lip, not including apical, marginal processes, 3/4 as long to longer than the petals ........................................................................................ C. barbatum Lip, not including apical, marginal processes, 1/2 as long as the petals ................................................................................................ C. rivularium Lip narrow, linguiform, basal callus tricuspid .......................... C. bicallosum Lip wide, trulliform, basal callus entire .................................................. 15 Plant flowering from the developing pseudobulb, petal margins ciliate; lip base cordate, margin alongside the basal cavity erect-spreading ............................................................................................. C. parguazense Plant flowering from the fully developed pseudobulb; petal margins smooth; lip base truncate, the margins alongside the basal cavity reflexed .................................................................................................... 16 Antennae long, divergent, not touching each other, reaching and often touching the sides of the lip callus ........................................... C. callosum Antennae short, convergent, touching each other, neither reaching nor touching the lip callus ............................................................. C. yavitäense Inflorescence always erect; lip not 3-lobed, never fimbriate .................. 18 Inflorescence erect or arching; lip 3-lobed or margins fimbriate ........... 21 Lip sack with 6 or 7 parallel, longitudinal lamellae on each side of the inner surface and an apical ridged callus on the apical inner surface .......................................................................................................... C. ferox

262

O RCHIDACEAE

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

24.

25(24). 25.

Lip sack smooth on the inner surface ..................................................... 19 Lip < 2 cm high, slightly emarginate at the apex ..................... C. maroaënse Lip > 2 cm high, not emarginate at the apex .......................................... 20 Plant lithophytic or terrestrial; lip apex narrowly recurved, column with a short clinandrium .................................................................... C. planiceps Plant epiphytic; lip apex broadly spreading or reflexed, column with a long, apicular clinandrium ......................................................... C. ×wendlingeri Inflorescence erect or arching; lip sack without basal calli .................... 22 Inflorescence arching or pendent; lip sack with 1 basal calli or with 2 calli, one on each side of the inner lip, near the base .................................. 23 Inflorescence erect; lip margins entire or finely dentate, not spreading or reflexed, sack without a callus ......................................... C. bergoldianum Inflorescence arching; lip margins fimbriate, spreading or reflexed, sack often with a V-shaped callus near the apex ..................... C. ×dunstervillei Lip with 1 transverse, tricuspid basal callus, margins crenate to dentate in the midlobe, dentate to fimbriate in the lateral lobes ............... C. gomezii Lip with 1 calli on each side of the inner lip near the base .................... 24 Lip hooded, with 2 wide, longitudinal, dentate calli at the base of the lateral lobes, midlobe apiculate, margins of lateral lobes fimbriate, partially enclosing the column ....................................................... C. costatum Lip sacciform, midlobe 3-lobed, with 2 narrow, tooth-like calli at the inner base of the lateral lobes, lateral lobes fimbriate or not, never enclosing the column ............................................................................................ 25 Margins of lateral lobes entire .......................................................... C. bicolor Margins of lateral lobes fimbriate ................................................. C. merchae

Catasetum barbatum (Lindl.) Lindl., Edwards’s Bot. Reg. 30: misc. 38. 1844. —Myanthus barbatus Lindl., Edwards’s Bot. Reg. 21: t. 1778. 1836. Catasetum comosum Cogn., J. Orchidées 6: 266. 1895. Epiphyte; inflorescence arching to pendent; flowers resupinate; sepals and petals green, spotted with maroon; lip margins and upper surface covered with long fimbriae. Wet forests, 50–1200 m; Delta Amacuro (Río Cuyubini), Bolívar (Río Carrao), Amazonas (Río Cataniapo, Río Sipapo, around San Fernando de Atabapo). Monagas; Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (widespread), Bolivia. ◆Fig. 250. Catasetum barbatum is an extremely variable species with at least a dozen synonyms, some grading into C. cristatum Lindl., many of which may be shown eventually to be valid species. Catasetum bergoldianum Foldats, Acta Bot. Venez. 3: 318. 1968. Lithophyte; inflorescences erect; flowers resupinate, yellowish green; lip often with

maroon stripes on inner surface. Lajas, 50– 100 m; Amazonas (near Puerto Ayacucho, Raudal de Maipures, basins of Río Cataniapo and Río Sipapo). Colombia. ◆Fig. 257. Catasetum bicallosum Cogn. in Mart., Fl. Bras. 3(5): 427. 1902. Catasetum stenoglossum Pabst, Arq. Bot. Estado São Paulo 3: 130. 1955. Epiphyte; inflorescence erect to arching; flowers resupinate; sepals and petals green with maroon spots; lip green, with a trifid callus at the base. Known only from the type, reported from the upper Río Orinoco river with no other data. ◆Fig. 252. Catasetum bicolor Klotzsch, Allg. Gartenzeitung 12: 337. 1854. Catasetum gongoroides Kraenzl., Repert. Spec. Nov. Regni Veg. 27: 254. 1930. Epiphyte; inflorescences arching to pendent; flowers resupinate; sepals and petals dark purple to maroon; lip complexly 3-lobed, the midlobe again 3-lobed, light brown, the margin often light green. Gallery forests, wet forests, 50–200 m; Bolívar (Río Suapure),

Catasetum 263

Amazonas (basins of Río Cataniapo and Río Ventuari). Panama, Colombia. ◆Fig. 259. Catasetum callosum Lindl., Edwards’s Bot. Reg. 26: misc. 77. 1840. Epiphyte; inflorescences arching to pendent; flowers resupinate, light green, lightly to densely spotted with maroon; lip with pronounced basal callus. Wet forests, ca. 600 m; Amazonas (Culebra). Coastal Cordillera; Colombia, Guyana, Brazil (Amazonas). ◆Fig. 254. Catasetum collare Cogn., J. Orchidées 6: 154. 1895. Epiphyte; inflorescences erect; flowers nonresupinate; lip greenish white, the margins green, strongly reflexed, finely denticulate. Wet forests, 50–500 m; Bolívar (Canaima), Amazonas (lower Río Cataniapo basin, upper Río Orinoco). Colombia, Ecuador, Brazil (Amazonas). ◆Fig. 247. Catasetum costatum Rchb. f., Gard. Chron. ser. 3, 1: 72. 1887. Epiphyte; inflorescences arching; flowers nonresupinate, light brownish maroon; lip 3lobed, the apical margins of lateral lobes fimbriate, the inner surface with 1 lamellum on each side. Wet forests, 500–1300 m; Amazonas (Cerro Duida, Sierra de la Neblina, Sierra Parima). Brazil (Amazonas). Catasetum cristatum Lindl., Trans. Hort. Soc. London 6: 83. 1824. Epiphyte; inflorescences arching; flowers generally resupinate; sepals and petals green, spotted with maroon; lip green or greenish cream, spotted with reddish maroon, the margins and/or surface with thick, fleshy papillae. Wet forests, 50–1000 m; Bolívar (between El Dorado and Santa Elena de Uairén), Amazonas (Río Cataniapo, around San Fernando de Atabapo). Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil (Amazonas, Pará). ◆Fig. 249. Catasetum cristatum is another variable species with a long list of synonyms, many of which may be shown eventually to be valid species. Catasetum discolor (Lindl.) Lindl., Edwards’s Bot. Reg. 27: sub t. 34. 1841. —Monachanthus discolor Lindl., Edwards’s Bot. Reg. 20: t. 1735. 1834.

Terrestrial or lithophytic; inflorescences erect; flowers orange, yellowish red, yellowish green, or brownish pink; lip margin serrate. Open, lowland and tepui sandy savannas, granitic rock outcrops, 1000–2300 m; Bolívar (Auyán-tepui, Gran Sabana, Roraima-tepui, Sororopán-tepui, Río Churún near Guarimba, Río Uaiparú), Amazonas (Cerro Aracamuni, Cerro Duida). Monagas; Colombia, Guyana, Suriname, French Guiana, widespread in Brazil. ◆Fig. 238. Catasetum ×dunstervillei G.A. Romero & Carnevali, Ann. Missouri Bot. Gard. 76: 458. 1989. Epiphyte; inflorescences arching; flowers resupinate, yellowish orange; lip sometimes spotted with maroon. Gallery and wet forests, 50–100 m; Bolívar (Río Chaviripa), Amazonas (around Puerto Ayacucho). Endemic. ◆Fig. 241. Catasetum ×dunstervillei is a showy natural hybrid between C. pileatum and C. discolor. Catasetum ferox Kraenzl., Gard. Chron. ser. 3, 18: 262. 1895. Epiphyte; inflorescence erect; flowers nonresupinate; sepals green with maroon spots; lip green, with 6 lamellae on its apical inner surface. Open savannas, 100–200 m; Amazonas (Maroa). Brazil (Amazonas). ◆Fig. 256. Catasetum gomezii G.A. Romero & Carnevali, Ann. Missouri Bot. Gard. 76: 455. 1989. Epiphyte; inflorescences arching; flowers resupinate; sepals and petals green, spotted with maroon; lip greenish cream, spotted with reddish maroon, the margins dentate. Wet forests, 50–100 m; Amazonas (Río Cataniapo). Endemic. ◆Fig. 260. Catasetum longifolium Lindl., Edwards’s Bot. Reg. 25: misc. 94. 1839. Epiphyte (always found hanging upsidedown under the leaf crown of the palm Mauritia flexuosa L.); leaves long and narrow, ribbon-like; inflorescence pendent; flowers nonresupinate, greenish orange; lip margin fimbriate. Morichales, 100–700 m; Bolívar (near Santa Elena de Uairén), Amazonas (Río Baría, Río Autana and its tributaries, Río Cuao). Colombia, Guyana, Suriname, French Guiana, Brazil (Amazonas, Pará). ◆Fig. 239.

264

O RCHIDACEAE

Catasetum macrocarpum Rich. ex Kunth, Syn. Pl. 1: 331. 1822. Epiphyte; inflorescences erect; flowers nonresupinate; lip green, often spotted with maroon, the apex 3-lobed, the midlobe a conical tooth. Gallery forests, 50–200 m; Bolívar (Río Caura, Río Suapure), Amazonas (Río Cataniapo). Coastal Cordillera of Distrito Federal and Miranda, Apure, Guárico; Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil (Amazonas, Bahia, Pará, Pernambuco). ◆Fig. 243. Catasetum maroaënse G.A. Romero & C. Gómez, Harvard Pap. Bot. 5: 179. 2000. Epiphyte, indistinguishable vegetatively from other members of the genus, except for the unusually dark green, thick leaves; inflorescences erect; flowers nonresupinate; lip green, yellowish green along the margins and within, the apex slightly emarginate. Humid forests, 100–200 m; Amazonas (near Maroa). Probably also in adjacent Colombia. Catasetum merchae G.A. Romero, Selbyana 10: 73. 1988. Epiphyte; inflorescences arching; flowers resupinate; sepals and petals green, spotted with maroon; lip green, the margins light brown, complexly 5-lobed and margins deeply fimbriate. Wet forests, 50–100 m; Amazonas (Río Cataniapo). Endemic. ◆Fig. 258. Catasetum microglossum Rolfe, Bot. Mag. 139: t. 8514. 1913. Epiphyte; inflorescence erect to arching; flowers resupinate; sepals and petals purple; lip greenish yellow, greatly abbreviate and saccate, with numerous filiform yellow or white appendages. Wet forests, 50–100 m; Amazonas (between Solano and San Carlos de Río Negro). Colombia, Ecuador. ◆Fig. 248. Catasetum parguazense G.A. Romero & Carnevali, Ann. Missouri Bot. Garden 76: 457. 1989. Catasetum bicallosum auct. non Cogn. 1894: sensu Dunst. & Garay, Orchids Venez. Ill. Field Guide 78. 1979. Epiphyte; inflorescence pendent; flowers resupinate; sepals and petals green, tinged with maroon or entirely maroon; lip green, spotted with maroon, the margin ciliate, the upper surface glandulose-verrucose. Wet forests, 50–1000 m; Bolívar

(Río Carrao basin, Río Parguaza). Brazil (Amazonas). ◆ Fig. 253. Catasetum pileatum Rchb. f., Gard. Chron. n.s. 17: 492. 1882. Catasetum bungerothii N.E. Br., Lindenia 2: 21, t. 57. 1886. Epiphyte; inflorescences arching; flowers showy, ivory white, resupinate; lip without a transverse callus. Gallery and wet forests, 50–200 m; Bolívar (Río Parguaza), Amazonas (upper Río Orinoco basin). Apure; Colombia, Brazil (Amazonas, Pará). ◆Fig. 242. Catasetum planiceps Lindl., Edwards’s Bot. Reg. 29: t. 9. 1843. Catasetum recurvatum Link, Icon. Pl. Rar. 2: 105, t. 42. 1844. Catasetum chloranthum Cogn., J. Orchidées 5: 251. 1894. Terrestrial or lithophytic; inflorescences erect; flowers green to yellowish green, nonresupinate. Sandy or rocky savannas, 50– 500 m; Bolívar (Cerro Altamira, Cerro Bolívar, Cerro Toribio, Río Paragua, Roraima-tepui), Amazonas (around Puerto Ayacucho). Coastal Cordillera; Panama, Colombia, Guyana, Brazil (Roraima). ◆Fig. 245. Catasetum rivularium Barb. Rodr., Gen. Spec. Orchid. 1: 130. 1877. Epiphyte; inflorescence arching to pendent; flowers resupinate; sepals and petals dark green, spotted with maroon; lip margins and upper surface covered with long fimbriae. Wet forests, 100–200 m; Amazonas (Caño Cabeza de Manteco, Río Autana). Brazil (Amazonas). ◆Fig. 251. Catasetum ×roseo-album (Hook.) Lindl., Edwards’s Bot. Reg. 26: misc. 65. 1840. —Monachanthus roseo-albus Hook., Bot. Mag. 67: t. 3796. 1840. Epiphyte (most often on a stem of the palm Mauritia flexuosa L. f.) or lithophyte; inflorescence erect, arching to pendent; flowers greenish white, yellow, pink, red, or maroon; lip lateral margins conspicuously fimbriate. Morichales, lajas, 50–100 m; Bolívar (Canaima, Río Parguaza), Amazonas (La Esmeralda, near Puerto Ayacucho, basins of Río Cataniapo and Río Sipapo). Colombia, Guyana, Suriname, French Guiana, widespread in northeastern Brazil. ◆Fig. 240. Catasetum ×roseo-album is a natural hybrid between C. discolor and C. longifolium.

Catasetum 265

Catasetum ×tapiriceps Rchb. f., Gard. Chron. ser. 3, 3: 136. 1888. Catasetum splendens Cogn., J. Orchideés 5: 302. 1894. Catasetum apertum Rolfe, Bull. Misc. Inform. Kew 1895: 284. 1895. Epiphyte; inflorescences erect, arching to pendent; flowers green, white, greenish white, or yellow, often spotted with maroon; lip with a prominent, transverse callus near the apex, the apex entire or 3-lobed, if 3lobed, the midlobe a flat or channeled, truncate, obtuse, or retuse flap. Flooded and gallery forests, 50–100 m; Bolívar (Río Parguaza), northeastern Amazonas. Apure; Colombia, Brazil (Amazonas). ◆Fig. 244. Catasetum ×tapiriceps is a natural hybrid between C. macrocarpum and C. pileatum. Catasetum ×wendlingeri Foldats, Acta Biol. Venez. 2: 167. 1958. Epiphyte; inflorescence erect to arching; flowers greenish yellow to yellowish orange; lip margin sometimes shallowly fimbriate, the apex conspicuously recurved. Wet forests, 50–100 m; Bolívar (Caripo), Amazonas (around Puerto Ayacucho). Endemic. ◆Fig. 246. Catasetum ×wendlingeri is a natural hybrid between C. pileatum and C. planiceps. Catasetum yavitaënse G.A. Romero & C Gómez, Harvard Pap. Bot. 3(1): 54. 1998. Epiphyte; inflorescence arching; flowers green to dark maroon. Wet forests, 100–200 m; Amazonas (Caño Pasa, Río Cataniapo basin, near Yavita). Endemic. ◆Fig. 255.

Staminate flowers

Fig. 238. Catasetum discolor

266

O RCHIDACEAE

Fig. 239. Catasetum longifolium

Fig. 240. Catasetum ×roseoalbum [C. discolor × C. longifolium]

Fig. 241. Catasetum ×dunstervillei [C. discolor × C. pileatum]

Fig. 242. Catasetum pileatum

Catasetum 267

Fig. 243. Catasetum macrocarpum

Fig. 245. Catasetum planiceps

Fig. 248. Catasetum microglossum

Fig. 244. Catasetum ×tapiriceps [C. macrocarpum × C. pileatum]

Fig. 246. Catasetum ×wendlingeri [C. pileatum × C. planiceps]

Fig. 249. Catasetum cristatum

Fig. 247. Catasetum collare

Fig. 250. Catasetum barbatum

268

O RCHIDACEAE

Fig. 251. Catasetum rivularium

Fig. 252. Catasetum bicallosum

Fig. 253. Catasetum parguazense

Fig. 254. Catasetum callosum

Fig. 256. Catasetum ferox

Fig. 255. Catasetum yavitaënse

Pistillate flower

Staminate flower Fig. 257. Catasetum bergoldianum

Fig. 258. Catasetum merchae

Fig. 259. Catasetum bicolor

Cattleya 269

Fig. 260. Catasetum gomezii

21. CATTLEYA Lindl., Coll. Bot. pls. 33, 37. 1824. [Subtribe Laeliinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Mostly epiphytes or lithophytes, rarely subterrestrial herbs, cespitose to creeping, mostly sun-loving. Rhizomes relatively thick; pseudobulbs heteroblastic (one internode) to homoblastic (many internodes), apically 1–3(4)-foliate, oblong, cylindric, or ellipsoid, terete or laterally compressed. Leaves conduplicate, articulate, coriaceous to thickly fleshy-coriaceous, erect or spreading, elliptic, ovate-elliptic to obovate, acute to rounded and obliquely 2-lobed at apex, sessile to subpetiolate. Inflorescences racemose, terminal on pseudobulb apex or originating at apex of a small aborted and leafless pseudobulb and then appearing to originate from rhizome, sometimes dense, 1–many-flowered, generally subtended by a conduplicate, elliptic to obovate spathe; peduncle generally stout; rachis thick, relatively short. Flowers resupinate, medium- to usually large-sized, showy, long-lasting, sometimes cleistogamous, fragrant; floral bracts small and inconspicuous, concave; pedicellate ovary terete, or somewhat 3-edged at apex. Perianth segments fleshy to submembranous, generally widely spreading; sepals subequal, free, mostly elliptic or elliptic-obovate, petals subequal to sepals, often broader. Lip erect, unlobed or 3-lobed, basal margins or lobes generally enfolding the column, rarely flat, apical portion of lobe erect to decurved and spreading, margins entire to fimbriate or wavy, apex rounded, truncate, emarginate, or obtuse, basally rounded or subcordate. Column erect, hemicylindric or clavate, stout, apically ± winged or not, curved or straight, footless; anther terminal, operculate, incumbent, 2-locular, each locule with a longitudinal septum; clinandrium with 1–3 teeth which partially conceal the anther; pollinia 4 (rarely 8), caudicles yellow; rostellum transverse; stigma ventral, transverse. Capsule ellipsoid, oblong, or subglobose, often with prominent ridges. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 45 species, 7 or 8 species in Venezuela, 3 of these in the flora area. Showy, often-cultivated orchids, the Cattleyas apparently have been subject to over-splitting based on inconspicuous features, especially in the Cattleya labiata

270

O RCHIDACEAE

complex, which may contain fewer than the currently accepted species. However, most of the recognized taxa are consistent within populations and may merit recognition as specific entities. Recent analyses of DNA sequence variation clearly indicates that many southeastern Brazilian taxa, formerly treated as members of the polyphyletic genus Laetia Lindl. (now restricted to Mesoamerica), would be better referred to Cattleya. Many Cattleya species are threatened with extinction due to overcollecting for horticultural purposes and due to habitat perturbation. Cattleya eldorado Linden has been collected in the state of Amazonas in Brazil near the Venezuelan border and likely occurs within the flora area. If found it would be most likely confused with C. jenmanii, with which it shares the 1-leaved flowers. However, C. eldorado is a smaller plant with an obovate lip, whereas the lip in C. jenmanii is elliptic. Key to the Species of Cattleya 1. 1. 2(1). 2.

Pseudobulbs 2- or 3-foliate; lip distinctly 3-lobed ......................... C. violacea Pseudobulbs 1-foliate; lip unlobed ............................................................. 2 Lip elliptic ...................................................................................... C. jenmanii Lip oblong, abruptly expanded at the apical 1/2 ..................... C. lawrenceana

Cattleya jenmanii Rolfe, Bull. Misc. Inform. Kew 1906: 85. 1906. Epiphyte or lithophyte, cespitose or creeping, erect, sun-loving to semishade-loving; flowers showy, in various shades of lavender or purple; lip with yellow and white zones as well as deep purple veins and a solid deep purple apical portion. Rain forests, 600–1100 m; Bolívar (basin of upper Río Caroní, Río Caura). Guyana. ◆Fig. 261. Cattleya lawrenceana Rchb. f., Gard. Chron. n.s. 23: 338. 1885. Epiphyte or lithophyte, cespitose to shortly creeping, sun-loving; flowers showy but medium-sized for the genus, purple with a deep purple apical zone on lip and some white and yellow in the throat. Rain or cloud forests or open rocky places, 400–1900 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Gran Sabana), Amazonas (Cerro Sipapo). Guyana, Brazil (Roraima). ◆Fig. 262.

Fig. 261. Cattleya jenmanii

Cattleya violacea (H.B.K.) Rolfe, Gard. Chron. ser. 3, 5: 802. 1889. —Cymbidium violaceum H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 341. 1815 [1816]. —Superba del Orinoco. Cattleya superba Schomb. ex Lindl., Sert. Orchid. pl. 22. 1838. Epiphyte, cespitose or shortly creeping, erect to arching; inflorescences 1–8-flowered; flowers showy, purple to deep purple, rarely pink; lip 3-lobed, deep purple. Rain forests, near sea level to 300(–600) m; widespread in the flora area. Apure; Amazon basin of Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 263.

Caularthron 271

Fig. 262. Cattleya lawrenceana

Fig. 263. Cattleya violacea

22. CAULARTHRON Raf., Fl. Tellur. 2: 40. 1836 [1837]. [Subtribe Laeliinae]. Epidendrum sect. Diacrium Lindl., J. Bot. (Hooker) 3: 81. 1841. Diacrium Lindl. ex Benth., J. Linn. Soc., Bot. 18: 312. 1881. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes or lithophytes, 25–150 cm tall in flower, erect to arching, mostly sunloving, cespitose. Rhizomes short but well developed, stout; pseudobulbs homoblastic (many internodes), fusiform to subcylindric, always internally hollow, basally stipitate and with 1 or 2 holes that often serve as entrances to ant colonies, with 3–7 distichous to subopposite leaves at the apex. Leaves conduplicate, articulate, fleshycoriaceous, oblong or oblong-elliptic, obtuse and emarginate. Inflorescences terminal, racemose or very rarely with 1 branch, laxly or ± densely few- to many-flowered, erect, stout; peduncle terete, elongate; rachis erect. Flowers resupinate, showy, lasting a few days, successive, with widely spreading segments, sometimes cleistogamous; floral bracts inconspicuous; pedicellate ovary terete, longer than perianth segments. Perianth white, often with a pink tinge; sepals free, subequal, oblong or ob-

272

O RCHIDACEAE

long-elliptic, acute, 10–32 mm long; petals subequal to sepals but often broader. Lip free, erect or ± spreading, unlobed or 3-lobed, basally with 2 calli; calli tooth-shaped, horn-shaped, or somewhat layered, with a cavity opening on the lip underside. Column short, arched, footless, concave, winged; anther terminal or subventral, operculate, incumbent, 2-locular or subtetralocular; clinandrium oblique, obtuse; pollinia 4, cartilaginous, laterally flattened with yellow caudicles; rostellum transverse; stigma ventral, suborbicular. Capsule ellipsoid. Central America, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 2 species, both in the flora area. Key to the Species of Caularthron 1. 1.

Lip deeply 3-lobed; petals (0.8–)15–23 mm wide ..................... C. bicornutum Lip unlobed or inconspicuously 3-lobed; petals < 11 mm wide ................... ............................................................................................C. bilamellatum

Caularthron bicornutum (Hook.) Raf., Fl. Tellur. 2: 41. 1836 [1837]. —Epidendrum bicornutum Hook., Bot. Mag. 61: t. 3332. 1834. —Diacrium bicornutum (Hook.) Benth., J. Linn. Soc., Bot. 18: 312. 1881. Diacrium amazonicum Schltr., Beih. Bot. Centralbl. 42(2): 108. 1925. Epiphyte or lithophyte, erect, sun-loving; flowers white with red spots on lip and sometimes on petals and sepals, very fragrant; sepals 20–32 mm long. Moist and semideciduous forests, 100–300 m; Bolívar (Rio Icabarú, Rio Uaiparú), Amazonas (basin of Río Casiquiare, Río Cataniapo, Río Negro, Río Ventuari). Monagas, Sucre; Colombia, Trinidad-Tobago, Brazil (Amazonas, Pará). ◆Fig. 264. Caularthron bicornutum is a showy species that is much sought after in cultivation. Although the size ranges overlap somewhat, it is usually much more robust than C. bilamellatum. Caularthron bilamellatum (Rchb. f.) R.E. Schult., Bot. Mus. Leafl. 18: 92, pls. 14, 15. 1958. —Epidendrum bilamellatum Rchb. f. in Walp., Ann. Bot. Syst. 6: 345. 1862. —Diacrium bilamellatum (Rchb. f.) Hemsl., Biol. Cent.-Amer., Bot. 3: 222. 1883. Diacrium venezuelanum Schltr., Repert. Spec. Nov. Regni Veg. 6: 41. 1919. Fig. 264. Caularthron bicornutum

Chaubardia

Epiphyte, or rarely a lithophyte, erect to ascendent; flowers white, sometimes suffused with pink on outside, very rarely entirely pale pink, fully opening or cleistogamous; sepals 10–21 mm long. Locally common in exposed situations in rain forests, 200–400 m; Bolívar (Altiplanicie de Nuria, near Puerto Ordaz, Represa Guri, lower Río Caura basin). Apure, Aragua, Barinas, Carabobo, Distrito Federal, Guárico, Lara, Monagas, Sucre, Zulia; Guatemala, Belize, Honduras, Costa Rica, Panama, Colombia, Trinidad-Tobago, Ecuador, Peru, Brazil. Caularthron indivisum (Bradf.) Garay & Dunst. was treated at the species level in Dunst. & Garay, Orchids Venez. Ill. Field Guide 107. 1979 (basionym: Epidendrum

273

indivisum Bradf. ex Griseb., Fl. Brit. W. I. 614. 1864). However, it is more likely just a peloric, actinomorphic form of Caularthron bilamellatum, which is zygomorphic, and we are here making the new combination for that form: Caularthron bilamellatum (Rchb. f.) R.E. Schult. f. indivisa (Bradf.) Carnevali & I. Ramírez, comb. nov. It is recognized nomenclaturally since it seems to occur sporadically along the distribution range of the species and has been collected several times and in at least three different countries. It is distinguished from the typical form by having the lip very similar to the petals and without well-developed callosities, whereas the typical form has the lip different from the petals and with callosities.

23. CHAUBARDIA Rchb. f., Bot. Zeitung (Berlin) 10: 671. 1852. [Subtribe Zygopetalinae]. by Gustavo A. Romero-González Cespitose epiphytes. Stem abbreviate, modified into very small, aphyllose pseudobulbs, entirely concealed by leaf sheaths. Leaves several, distichous, membranous, narrowly oblong-ovate to oblong-obovate, abruptly acute to acuminate, plicate, articulate. Inflorescences lateral, arising from the axil of the leaf sheaths, erect to arching, 1-flowered, shorter than the leaves. Flowers resupinate; floral bracts concave, appressed, ovate-triangular to tubular-funnel-shaped, < 1/2 as long as the pedicellate ovary. Sepals and petals membranous, erect-spreading to spreading, the margins sometimes strongly recurved; dorsal sepal narrowly oblong-ovate to oblongelliptic; lateral sepals oblong-elliptic to narrowly ovate, oblique, the inner basal margins sometimes strongly inrolled; petals broadly ovate to oblong-obovate, sometimes slightly oblique; lip clawed, the claw short and broad or long and narrow, S-shaped, keeled, blade entire, auriculate, apiculate, or divided into a conduplicate, concave hypochile and a narrowly elliptic-ovate, acuminate, slightly concave epichile attached to the abaxial surface of the hypochile; disk with a radially keeled, semicircular callus. Column elongate, slightly arched, semiterete, broadly winged in the lower 1/2–2/3, produced into a short foot, sometimes abaxially and basally pubescent, the rostellum transverse, plate-like, emarginate; anther terminal; pollinia 4, yellow, cartilaginous, dorsoventrally flattened, subequal, obovate, attached in 2 pairs to a very short, apparently trapeziform tegula. Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 3 species, 1 in Venezuela. Chaubardia is closely related to Chaubardiella, but is distinguished by the presence of small, aphyllous or inconspicuous pseudobulbs and a transverse, platelike, emarginate rostellum.

274

O RCHIDACEAE

Chaubardia surinamensis Rchb. f., Bot. Zeitung (Berlin) 10: 672. 1852. Hoehneella santos-nevesii Ruschi, Publ. Arq. Publ. Estado Espíritu Santo 3. 1945. Epiphyte; pseudobulbs compressed, small (to 1 cm), generally concealed by foliar sheaths; inflorescences 1-flowered; sepals and petals green; lip white. Wet forests, 300–500 m; Bolívar (upper Río Paragua, Río Erebato, Salto Chalimano on Río Paramichi). Monagas; Guyana, Suriname, Ecuador, Brazil (Espírito Santo). ◆Fig. 265. Lip Fig. 265. Chaubardia surinamensis

24. CHAUBARDIELLA Garay, Orquideología 4: 146. 1969. [Subtribe Zygopetalinae]. by Gustavo A. Romero-González Cespitose epiphytes. Stem abbreviate, not modified into pseudobulbs. Leaves several, distichous, membranous, narrowly oblong-ovate to oblong-obovate, acute, plicate, articulate. Inflorescences lateral, arising from the axil of the leaf sheaths, arching to subpendent, 1-flowered, shorter than the leaves. Flowers resupinate or not; floral bracts tubular-funnel-shaped, cucullate, < 1/2 as long to almost as long as the pedicellate ovary. Sepals and petals subfleshy, spreading, the margins sometimes strongly recurved; sepals narrowly ovate to elliptic-obovate; dorsal sepal acute, the lateral sepals short-acuminate, sometimes slightly oblique and/or the inner basal margins strongly inrolled; petals obovate to elliptic-obovate, sometimes slightly oblique; lip sessile, articulate with the column foot, fleshy, rigid, basally subglobose, saccate, or cochleate, with a reflexed, acute or obtuse apex, entire or 2lobed, if 2-lobed the 2 lobes falcate or not, and with a transverse, antrorse, subtriangular or semicircular, entire or 2-lobed callus, the callus sometimes subquadrate with both sides with sharp teeth, or shallow, truncate, plate-like, bifid above the base. Column short, thickened, semiterete, apically winged or not, produced into a short foot, the rostellum filiform, elongate, arched; anther terminal; pollinia 4, yellow, cartilaginous, dorsoventrally flattened, subequal, obovate or linear-obovate, attached in 2 pairs to a common point on a subtriangular, sometimes hooked tegula. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru; 9 or 10 species, 1 in Venezuela. Chaubardiella is closely related to Chaubardia Rchb. f. and Stenia Lindl. but, in the flora area, it is distinguished from Chaubardia by the absence of pseudobulbs and the filiform rostellum, and from Stenia by the lip articulate with the column foot.

Cheiradenia 275

Chaubardiella tigrina (Garay & Dunst.) Garay, Orquideología 4: 149. 1969. —Chaubardia tigrina Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 2: 72. 1961. —Stenia tigrina (Garay & Dunst.) Foldats, Acta Bot. Venez. 3: 423. 1968. Plant without pseudobulbs; inflorescence 1-flowered; flowers pale green-brown with transverse maroon stripes; lip callus waxy, yellow-brown, with about 10 radiating ridges. Wet forests, 600–800 m; Bolívar (La Escalera to Cerro Venamo region, base of Salto Angel). Colombia, Ecuador, Peru. ◆Fig. 266.

Fig. 266. Chaubardiella tigrina

25. CHEIRADENIA Lindl., Fol. Orchid., Cheiradenia 1. 1853. [Subtribe Zygopetalinae]. by Gustavo A. Romero-González Cespitose epiphytes. Stem very short, modified into small pseudobulbs of several internodes. Leaves several, distichous, membranous, narrowly oblong-ovate to oblong-obovate, acuminate, plicate, articulate. Inflorescences lateral, arising from the axil of the leaves, erect, 1–few-flowered, racemose, rarely branched, as long as to longer than the leaves. Flowers resupinate; floral bracts concave, acuminate, < 1/2 as long as the pedicellate ovary. Sepals and petals membranous, concave, spreading; sepals elliptic to obovate; lateral sepals sometimes slightly oblique; petals widely ovate to elliptic. Lip membranous, articulate with the column foot, entire or somewhat 3-lobed; disk with a transverse, antrorse, excavated, dentate ridge. Column short, semiterete, subclavate, produced into a short foot; anther terminal; pollinia 4, yellow, cartilaginous, dorsoventrally flattened, subequal, obovate, attached in 2 pairs to a trullate tegula. Venezuela, Guyana, Suriname, French Guiana, Brazil; 1 species.

276

O RCHIDACEAE

Cheiradenia cuspidata Lindl., Folia Orchid., Cheiradenia 1. 1853. Cheiradenia imthurnii Cogn. in Mart., Fl. Bras. 3(6): 481. 1906. Small epiphyte (pseudobulbs to 1 cm and leaves to 10 cm); leaves velvety green; inflorescence erect; flowers ≤ 4, pinkish white with some dark maroon markings. Wet forests, 300–900 m; Bolívar (Altiplanicie de Nuria, Campamento La Reforma–CVG, La Escalera to Cerro Venamo region, Río Aonda on Auyán-tepui, Río Chicanán, Río Paragua, base of Salto Angel, Serranía de Imataca). Guyana, Suriname, French Guiana, Brazil (Amapá). ◆Fig. 267.

Fig. 267. Cheiradenia cuspidata

26. CHELYORCHIS Dressler & N.H. Williams in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1130. 2000. [Subtribe Oncidiinae]. Oncidium sect. Oblongata Kraenzl. in Engl., Pflanzenr. IV. 50(Heft 80): 233. 1922, pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytic herbs. Rhizome abbreviate; stems pseudobulbous; pseudobulbs heteroblastic (one internode), ancipitous (compressed laterally), appressed over the bark of the host tree, suborbicular, the surface bumpy, often purple spotted, basally clothed with several sheaths lacking foliar blades, apically 1- or 2(3)-foliate. Leaves conduplicate, articulate with their sheaths, fleshy-coriaceous. Inflorescences lateral, erect, originating from base of the pseudobulb, usually a panicle, more rarely a raceme, usually surpassing the leaves, peduncle and rachis terete; floral bracts inconspicuous; pedicellate ovary conspicuous, usually terete. Flowers resupinate, relatively large and showy, long lasting, bright yellow, the sepals blotched or barred with maroon, the petals spotted with the same color, lip bright yellow, callus yellow with fine purple spots; perianth segments spreading, subfleshy, the sepals concave, subequal, free, petals larger and showier than the sepals. Lip adnate to the base of the column at the very base, spreading, larger and showier than sepals and petals, the base forming a right angle to the column, then flexed downward and forming a 180° angle to the column, blade entire or the lateral lobes very reduced, central lobe spreading, transversely oblong , suborbicular or reniform, apically emarginate; disk conspicuously callose, the callus fleshy, composed of 5 teeth, 2 transversal pairs and a central, longitudinal one. Column relatively short, footless, erect; lateral margins near the stigma with a pair of small, flabellate-reniform wings; anther subventral, operculate, incumbent, strongly convex, semiglobose, 1-locular or imperfectly 2locular; clinandrium petaloid, small, shallowly 3-dentate: pollinia 2, narrowly oblong, stipes reniform, short and wide, viscidium relatively small.

Chelyorchis 277

Guatemala to Panama, Colombia, Venezuela, Trinidad-Tobago, Ecuador, Peru; 1 species. The single species of this genus has long been suspected not to be related to other members of Oncidium. It has been recently segregated from Oncidium based on molecular evidence. This also indicated that Chelyorchis is related to a clade formed by various Mesoamerican taxa, currently placed in the genera Osmoglossum (Schltr.) Schltr. and Palumbina Rchb. f. and not to the core of Oncidium. Chelyorchis ampliata (Lindl.) Dressler & N.H. Williams in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide, 1130. 2000. —Oncidium ampliatum Lindl., Gen. Sp. Orchid. Pl. 202. 1833. Oncidium bernouillanum Kraenzl. in Engl., Pflanzenr. IV. 50(Heft 80): 231. 1922. Epiphyte, usually sun-loving, erect; leaves 8–50 cm long; flowers pale or bright yellow with bright brown or maroon spots on base of petals and lip, lip 12–24 × 16–32 mm. Dry or deciduous, rarely humid forests, near sea level to 200 m; Delta Amacuro (Río Amacuro), Bolívar (lower Rio Caroní). Aragua, Barinas, Distrito Federal, Guárico, Lara, Monagas, Yaracuy, Zulia; extra-Venezuelan distribution as in genus. ◆Fig. 268. The thickly coriaceous, flattened pseudobulbs, which are usually purple-spotted and appressed to the host’s bark, are unique to this species. The plant is widely cultivated for its showy flowers. Large, well-cultivated plants make spectacular horticultural specimens.

Fig. 268. Chelyorchis ampliata

278

O RCHIDACEAE

27. CLEISTES Rich., Mém. Mus. Hist. Nat. 4: 31. 1818. [Subtribe Pogoniinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial or humicolous herbs, erect, mostly sun-loving, often in boggy or marshy areas. Roots fibrous, fleshy or tuberous. Stems unbranched or rarely branching, subcylindric. Leaves convolute, not articulate, subfleshy to rarely subcoriaceous, distichous to ± spiraled, solitary to several per stem, the leaves or the whole plant frequently glaucous; sheaths clasping; blades sometimes reduced and bract-like or totally absent. Inflorescences racemose, terminal, few-flowered, or the flowers solitary and axillary at base of leaf-like bracts. Flowers resupinate, shortlived, usually showy, greenish, yellow, white, purple, pink, or red; floral bracts similar to the leaves but smaller; pedicellate ovary with an abscission layer below the perianth, relatively long, subterete. Perianth segments membranous to subfleshy, mostly campanulate; sepals free, usually narrowly elliptic or narrowly oblong-elliptic, the laterals somewhat oblique, sometimes widely spreading; petals similar to sepals, often wider and shorter or differently colored, parallel to column. Lip free from column and parallel to it, basally with 2–4 pedunculate calli, unlobed or 3lobed, flat or more frequently concave, inner surface generally with well-developed longitudinal calli or lamellae, frequently tuberculate. Column relatively long, erect or forming an angle with respect to ovary, usually tapering basally, apically lobed or unlobed, footless; anther terminal, incumbent; clinandrium mostly 3-lobed or otherwise lobed or lacerate; pollinia 2, pollen soft and mealy, coherent in tetrads, without viscidium; rostellum transverse; stigma entire, ventral. Capsule ellipsoid. U.S.A. (Florida, Louisiana, Kentucky, North Carolina), Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, southern Brazil; ca. 30 species, 9 in Venezuela, 8 of these in the flora area. Plants of this genus inhabit open places such as savannas, rocky outcrops, or bogs. The aerial portion of the plant is seasonal or very rarely perennial, depending on the length of the dry season. Key to the Species of Cleistes 1. 1. 2(1). 2. 3(1). 3. 4(3). 4. 5(3). 5. 6(5).

Leaves very reduced, sheath-like, or absent; flowers pale yellow or rarely white ....................................................................................................... 2 Leaves well developed; flowers mostly pink, purple, red, or white, rarely pale yellow .............................................................................................. 3 Plants often < 10(–15) cm tall; dorsal sepal 12–14 mm long; lip 3-lobed ....................................................................................................... C. huberi Plants 23–80 cm tall; dorsal sepal 18–20 mm long; lip unlobed ..... C. stricta Lip unlobed ................................................................................................. 4 Lip 3-lobed .................................................................................................. 5 Lip 3.5 cm long; on tepui summits or slopes above 1600 m ....... C. parviflora Lip (4–)5 cm long; in savannas or ecotones below 1400 m ................ C. rosea Lip 22 mm long; flowers mostly pink or purple ........................................ 6 Lip 17 mm long; flowers yellow or white................................................... 7 Basal 1/2 of longitudinal callus of lip covered with membranous keels; clinandrium margin laciniate ....................................................... C. lepida

Cleistes 279

6. 7(5).

7.

Basal 1/2 of longitudinal callus of lip without membranous keels; clinandrium margin not laciniate ............................................... C. triflora Leaves several; inflorescences 1–4-flowered; sepals and petals white; sepals at least 7 times longer than wide; plants of lowland savannas, usually below 600(–900) m .................................................................. C. tenuis Leaf solitary; inflorescence 1-flowered; sepals and petals greenish yellow to yellow; sepals ca. 5 times longer than wide; plants of tepui summits or slopes, 900–2200 m ............................................................. C. unifoliata

Cleistes huberi Carnevali & I. Ramírez, Ann. Missouri Bot. Gard. 77: 549. 1990. Terrestrial to humicolous; appressed to the stem; inflorescences 1–3-flowered; flowers campanulate, creamy white to pale yellow; dorsal sepal to 12 mm long. Savannas or scrublands on tepui summits, 2100–2800 m; Bolívar (Aparamán-tepui, Kukenán-tepui, Roraima-tepui, Tramen-tepui, Yuruaní-tepui). Endemic. ◆Fig. 269. Cleistes lepida (Rchb. f.) Schltr., Arq. Bot. Estado São Paulo 1: 179. 1926. —Pogonia lepida Rchb. f., Xenia Orchid. 2: 90. 1865. Cleistes moritzii (Rchb. f.) Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 4: 54. 1966. —Pogonia moritzii Rchb. f., Xenia Orchid. 2: 89. 1865. Terrestrial to 1 m tall; flowers opening 2 or 3 successively, whitish or pink. Savannalike associations on tepuis; 2000–2300 m; Bolívar (Macizo del Chimantá [Toronótepui]). Ecuador, Brazil. The Venezuelan collections may represent a different species. Cleistes parviflora Lindl., Gen. Sp. Orchid. Pl. 410. 1840. —Pogonia parviflora (Lindl.) Rchb. f., Xenia Orchid. 2: 90. 1865. Terrestrial to humicolous, 30–100 cm tall; flowers campanulate, deep purple or reddish. Trachypogon savannas, forest ecotone and savannas on tepuis (1100–)1500–2600 m; common in Bolívar and Amazonas. Guyana, Suriname, French Guiana, Brazil. Cleistes rosea Lindl., Gen. Sp. Orchid. Pl. 410. 1840. —Pogonia rosea (Lindl.) Rchb. f., Xenia Orchid. 2: 89. 1865. Terrestrial to 1.5 m tall; inflorescences 1– 3-flowered; sepals 4–7 cm long, widely

spreading; petals subparallel to column; lip with a bright yellow longitudinal callus. Marshy open places, savannas, forest ecotones, open tepui associations, 50–1300 m; Bolívar (widespread), Amazonas (widespread). Aragua, Distrito Federal, Mérida, Miranda, Yaracuy; Costa Rica, Panama, Colombia, Trinidad-Tobago, Guyana, Ecuador, Peru, Brazil. ◆Fig. 270. Two forms are known from the flora area, f. rosea with rose, pink, or purple petals and f. pallida Carnevali & I. Ramírez with greenish cream or greenish white petals. Cleistes stricta (C. Schweinf.) Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 5: 54. 1972. —Pogonia stricta C. Schweinf., Fieldiana, Bot. 28: 167, fig. 25. 1951. Terrestrial; leaves absent or bract-like and then narrowly elliptic; flowers small for the genus, not widely opening; sepals yellow, 15–20 mm long; petals white; lip white with purple apex. Locally common in savannas on tepuis, often in swampy places, 1500–2200 m; Bolívar (Jaua-Sarisariñama massif, Macizo del Chimantá), Amazonas (Cerro Aracamuni, Sierra de la Neblina). Endemic. Cleistes tenuis (Rchb. f.) Schltr., Arq. Bot. Estado São Paulo 1: 180. 1926. —Pogonia tenuis Rchb. f., Xenia Orchid. 2: 91. 1865. Slender terrestrial to 50 cm tall; perianth white; sepals 15–19 mm long; lip with purple oblique stripes. White-sand, often swampy, savannas 50–400(–1200) m; Bolívar (Río Uairén), Amazonas (basin of Río Casiquiare, Río Negro, Río Sipapo). Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil. Cleistes triflora (C. Schweinf.) Carnevali & I. Ramírez in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1130.

280

O RCHIDACEAE

2000. —Pogonia triflora C. Schweinf., Fieldiana, Bot. 28: 169, fig. 26. 1951. Herb to 70 cm tall; flowers 2 or 3, campanulate, opening in slow succession; perianth pink or pale purple; sepals to 34 mm long; lip with purple stripes. Widespread in exposed places on Trachypogon and tepui savannas, 1300–1900 m; Bolívar (Auyán-tepui, Kavanayén, La Escalera to Cerro Venamo region, Roraima-tepui). Anzoátegui, Mérida, Táchira, Trujillo; Colombia. ◆Fig. 271.

Cleistes unifoliata (C. Schweinf.) Carnevali & I. Ramírez, Ann. Missouri Bot. Gard. 77: 551. 1990. —Pogonia unifoliata C. Schweinf., Fieldiana, Bot. 28: 171, fig. 27. 1961. Terrestrial or humicolous, to 20 cm tall; leaf solitary, borne in the distal 1/3 of the stem; inflorescences 1-flowered; flowers not widely spreading, pale yellow or greenish yellow; sepals 14–15 mm long. Tepui summits, 900–2200 m; Bolívar (Amaruay-tepui, Cerro Guanacoco, Macizo del Chimantá, Ptari-tepui). Suriname.

Fig. 270. Cleistes rosea Fig. 269. Cleistes huberi

Fig. 271. Cleistes triflora

Clowesia 281

28. CLOWESIA Lindl., Edwards’s Bot. Reg. 29: misc. 25. 1843. [Subtribe Catasetinae]. by Gustavo A. Romero-González Epiphytes. Stems modified into pseudobulbs of several internodes, ovoid to slender-ovoid, when young entirely concealed by distichous, scarious sheaths, eventually naked. Leaves several, distichous, convolute, plicate, articulate, deciduous at the end of the growing season. Inflorescences 1–several, lateral, arising from the base of the pseudobulbs, arching or pendent, racemose, few- to many-flowered. Flowers small to medium-sized, resupinate, bisexual, protandric, fragrant, often showy; floral bracts lanceolate, funnel-shaped. Sepals and petals membranous; lip fleshy, shallowly to deeply concave, entire or obscurely 3-lobed, sometimes extending rearward to 1/2 the length of the pedicellate ovary, the margins entire or sometimes fimbriate. Column short to somewhat elongate, semiterete, clavate, winged, with a conspicuous foot; anther somewhat ventral to terminal, operculate, imperfectly bilocular, membranous, apiculate or not; pollinarium with 2 yellow pollinia, ovate and dorsoventrally flattened to subspherical, cartilaginous, with a tubular or complexly folded tegula, and an adhesive, elliptic or lunate viscidium. Southern Mexico, Central America, Colombia, Venezuela, Ecuador, Peru, Brazil; 6 species, 1 in Venezuela. Clowesia warczewiczii (Lindl. & Paxton) Dodson, Selbyana 1: 136. 1975, “warczewitzii.” —Catasetum warczewiczii Lindl. & Paxton, Paxt. Fl. Gard. 1: 45. 1853 [1850]. Catasetum scurra Rchb. f., Gard. Chron. 2: 1003. 1872. Epiphyte; inflorescence pendent; flowers white to greenish white striped with pale green; column short, without antennae. Wet forests, 50–500 m; Bolívar (Río Paragua), Amazonas (Río Cataniapo, Río Sipapo). Honduras, Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil (Amazonas). ◆Fig. 272.

Fig. 272. Clowesia warczewiczii

282

O RCHIDACEAE

29. COCHLEANTHES Raf., Fl. Tellur. 4: 45. 1836 [1838]. [Subtribe Zygopetalinae]. by Gustavo A. Romero-González Cespitose epiphytes or rarely terrestrial herbs. Stem abbreviate, not modified into pseudobulbs. Leaves numerous, distichous, in a flabelliform cluster, membranous, narrowly ovate-linear to obovate, acute to acuminate, plicate, articulate. Inflorescences lateral, arising from the axil of the leaf sheaths, erect to arching, 1-flowered, shorter than to almost as long as the leaves. Flowers rather fleshy, resupinate, showy; floral bracts concave, appressed, ovate-triangular to tubular-funnel-shaped, shorter than to almost as long as the pedicellate ovary. Sepals and petals spreading to erect-spreading; sepals narrowly linear-ovate to obovate, concave, the lateral ones sometimes slightly oblique and/or the inner basal margins strongly inrolled; petals oblong, elliptic, or obovate, acute, sometimes slightly oblique and/or undulate. Lip sessile to shortly clawed, articulate with the column foot, circular, ovate, quadraterhombic or subquadrate, slightly 3-lobed, the margin most often undulate; disk with a conspicuous, antrorse, thickened, radially keeled callus. Column elongate, slightly arched, semiterete or sometimes subpentagonal in cross section, apically wingless or with 2 short wings, abaxially shallowly concave, produced into a short foot, sometimes abaxially and/or basally pubescent; rostellum filiform, acuminate; anther terminal; pollinia 4, yellow, cartilaginous, dorsoventrally flattened, subequal, obovate, attached in 2 pairs to a trullate to subtriangular tegula. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 14 species, 3 in Venezuela, 1 of these in the flora area. Cochleanthes flabelliformis (Sw.) R.E. Schult. & Garay, Bot. Mus. Leafl. 18: 324. 1959. —Epidendrum flabelliformis Sw., Prodr. 123. 1788. Epiphyte without pseudobulbs; inflorescence 1-flowered; sepals and petals pale green-white; lip slightly 3-lobed, white with purple veins and a fan-shaped callus near the base. Wet forests, ca. 600 m; Bolívar (Altiplanicie de Nuria). Coastal Cordillera of Miranda, Monagas; West Indies, Colombia, Trinidad-Tobago, Ecuador, Brazil (Minas Gerais, Paraná, São Paulo). ◆Fig. 273.

Fig. 273. Cochleanthes flabelliformis

Comparettia 283

30. COMPARETTIA Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 42, t. 73. 1836. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Twig epiphytes. Rhizomes thin, short; pseudobulbs heteroblastic (one internode), small, 1-foliate, clothed by 2 or more distichous sheaths, these frequently with foliar blades. Leaves coriaceous, articulate, narrow-oblong to elliptic. Inflorescences lateral, originating from pseudobulb base, racemose or with few branches, few- to many-flowered, erect to arching. Flower with a long sepal spur; floral bracts inconspicuous; pedicellate ovary terete, longer than sepals. Dorsal sepal free, protuberant at the base with an elongate spur; lateral sepals fused into a cylindric spur; petals similar to dorsal sepal but wider. Lip continuous with the base of the column, spreading, longer than perianth segments, clawed, dilated into a 2-lobed or subpandurate blade, basally with 2 tails or elongated linear spurs, included in the spur; lateral lobes auriculate, middle lobe larger than the others, emarginate; disk in basal zone callose or carinate. Column erect, wide, semiterete, apically grooved, wingless, footless; anther terminal, operculate, incumbent, 1-locular; clinandrium inconspicuous; pollinia 2, cartilaginous, broadly ovoid; rostellum transverse; stigma ventral. Capsule oblong, elliptic to ovoid, obtusely trigonous. Southern Mexico, Central America, West Indies, Colombia, Venezuela, French Guiana, Ecuador, Peru, Brazil, Bolivia; 8 species, 1 in Venezuela.

Fig. 274. Comparettia falcata

284

O RCHIDACEAE

Comparettia falcata Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 42. 1836. Twig epiphyte; flowers pink to purple-red; lip ca. 1 cm broad. Rain and cloud forests, 1000–2000 m; Bolívar (La Escalera to Cerro

Venamo region), Amazonas (Sierra de la Neblina). Aragua, Carabobo, Distrito Federal, Falcón, Lara, Mérida, Miranda, Portuguesa, Yaracuy; widespread in Neotropics. ◆Fig. 274.

31. CORYANTHES Hook., Bot. Mag. 58: t. 3102. 1831. [Subtribe Stanhopeinae]. by Gustavo A. Romero-González Cespitose, robust epiphytes, usually in ant gardens. Stem modified into ovoid, sulcate pseudobulbs of 1 internode. Leaves 2 or 3, narrowly to broadly ovate, acute to acuminate, plicate, articulate. Inflorescences 1–several, lateral, arising from the base of the pseudobulbs, erect, arching to pendent, racemose, few-flowered. Flowers fleshy, waxy, morphologically complex, resupinate, short-lived (2 or 3 days in species from flora area), often showy; floral bracts concave, narrowly to broadly ovate, shorter than the pedicellate ovary. Sepals and petals membranous; sepals spreading or strongly reflexed, the margins undulate, the lateral sepals much larger than the dorsal one; petals much smaller than the sepals, oblong, falcate, the margins undulate. Lip fleshy, rigid, conspicuously clawed, divided into a spreading, disciform, concave or helmet-shaped hypochile, a semitubular mesochile, with or without transverse ridges, and a much larger epichile, cup-shaped, slightly but abruptly 3-lobed in front. Column terete, generally dilated and recurved near the apex, apically winged, footless, basally with 2 shallowly falcate fluid-producing glands; anther terminal, operculate, incumbent, 1-locular; pollinarium with 2 yellow pollinia, obovate, cleft, cartilaginous, and dorsoventrally flattened, the tegula short, ligulate, recurved, the viscidium crescent-shaped. Southern Mexico, Central America, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 20 species, 9 in Venezuela, 8 of these in the flora area. Key to the Species of Coryanthes 1. 1. 2(1).

2.

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

Outer surface of the mesochile smooth, without transverse lamellae ..... 2 Outer surface of the mesochile with transverse lamellae ........................ 3 Margin of the hypochile with a pointy process; projection of the mesochile (this seen only after removing the hypochile) emarginate to deeply bifurcate .................................................................................. C. albertinae Margin of the hypochile shallowly and broadly emarginate; projection of the mesochile (this seen only after removing the hypochile) entire, lanceolate ................................................................................. C. maculata Inflorescence erect; flowers small (petals < 3 cm long) ............................ 4 Inflorescence pendent; flowers large (petals > 5 cm long) ........................ 5 Hypochile deeply emarginate, its concavity facing the claw ........... C. pegiae Hypochile entire, its concavity facing the mesochile ...................... C. rutkisii Hypochile concealing ≥ 2/3 of the mesochile ............................................... 6 Hypochile concealing ≤ 1/2 of the mesochile ............................................... 7 Hypochile concealing 2/3 of the mesochile; projection of the mesochile (this seen only after removing the hypochile) 3, tooth-like ... C. cataniapoënsis

Coryanthes 285

6.

7(5). 7.

Hypochile completely concealing the mesochile; projection of the mesochile (this seen only after removing the hypochile) 2, ridge-like ..................................................................................................... C. gomezii Hypochile shallowly concave, almost flat on lateral view ........... C. feildingii Hypochile deeply concave, helmet-like ..................................... C. macrantha

Coryanthes albertinae H. Karst., Auswahl Gew. Venez. 5, t. 1. 1848. Epiphyte; inflorescence 40–50 cm long, pendent, 3–6-flowered; flowers pink, whitish yellow, or orange, lightly spotted with maroon; dorsal sepal to 3.5 cm long. Wet forests, 50–500 m; Bolívar (Gran Sabana according to Dunsterville), Amazonas (Río Cataniapo, upper Río Orinoco). Coastal Cordillera, Barinas; Mexico, Central America, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil.

Fig. 275. Coryanthes gomezii

Coryanthes cataniapoënsis G.A. Romero & Carnevali, Ann. Missouri Bot. Gard. 76: 454. 1989. Epiphyte; inflorescence (18–)20–25 cm long, pendent, 1- or 2-flowered; flowers white, heavily spotted with maroon; dorsal sepal 5–6 cm long. Wet forests, ca. 100 m; Amazonas (Río Cataniapo). Endemic. ◆Fig. 276. Coryanthes feildingii Lindl., J. Hort. Soc. London 3: 15. 1848. Epiphyte; inflorescence 25–30 long, recurved to pendent, generally 1-flowered; flower whitish yellow, heavily spotted with maroon; dorsal sepal 5–6 cm long. Wet forests, 400–500 m; Bolívar (Río Icabarú, upper Río Caura). Guyana. Coryanthes gomezii G.A. Romero & G. Gerlach, Harvard Pap. Bot. 5: 180. 2000. Epiphyte; inflorescence 20 cm long, pendent, 2-flowered; flowers whitish pink, the hypochile wine red; dorsal sepal 3.5 cm long. Wet forests, 100–200 m; Amazonas (Río Cataniapo). Endemic. ◆Fig. 275. Coryanthes gomezii is a rare species known only from the type collection. Coryanthes macrantha (Hook.) Hook., Bot. Mag. 58: sub t. 3102. 1831. —Gongora macrantha Hook., Bot. Misc. 2: 151, t. 80. 1831. Epiphyte; inflorescence 13–18 cm long, pendent, generally 1- or 2-flowered; flowers whitish yellow to dark orange, heavily spot-

Fig. 276. Coryanthes cataniapoënsis

286

O RCHIDACEAE

Fig. 277. Coryanthes macrantha

Fig. 278. Coryanthes rutkisii

Fig. 279. Coryanthes pegiae

Cranichis 287

ted with maroon; dorsal sepal 5–6 cm long. Wet forests, 50–500 m; Bolívar (near Isla Anacoco, Río Caura, between Río Icabarú and Uaiparú), Amazonas (Río Cataniapo). Distrito Federal. Trinidad-Tobago, Guyana, Peru. ◆Fig. 277. Coryanthes maculata Hook., Bot. Mag. 58: t. 3102. 1831. Epiphyte; inflorescence 40–50 cm long, pendent, generally 4–6-flowered; flowers whitish yellow to yellow, with the edge of the epichile pink to rose, the inside surface of the epichile spotted with maroon; dorsal sepal to 4 cm long. Wet forests, 800–1000 m; Bolívar (Río Caruay). Guyana, Suriname, French Guiana.

Coryanthes pegiae G.A. Romero, Selbyana 9: 147. 1986. Epiphyte; inflorescence 22–32 cm long, erect, generally 1- or 2-flowered; flower whitish yellow, heavily spotted with maroon; dorsal sepal 2–2.5 cm long. Wet forests, 50–100 m; Amazonas (Raudal Rabipelado, Río Cataniapo). Endemic. ◆Fig. 279. Coryanthes rutkisii Foldats, Bol. Soc. Venez. Ci. Nat. 28: 231. 1969. Epiphyte; inflorescence 25–30 cm long, erect, 1- or 2-flowered; flowers whitish yellow, spotted with maroon; dorsal sepal 2–2.5 cm long. Wet forests, 1000–1500 m; Bolívar (Luepa). Brazil. ◆Fig. 278.

32. CRANICHIS Sw., Prodr. 120. 1788. [Subtribe Cranichidinae]. Ocampoa A. Rich., Ann. Sci. Nat. Bot. sér. 3, 3: 31. 1845. Cystochilum Barb. Rodr., Gen. Spec. Orchid. 1: 197. 1877. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial or humicolous herbs, rarely subepiphytes, cespitose. Roots fasciculate, fleshy, villous. Leaves thin to subfleshy, commonly basal, rarely cauline, sometimes with spots, often with a distinct petiole, usually present at anthesis. Inflorescence terminal, racemose, usually lax; peduncle with few tubular bracts. Flowers small, not resupinate; perianth segments usually greenish or white, lip often with showy, prominent, reticulate, green or purplish veins; pedicellate ovary elongate; sepals free, thin; petals usually narrower than sepals; lip fleshier than other perianth segments, usually concave or cochleate, often lobed. Column fleshy, short; clinandrium cyathiform; rostellum prominent, acute, erect; stigma relatively large, terminal; anther dorsal, erect; pollinia 4, with a small hamulus, brittle in chunks, with a small, globose viscidium. U.S.A. (Florida), Mexico, Central America, West Indies, Colombia, Venezuela, Suriname, French Guiana, Ecuador, Peru, Amazonian and southern Brazil, Bolivia, northern Argentina; ca. 60 species, 9–11 species in Venezuela, 1 of these in the flora area. Cranichis is a genus in need of a modern revision. Cranichis diphylla Sw., Prodr. 120. 1788. Cranichis monophylla Lindl., Orchid. Linden. 27. 1846. Cranichis guatemalensis Schltr., Repert. Spec. Nov. Regni Veg. 2: 129. 1906. Cranichis nigrescens Schltr., Repert. Spec. Nov. Regni Veg. 10: 482. 1911. Cranichis ovatilabia Schltr., Repert. Spec. Nov. Regni Veg. 7: 59. 1920. Cranichis stictophylla Schltr., Repert. Spec. Nov. Regni Veg. 7: 62. 1920. Cranichis alfredi Schltr., Repert. Spec.

Nov. Regni Veg. 19: 82. 1923. Terrestrial herb 15–20 cm tall; flowers white with green veins; dorsal sepal 1.5–3.5 mm long. Moist forests, 500–1000 m; Bolívar (Río Maurak headwaters near El Paují). Aragua, Distrito Federal, Miranda, Mérida; southern Mexico, Central America, West Indies, Colombia, Suriname, Ecuador, Peru. ◆Fig. 280. Cranichis diphylla is the only representative in the Guayana Highlands of this mainly Andean genus.

288

O RCHIDACEAE

Fig. 280. Cranichis diphylla

33. CRYPTARRHENA R. Br., Bot. Reg. 2: t. 153. 1816. [Subtribe Cryptharrheninae]. Orchidofunckia A. Rich. & Galeotti, Ann. Sci. Nat. Bot. sér 3, 3: 24. 1845. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, sometimes twig epiphytes, small to medium-sized, erect to pendulous, cespitose, shade-loving. Pseudobulbs absent or present, when present, small, flattened, 1- or 2-leaved apically, enveloped by several distichous, imbricate leafbearing sheaths. Leaves conduplicate, articulate with their sheaths, coriaceous to ± fleshy, apically obliquely 2-lobed. Inflorescences lateral, originating from the axils of the leaf sheaths or from pseudobulb base, laxly to ± densely many-flowered racemes, usually longer than leaves, suberect, arching to pendulous. Flowers resupinate, small, widely spreading; floral bracts small, linear; pedicellate ovary subterete. Sepals free, similar, the lateral ones ± oblique; petals similar to dorsal sepal but smaller. Lip spreading, stalked, with a callus on the stalk, expanded into a 4-lobed blade from the keeled claw with linear, ovate-triangular to subquadrate lobes, apically emarginate. Column erect, short, broadened toward apex, wingless, footless; anther terminal to subventral, operculate, incumbent; clinandrium with an erose, wide margin that completely covers the anther; pollinia 4, cartilaginous,

Cryptocentrum 289

caudicles long, narrow, thickened toward apex, viscidium small; rostellum horizontal; stigma ventral. Capsules obovoid-ellipsoid. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 3 or 4 species, 1 in Venezuela. Cryptarrhena kegelii Rchb. f., Bot Zeitung (Berlin) 10: 766. 1852. Epiphyte; racemes short; flowers minute, complex, green; sepals 3–5 mm long. Rain forests, in moist places on moss, 600–800 m; Bolívar (Altiplanicie de Nuria, Auyán-tepui). Miranda, Táchira; Mexico, Central America, West Indies, Guyana, Suriname, Colombia, Ecuador, Peru, Brazil. ◆Fig. 281.

Fig. 281. Cryptarrhena kegelii

34. CRYPTOCENTRUM Benth. & Hook. f., Gen. Pl. 3: 557. 1883. [Subtribe Maxillarinae]. Pittierella Schltr., Repert. Spec. Nov. Regni Veg. 3: 80. 1907. by Germán Carnevali and Ivón M. Ramírez-Morillo Minute to medium-sized epiphytes, cespitose. Rhizome short to almost absent; stems short, not pseudobulbose thickened, cylindric in their basal portion, covered with leaf-bearing sheaths which are distichous, imbricate. Leaves articulate with sheaths, conduplicate to terete, fleshy to coriaceous, distichous or polystichous, crowded on the short stem. Inflorescences originating from sheath axils or from the base of the stem, 1-flowered, pedunculate, few to many simultaneously, usually shorter than the leaves, but much longer than leaves in some species; peduncle covered with tubular sheaths. Flowers small, greenish to yellowish, sometimes purple or reddish-tinged, or totally red-purple; resupinate, fleshy; floral bract tubular, completely covering pedicellate ovary. Sepals connate in the basal portion, free and spreading in the apical; dorsal sepal generally shorter than the laterals; lateral se-

290

O RCHIDACEAE

pals basally produced into an elongate spur, totally hidden by the floral bract; petals similar to the dorsal sepal or shorter, free. Lip decurrent on column, basally produced into an elongate spur which is included in and is a little shorter than the sepal spur; blade of lip simple, usually lanceolate to elliptic-lanceolate. Column footless, short, usually with a pair of auricles apically; anther apical, operculate, incumbent, unilocular; pollinia 4, with a small viscidium; stigma ventral. Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Ecuador, Peru, Bolivia; 17 species, 2 in Venezuela, both in the flora area. Key to the Species of Cryptocentrum 1.

1.

Leaves distichous on the stem, 5–15(–20) cm long, always much longer than the stem; floral bract long and enveloping nearly all the spur ..................................................................................... C. dunstervilleorum Leaves spiralled on the stem, 1.5–2.5 cm long, shorter or ± equal to the stem in mature plants; floral bract short and only enveloping the apex of the spur ................................................................................ C. peruvianum

Cryptocentrum dunstervilleorum Carnevali & G.A. Romero, Orchidee (Hamburg), Beih. 1: 3. 1994. Cryptocentrum gracillimum auct. non Ames & C. Schweinf. 1925: sensu Dunst. & Garay, Venez. Orchid. Ill. 2: 79. 1961; Foldats in Lasser, Fl. Venez. 15(5): 442. 1970. Epiphyte, almost always growing buried in bryophytes. Inflorescences usually several

simultaneously, long-pedunculate; flowers yellow-green. Cloud forests, 600–1400 m; locally common in eastern Bolívar (Altiplanicie de Nuria, La Escalera to Cerro Venamo region, headwaters of Río Chicanán). Guyana. ◆Fig. 282. Cryptocentrum peruvianum (Cogn.) C. Schweinf., Bot. Mus. Leafl. 12: 188. 1946. —Centroglossa peruviana Cogn., Bull.

Fig. 282. Cryptocentrum dunstervilleorum

Cyclopogon 291

Soc. Roy. Bot. Belgique 43: 331. 1906. Colombia, Venezuela, Ecuador, Peru; 3 subspecies, 1 in Venezuela. C. peruvianum subsp. peruvianum Cryptocentrum hoppii Schltr., Repert. Spec. Nov. Regni Veg. 27: 103. 1924.

Epiphyte, usually growing on twigs buried in bryophytes; leaves acute; inflorescences usually few, short-pedunculate; flowers yellow or green, campanulate, petals 3.5– 6 mm long. Rain or cloud forests, 500–1600 m; Amazonas (Caño Piedra on Cerro Sipapo massif). Colombia, Ecuador, Peru.

35. CYCLOPOGON C. Presl, Reliq. Haenk. 1: 93. 1827. [Subtribe Spiranthinae]. Beadlea Small, Fl. S.E. U.S. 319. 1903. Spiranthes Rich., De Orchid. Eur. 20, 28, 36. Aug.–Sept. 1817, nom. cons., pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial, slender herbs. Roots fasciculate, fusiform. Leaves basal, not articulate, rosulate, commonly pseudopetiolate, concolorous or variegated. Inflorescence erect, variously bracteate, terminated by a loose- to dense-flowered spike. Flowers resupinate; floral bracts conspicuous; pedicellate ovary cylindric to fusiform, slightly twisted, sessile. Sepals free, subparallel, globose at the base; lateral sepals oblique, with base of column forming a short mentum; petals connivent with dorsal sepal, clawed, narrower than sepals. Lip clawed, sagittate-auriculate or rarely cordate at base, unlobed to 3-lobed, fleshier than other perianth segments, lateral margins joined with sides of column. Column erect, free from dorsal sepal, ± elongate with a short, oblique base at top of ovary; anther concave-hooded; pollinia 4, puberulent, clavate, with a small viscidium originated on the abaxial surface of the rostellum; rostellum undivided, soft, pliable, longer than wide, linear-oblong to variously triangular; stigmas 2, free to joined. U.S.A. (Florida), Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, southern Brazil, Bolivia, Paraguay, northern Argentina, Uruguay; ca. 55 species, 5 or 6 in Venezuela, 2 of these in the flora area. The authors have seen live, sterile material collected near the base of Cerro Marahuaka of a third, as yet unidentified, species. Key to the Species of Cyclopogon 1. 1.

Lip broadest below apical constriction; leaves velvety purple-green above; pseudopetiole often shorter than leaf ................................ C. cranichoides Lip broadest above apical constriction; leaves green above; pseudopetiole often longer than leaf ..................................................................... C. elatus

Cyclopogon cranichoides (Griseb.) Schltr., Beih. Bot. Centralbl. 37(2): 387. 1920. —Pelexia cranichoides Griseb., Cat. Pl. Cub. 29. 1866. —Beadlea cranichoides (Griseb.) Small, Fl. S.E. U.S. 319. 1903. —Spiranthes cranichoides (Griseb.) Cogn. in Urb., Symb. Antill. 6: 338. 1909. Terrestrial or subterrestrial; roots tuberous; leaves basal, ca. 5, with 5 lines of dull green dots between veins, dull green-magenta below; inflorescences greenish, yellow-

ish, or purplish, basally glabrous, apically pubescent, many-flowered; dorsal sepal maroon-brown; lip white. Lower montane forests, 300–700 m; Bolívar (Altiplanicie de Nuria). U.S.A. (Florida), Mexico, Central America, West Indies, Colombia, Ecuador. ◆Fig. 283. Cyclopogon elatus (Sw.) Schltr., Beih. Bot. Centralbl. 37(2): 387. 1920. —Satyrium elatum Sw., Prodr. 119. 1788. —Spiranthes elata (Sw.) Rich., De Orchid. Eur. 37.

292

O RCHIDACEAE

1817. —Beadlea elata (Sw.) Small ex Britton, Brooklyn Bot. Gard. Mem. 1: 38. 1918. Erect, cespitose, humicolous to terrestrial; flowers small, not opening widely; sepals externally greenish or brownish; lip white. Lower montane forests, 500–600 m; Bolívar (Altiplanicie de Nuria). Monagas; U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, southern Brazil, Paraguay, northern Argentina, Uruguay. Cyclopogon elatus is broadly defined and may include several different taxa.

Flower

Lip

Fig. 283. Cyclopogon cranichoides

Cycnoches 293

36. CYCNOCHES Lindl., Gen. Sp. Orchid. Pl. 154. 1832. [Subtribe Catasetinae]. Cycnauken Lem., Fl. Serres Jard. Eur. 1: 87. 1845. by Gustavo A. Romero-González Epiphytes, often on rotten tree trunks, or rarely terrestrial. Stems modified into pseudobulbs of several internodes, slender, elongate, fusiform-cylindric, when young entirely concealed by distichous, scarious sheaths, eventually naked. Leaves several, distichous, convolute, plicate, articulate, deciduous at the end of the growing season. Inflorescences 1–several, lateral, arising from the base of the apical internodes, arching or pendent, few- to many-flowered. Flowers nonresupinate, unisexual, staminate and pistillate ones often sexually dimorphic, rarely also with polymorphic intermediate ones; floral bracts lanceolate, funnel-shaped. Staminate flowers whitish green or various colors, spotted or not; sepals and petals membranous, the lateral sepals strongly reflexed or not; lip sessile or clawed, the margin with finger-like, clavate processes; disk ventricose, fleshy-thickened, or sometimes thin, lightly concave; column elongate, arched, terete, slender-clavate, with a conspicuous foot; anther somewhat ventral, operculate, unilocular, membranous; pollinarium with 2 yellow pollinia, cartilaginous, spherical, with a semitubular tegula, and a large, adhesive, circular or elliptic viscidium. Pistillate flowers whitish green or green; sepals and petals fleshy, strongly reflexed; lip ventricose, fleshythickened; column short and thick-clavate, with a conspicuous foot; pollinarium and anther poorly developed, deciduous at anthesis; stigma a narrow transverse cleft near the apex. Intermediate flowers mostly sterile, color and morphology a continuum between the pistillate and the staminate flowers. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 20–25 species, 5 in Venezuela, all in the flora area. Plants in this genus can produce pistillate, staminate, or rarely intermediate flowers, sometimes mixed in the same raceme. As pistillate flowers are sometimes indistinguishable between species, only the characters of staminate flowers (those presenting elongate, slender columns with fully developed pollinaria) are considered in this key and in the species descriptions. Key to the Species of Cycnoches 1. 1. 2(1). 2. 3(1). 3. 4(3). 4.

Flowers small; dorsal sepal 1.5–4 cm long ................................................ 2 Flowers large; dorsal sepal 6–11 cm long .................................................. 3 Flowers yellowish green and white with few maroon spots; lip fleshy, the margins entire ............................................................................... C. haagii Flowers purple or purple-spotted throughout; lip membranous, with marginal processes .................................................................. C. thurstonorum Flowers brownish green, with purple spots; lip oblong-lanceolate to lanceolate, without a prominent callus ........................................ C. loddigesii Flowers yellowish green and white; lip ovate to ovate-lanceolate, with a prominent dark green callus ................................................................. 4 Callus of the lip dark greeen, dorsal sepal > 7 cm ................. C. chlorochilon Callus of the lip yellowish greeen, dorsal sepal ≤ 6.5 cm ................. C. lusiae

294

O RCHIDACEAE

Cycnoches chlorochilon Klotzsch, Allg. Gartenzeitung 6: 225. 1838. —Cycnoches ventricosum var. chlorochilon (Klotzsch) P. Allen, Orchid J. 1: 401. 1952. Epiphyte; inflorescence pendent; flowers white to greenish white, nonresupinate; lip ventricose, greenish white to ivory, with a prominent, green, basal callus; column pale yellowish green. Wet forests, 50–100 m; Bolívar (Río Cuchivero). Coastal Cordillera, Yaracuy, Zulia; Panama, Colombia, Guyana.

Several male specimens of euglossine bees, Eulaema cingulata Fabr., captured in Río Cataniapo, Amazonas state, carried pollinaria referable to this species. Cycnoches haagii Barb. Rodr., Gen. Spec. Orchid. 2: 221. 1882. Cycnoches versicolor Rchb. f., Gard. Chron. ser. 3, 4: 596. 1888. Epiphyte; inflorescence arching or pendent; flowers greenish white (yellowish green

Fig. 284. Cycnoches haagii

Fig. 285. Cycnoches thurstonorum

Cycnoches 295

Fig. 286. Cycnoches lusiae

when wilting), nonresupinate; lip fleshy, irregularly spotted with maroon, the margins entire; column terete, elongate. Wet forests, 50–1000 m; Bolívar (Kavanayén, Río Paragua), Amazonas (Río Cataniapo, Río Padamo). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, (Amazonas, Pará, Goiás, Matto Grosso). ◆Fig. 284. Cycnoches loddigesii Lindl., Gen. Sp. Orchid. Pl. 154. 1832. Cycnoches cucullata Lindl., Edwards’s Bot. Reg. 23: sub t. 1951. 1837. Epiphyte; inflorescence arching to pendent; flowers greenish brown, nonresupinate; lip fleshy, narrowly oblong-lanceolate, acute, white to pale pink or lightly spotted with maroon, the apex often with greenish maroon veining; column dark maroon. Wet forests,

200–1000 m; Bolívar (Altiplanicie de Nuria, Cerro Muribata, Río Caura, Salto Pará). Sucre; Guyana, Suriname, French Guiana, Brazil (Amazonas). Cycnoches lusiae G. Romero & Garay, Harvard. Pap. Bot. 476. 1999. Epiphyte; inflorescence pendent; flowers white to greenish white, nonresupinate; lip ventricose, greenish white to ivory, with a prominent, yellowish green, basal callus; column pale yellowish green. Wet forests; Bolívar (Distrito Cedeño). Endemic. ◆Fig. 286. Cycnoches thurstonorum Dodson, Icon. Pl. Trop. ser. 2, 5: t. 432. 1989. Epiphyte; inflorescence pendent; flowers white to greenish white, nonresupinate; lip white, spotted with maroon, lanceolate,

296

O RCHIDACEAE

acuminate, the margins with clavate, fingerlike processes; column dark red-maroon. Wet forests, ca. 100 m; Amazonas (San Carlos de Río Negro). Colombia, Ecuador. ◆Fig. 285.

The identification of this taxon as Cycnoches thurstonorum should be regarded as tentative until more material is collected to make a definitive determination.

37. CYRTOPODIUM R. Br. in W.T. Aiton, Hortus Kew. ed. 2, 5: 216. 1813. [Subtribe Cyrtopodiinae]. by Gustavo A. Romero-González Mostly terrestrial or lithophytic herbs, sometimes epiphytic herbs outside the flora area, cespitose. Stem modified into pseudobulbs, ovoid or fusiform to cylindric, of several elongate internodes, when young entirely concealed by distichous, scarious sheaths, eventually naked. Leaves several, distichous, coriaceous, ovate to obovate, acuminate, plicate, articulate, deciduous at the end of the growing season. Inflorescences lateral, arising from the base of the pseudobulbs, erect, racemose to paniculate, many-flowered, usually much longer than leaves, in the flora area most often arising with the new, developing growth. Flowers resupinate, often showy; floral bracts linear-ovate to elliptic, often conspicuous, showy, the margins undulate. Sepals and petals membranous, spreading, often conspicuously undulate; sepals narrowly elliptic or ovate to obovate, the lateral ones laterally adnate to the base of the column foot, sometimes wider than the dorsal one and/or slightly oblique; petals usually wider than the sepals, obovate to elliptic-obovate. Lip subsessile to clawed, adnate to the apex of the column foot, basally cordate to cuneate, conspicuously 3lobed, the lateral lobes erect to spreading, subcircular to narrowly elliptic, sometimes slightly falcate, midlobe ovate, kidney-shaped, semicircular, to tranversely elliptic, sometimes spatulate, apically truncate, sometimes shallowly emarginate or 2-lobed, the margins entire, undulate and/or verrucose; disk with a keeled, tuberculate, cristate, or verrucose callus. Column short to somewhat elongate, slightly arched, clavate to subclavate, semiterete, with a conspicuous foot; anther terminal, incumbent, operculate, imperfectly 2-locular; pollinia 2, yellow, cartilaginous, sulcate, subtriangular in outline, semiterete in cross section, attached to a short, trullate viscidium. U.S.A. (Florida), Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina; 30 species, 7 in Venezuela, 5 of these in the flora area. Key to the Species of Cyrtopodium 1. 1. 2(1). 2. 3(2). 3. 4(3). 4.

Lip midlobe margin conspicuously verruculose-tuberculate ... C. graniticum Lip midlobe margin not conspicuously verruculose-tuberculate ............. 2 Mature, flowering pseudobulbs large, > 25 cm long; lip lateral lobes bright yellow ..................................................................................... C. andersonii Mature, flowering pseudobulbs small, to 15–20 cm long; lip lateral lobes spotted or solid dark orange or maroon ................................................ 3 Lip base cuneate, the midlobe conspicuously clawed ............. C. parviflorum Lip base subcordate to deeply cordate, the midlobe not clawed .............. 4 Flowers yellow with orange-green markings; lip 14–20 mm wide between lateral lobes, the base subcordate ........................................... C. cristatum Flowers yellow with purple-brown spots; lip 9–13 mm wide between lateral lobes, the base deeply cordate ...................................................... C. fowliei

Cyrtopodium 297

Cyrtopodium andersonii (Lamb. ex Andrews) R. Br. in W.T. Aiton, Hortus Kew ed. 2, 5: 216. 1813. —Cymbidium andersonii Lamb. ex Andrews, Bot. Repos. 10: t. 651. 1812. Cyrtopodium flavescens Cogn., J. Orchidées 6: 74. 1895. Inflorescence paniculate; flowers bright yellow. Sandy, rocky savannas, 300–1500 m; Delta Amacuro (Serranía de Imataca), Bolívar (between El Pao and San Félix, Gran Sabana, Río Caura), Amazonas (Río Yureba). Guyana, Brazil. Cyrtopodium cristatum Lindl., Edwards’s Bot. Reg. 27: sub t. 8. 1841. Inflorescence racemose (rarely paniculate); flowers bright yellow with dark orange and light green markings. Sandy savannas, 100–1800 m; Bolívar (Gran Sabana), Amazonas (upper Río Ventuari). Monagas; Colombia, Guyana, Brazil (Bahia, Minas Gerais, Pará). ◆Fig. 287. Cyrtopodium fowliei L.C. Menezes, Orchid Digest 59: 17. 1995. Inflorescence racemose; sepals and petals bright yellow with maroon markings, lip orange with yellow markings. Sandy savannas, 800–900 m; Bolívar (San Ignacio). Guyana, Suriname, Brazil. A color photograph of Cyrtopodium fowliei was published in Amer. Orchid Soc. Bull. 60: 445. 1991. Cyrtopodium graniticum G.A. Romero & Carnevali, Harvard Pap. Bot. 2: 512. 1999. Inflorescence paniculate; flowers bright yellow with maroon and green markings. Granitic outcrops, 50–100 m; Amazonas (Piedra Cocuy, Puerto Ayacucho). Endemic. Collections from the flora area previously identified as Cyrtopodium punctatum (L.) Lindl. are here referred to C. graniticum. In C. punctatum, the lateral lobes of the lip are longer than the midlobe and the apex of the midlobe is truncate; in C. graniticum, the lateral lobes of the lip are shorter than the midlobe and the apex of the midlobe is obtuse. Cyrtopodium parviflorum Lindl., London J. Bot. 2: 672. 1843. Cyrtopodium broadwayi Ames, Orchidaceae 7: 51, t. 112. 1922.

Fig. 287. Cyrtopodium cristatum

298

O RCHIDACEAE

Inflorescence racemose (rarely paniculate); flowers bright yellow with deep maroon and green markings. Sandy savannas, 200– 1800 m; Bolívar (Altiplanicie de Nuria,

Auyán-tepui, near Kavanayén, Río El Toro, Roraima-tepui, near Santa Elena de Uairén). Trinidad, Guyana, Suriname, French Guiana, Brazil (Amazonas, Goiás, Mato Grosso).

38. DIADENIUM Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 41, t. 71. 1835. [Subtribe Oncidiinae]. Chaenanthe Lindl., Edwards’s Bot. Reg. 24: misc. 38. 1838. by Germán Carnevali and Ivón M. Ramírez-Morillo Minute to small twig epiphytes, cespitose, in some species the whole plant purple-tinged. Rhizome short, thin; pseudobulbs orbicular, ovoid to fusiform, apically 1-foliate, totally or partially enveloped by sheaths of which the innermost 1– 4 are leaf-bearing. Leaves conduplicate, thinly coriaceous, elliptic to oblong-elliptic, acute. Inflorescences lateral, originating from pseudobulb base, racemose or 2- or 3pinnate panicles, few- to many-flowered; peduncle and rachis thin, articulations covered by small sheaths. Flowers small, resupinate or not, successive or several in simultaneous anthesis in each branch, pink to purple; floral bracts small, acute; pedicellate ovary terete, shorter to longer than sepal spur. Dorsal sepal free, elliptic to oblong-elliptic; lateral sepals long-decurrent on the elongate column foot, the free portion connate into an apically bifid to retuse synsepal, which is oblong to oblongelliptic, the basal part sometimes forming a mentum or a ± elongate spur which contains the apical portion of the column foot and the basal portion of the lip; petals similar to dorsal sepal or broader. Lip divided into a basal narrow claw that runs parallel to the column foot and is sometimes enclosed by the sepal spur, and a dilated apical blade; blade unlobed or basally lobed, apex obtuse, truncate to 2-lobed. Column ± cylindrical, basally produced into an elongate foot; anther apical, incumbent, operculate; pollinia 2, ovoid to ellipsoid, tegula elongate, viscidium short, narrow; stigma ventral, elliptic to slit-like. Venezuela, Ecuador, Peru, Brazil, Bolivia; 2 species, 1 in Venezuela. Diadenium barkeri (Lindl.) Benth. & Hook. f., Gen. Pl. 3: 558. 1880. —Chaenanthe barkeri Lindl., Edwards’s Bot. Reg. 24: misc. 38. 1838. Herb 5–7 cm tall, frequently on small shrubs, often purple-tinged; flowers small, pinkish or violet-pink. Open places in rain forests, 500–900 m; rare and local in Bolívar (near Perai-tepui and Icabarú). Ecuador, Peru, Amazonian Brazil, Bolivia. ◆Fig. 288.

Fig. 288. Diadenium barkeri

Dichaea 299

39. DICHAEA Lindl., Gen. Sp. Orchid. Pl. 208. 1833. [Subtribe Zygopetalinae]. Fernandezia Ruiz & Pav., Fl. Peruv. Prodr. 123. 1794, pro parte. Epithecia Knowles & Westc., Fl. Cab. 2: 167, t. 87. 1838. Dichaeopsis Pfitzer in Engl. & Prantl, Nat. Pflanzenfam. 2(6): 207. 1889. by Germán Carnevali and Ivón M. Ramírez-Morillo Generally epiphytes, rarely lithophytes or subterrestrial herbs, minute to rather large, cespitose, pendulous, arching to erect or prostrate. Roots thin, sometimes very thick and fleshy, emerging basally or from the internodes in cases of prostrate stems. Stems monopodial or pseudomonopodial with indeterminate growth but rarely getting beyond a certain length for each particular species, totally enveloped by leaf sheaths, frequently branching basally or, more rarely, from distal internodes. Leaves distichous, thin to succulent, marcescent (remaining on the plants and eventually disintegrating there) or articulate with their sheaths; sheaths imbricate, blades usually oblong or linear-oblong, sometimes elliptic or narrowly ovate-elliptic, rounded to acute or acuminate, frequently apiculate or subapically mucronate, margins smooth or ciliolate to finely dentate, the cilia or teeth sometimes restricted to the apical part of the leaves. Inflorescences 1-flowered, solitary, opposite the leaves; peduncle usually filiform, mostly shorter than leaves; floral bracts usually enveloping 1/2 or more of the ovary, tubular, frequently apiculate or mucronate; ovary obconical to fusiform, surface smooth to densely muricate, usually with a linear bracteole basally, longer to shorter than the ovary. Flowers resupinate, usually campanulate or with spreading perianth segments, short-lived. Perianth segments membranous to fleshy, 3–20 mm long, 1–5-veined, oblong-elliptic to ovate, rounded to acuminate, the external surface varying from smooth to verruculose, usually white or greenish, frequently tinged, spotted, or blotched with red, purple, pink, or violet; lateral sepals generally somewhat oblique. Lip rigidly attached to column base, usually attenuated at base, blade usually anchor-shaped, rarely pandurate, margins smooth or finely ciliolate, color as in other perianth segments but usually with larger zones of pink, red, purple, or violet, generally ecallose. Column short to rarely somewhat elongate, subcylindric, forming an angle with the ovary; anther terminal, operculate, incumbent; clinandrium shallow; pollinia 4, ovoid, waxy, tegula well developed, short to elongate, viscidium small; stigma ventral but sometimes appearing as frontal due to angle in column, rather large, frequently with an infrastigmatic, often pubescent ligule. Capsules smooth to muricate. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 70 species, 20–25 in Venezuela, 15 of these in the flora area. Understanding the systematics of Dichaea has been difficult because the flowers do not rehydrate easily, making them hard to interpret from herbarium specimens. Furthermore, the plants are usually found with only one or few flowers at a time and these are often lost during the drying process. The recent emphasis on field work and the collection of new material has improved our understanding of Dichaea during the last two decades, coupled with an increase of known, better documented, recently described taxa.

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

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

4.

5(3).

5.

6(5).

6.

7(1). 7. 8(7).

Leaves not articulate with their sheaths, marcescent; ovary always muricate ................................................................................................. 2 Leaves articulate with their sheaths, eventually deciduous; ovary smooth or muricate ............................................................................................. 7 Plants small, usually < 10 cm long; leaves ciliate-denticulate throughout, 2.5–3 mm wide; claw of the lip with a 2-lobed callus basally ................ ................................................................................................. D. hystricina Plants usually > 15 cm long; leaves ciliate-denticulate only in the apical 1/2, mature leaves > 5 mm wide; claw of lip ecallose ............................ 3 Mature leaves usually (25–)28–40 mm long, 4–5 times longer than wide, apex acute to acuminate ........................................................................ 4 Mature leaves usually (5–)10–20(–25) mm long, 2–3.5 times longer than wide, apex rounded, obtuse to acute ..................................................... 5 Lip obovate-pandurate (fiddle-shaped with the apex the larger end), not anchor-shaped; mature leaves narrowly ovate-elliptic to narrowly elliptic, acute, the upper surfaces of the leaves all facing the same side of the plant; sepals elliptic obovate, obtuse and minutely apiculate .................................................................................................... D. pendula Lip anchor-shaped; mature leaves narrowly elliptic to triangular-elliptic, acuminate or narrowly acute, the upper surfaces of the leaves all perpendicular to the stem and then facing upward or downward; sepals narrowly elliptic, acute to acuminate ............................. D. camaridioides Leaves acute and shortly acuminate, narrowly ovate to narrowly elliptic; stem internodes somewhat elongate and foliar blades not overlapping, then leaves laxly arranged along the axis of the plant; petals obtuse or broadly acute; infrastigmatic ligule longer than stigmatic surface ................................................................................................ D. tachirensis Leaves abruptly short-acuminate to mucronate, broadly elliptic or oblong-elliptic to oblong; stem internodes abbreviated and then foliar blades overlapping, thus leaves densely arranged almost along the whole length of the plant; petals obtuse to acute and acuminate; infrastigmatic ligule shorter than stigmatic surface ........................... 6 Leaves with reticulate cross veins, usually narrowly oblong-elliptic to oblong; sepals glabrous without; petals obtuse to obtusely acute; claw of the lip narrow, its margins and the basal margins of the lip blade glabrous ................................................................................. D. splitgerberi Leaves without reticulate cross veins, usually broadly elliptic; sepals verruculose without; petals acute; claw of the lip broad, its margins and those of the basal portion of the lip blade minutely ciliate .................................................................................................... D. latifolia Ovary muricate .......................................................................................... 8 Ovary smooth ........................................................................................... 10 Leaves 40–75 × 12 mm; sepals 7–15 mm long ............................... D. robusta

Dichaea 301

8. 9(8).

9.

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

13.

14(12). 14.

Leaves 13–17 × 3–7 mm; sepals 4.8–5 mm long ....................................... 9 Leaves 15–25 × 4–7 mm; 3–4 times longer than wide, apex obtuse or rounded, abruptly aristate; lip 4.2–4.2 mm long, 4.5 mm wide across the expanded lateral lobes ............................................................ D. kegelii Leaves 25–35 × 3–4 mm, at least 6 times longer than wide, apex acute or acuminate, long aristate; lip 4–4.2 mm long, 5–5.5 mm across the expanded lateral lobes .......................................................... D. venezuelensis Leaves (3.5–)4.5–10 cm long; infrastigmatic ligule at least 1/2 the length of the column .................................................................................. D. trulla Leaves < 3 cm long; infrastigmatic ligule < 1/3 the length of the column .............................................................................................................. 11 Petals conspicuously wider than lateral sepals; leaves ± glaucous when fresh ...................................................................................... D. panamensis Petals ± equal in width to lateral sepals; leaves not glaucous when fresh .............................................................................................................. 12 Lip apically rounded, truncate, or rarely obtuse .................................... 13 Lip apically acute ..................................................................................... 14 Leaves fleshy, the lower surface with veins hardly noticeable; plants usually much branching, often growing at river edges, flooded for part of the year; ovary not subtended by a bracteole; lateral veins of the dorsal sepal confluent with the central vein at the apical 1/3; lateral lobes of the lip obtuse, lip apically truncate .................................. D. ancoraelabia Leaves membranous-subfleshy, the lower surface with 2 or 3 veins conspicuous; plants with few branches, usually growing inside terra firme forests; ovary subtended by a small but conspicuous bracteole; lateral veins of the dorsal sepal not confluent with the central vein; lateral lobes of the lip acute; lip apically rounded to broadly obtuse ...... D. hookeri Leaves < 3.5 times longer than wide; petals broadly elliptic, obtuse; infrastigmatic ligule conspicuous ................................................... D. picta Leaves > 5.5 times longer than wide; petals narrowly ovate-elliptic, acute; infrastigmatic ligule absent or inconspicuous ............ D. trinitensis

Dichaea ancoraelabia C. Schweinf, Amer. Orchid Soc. Bull. 16: 614. 1947. Epiphyte 5–50 cm long, pendulous; stem usually unbranched; flowers white with some red spotting. Riparian rain forests, 100–300 m; Amazonas (Raudal Remo on Rio Siapa, basin of Río Casiquiare, Río Cunucunuma, Rio Ocamo). Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Amazonas, Pará, Roraima). Dichaea ancoraelabia is locally common in many places in southern Amazonas growing as a low epiphyte at the edges or rivers, spending part of the rainy season totally or partially submerged.

Dichaea camaridioides Schltr., Repert. Spec. Nov. Regni Veg. Beih. 7: 201. 1920. Dichaea brachypoda auct. non Rchb. f 1866: sensu Foldats in Lasser, Fl. Venez. 15(5): 447. 1970; Dunst. and Garay, Venez. Orchid. Ill. 3: 73. 1965. Epiphyte, cespitose, erect, arched to pendulous; flowers cream-colored, purple-speckled, margins green. Cloud forests on tepui slopes or summits, 1200–1300 m; Amazonas (Cerro Marahuaka, upper Río Yameduaka near Cerro Duida). Lara, Táchira, Yaracuy; Costa Rica, Colombia.

302

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Dichaea hookeri Garay & Sweet, J. Arnold Arbor. 53: 395. 1972. Dichaea rendlei var. trinitensis auct. non (Gleason) Foldats 1959: sensu Dunst. and Garay, Venez. Orchid. Ill. 1: 101. 1959; Foldats in Lasser, Fl. Venez. 15(5): 473. 1970. Epiphyte, cespitose, pendulous to arching, shade-loving; stems unbranched or branching basally, to 15 cm long; flowers small, white or greenish white, sometimes red-spotted, campanulate; sepals 5–8 mm long. Rain forests; 100–500 m; Delta Amacuro (Río Acure), Bolívar (Río Caroní basin, southwest of Urimán), Amazonas (basin of Río Atabapo, Río Negro, Río Ventuari). Miranda, Nueva Esparta; Lesser Antilles, Trinidad-Tobago.

The recently rediscovered Dichaea kegelii is easy to recognize among the species with articulate leaves and muricate ovaries by its short, relatively broad, apically obtuse or rounded leaves. The name D. kegelii has been erroneously applied to D. splitgerberi, a member of subgenus Dichaea (with leaves not articulated to their sheaths). A good illustration of this species is published in Dunsterville and Garay, Venez. Orchid. Ill. 1: 97. 1959.

Dichaea hystricina Rchb. f., Flora 48: 279. 1865. Epiphyte, suberect, spreading to pendulous, often growing among bryophytes, shade-loving; stem unbranched or branched, usually < 8 cm long, rarely to 15 cm; flowers minute, campanulate, white or greenish white with some violet-pink spots, sepals 4–6 mm long; lip sometimes entirely violet. Cloud forests, 600–1700 m; Bolívar (La Escalera to Cerro Venamo region, basin of Río Caroní, Río Cuyuní, Serrania de Maigualida), Amazonas (Cerro Aracamuni, Cerro Marahuaka, Sierra de la Neblina). Widespread in northern Venezuela; Guatemala to Panama, Colombia, Ecuador, Brazil (Amazonas, Roraima). ◆Fig. 290.

D. latifolia var. longa (Schltr.) Folsom, Orch. Dig. 60: 154. 1996. —Dichaea longa Schltr., Repert. Spec. Nov. Regni Veg. Beih. 10: 54. 1922. Dichaea muricata auct. non (Sw.) Lindl. 1833: sensu Dunst. and Garay, Venez. Orchid. Ill. 2: 89. 1961; Foldats in Lasser, Fl. Venez. 15(5): 460. 1970. Epiphyte, pendulous, shade-loving; stems to 1 m long, unbranched or branching in the lower 1/2; flowers campanulate; sepals greenish, sometimes violet-spotted; sepals 7–9 mm long; lip white with violet zones. Rain to cloud forests, 500–1500 m; Bolívar (Cerro Marutaní, Sierra Pakaraima, Urimán). Coastal Cordillera, Portuguesa; Colombia, Ecuador, Peru, Bolivia.

Dichaea kegelii Rchb. f., Linnaea 41: 129. 1877. Dichaea morrisii auct. non Fawc. & Rendle 1910: sensu Dunst. and Garay, Venez. Orchid. Ill. 1: 97. 1959; Foldats in Lasser, Fl. Venez. 15(5): 454. 1970, pro parte. Epiphyte, pendulous, shade-loving; stems 25–30 cm long; flowers white; sepals 4.8–5 mm long. Rain forests, 300–400 m; Bolívar (Cerro Ichún, 93 km south of El Dorado). Guyana, Suriname, French Guiana, Ecuador(?).

Dichaea panamensis Lindl., Gen. Sp. Orchid. Pl. 209. 1833. Epiphyte, erect, arched to pendulous, shade-loving, cespitose; stems usually unbranched, 4–15 cm long; roots very fleshy; leaves glaucous, articulated with their sheaths; flowers small, campanulate, perianth segments white or greenish white; lip white; column sometimes red. Rain forests; 50–600 m; Delta Amacuro (Sacupana), Bolívar (El Paují, near Cerro Guaiquinima), Amazonas (basins of Río Cataniapo and Río Sipapo). Apure, Barinas, Portuguesa, Táchi-

Dichaea latifolia Lindl., Gen. Sp. Orchid. 208. 1833. West Indies, Colombia, Ecuador, Peru, Bolivia. 2 varieties, both in Venezuela, 1 in the flora area.

Dichaea 303

ra; Mexico, Central America, Colombia, Ecuador, Peru, Brazil (Amazonian states). Dichaea pendula (Aubl.) Cogn. in Urb., Symb. Antill. 4: 182. 1903. —Limodorum pendulum Aubl., Hist. Pl. Guiane. 819. 1775. Dichaea pendula var. swartzii C. Schweinf., Bot. Mus. Leafl 17: 62. 1955, nom. illeg. —Dichaea swartzii (C. Schweinf.) Garay & H.R. Sweet, J. Arnold Arbor. 53: 397. 1972. Epidendrum echinocarpum Sw., Prodr. 124. 1788, nom. superfl. —Dichaea echinocarpa (Sw.) Lindl., Gen. Sp. Orchid. Pl. 208. 1833, nom. illeg. Epiphyte, shade-loving, medium- to largesized for the genus, pendulous; flowers subcampanulate; perianth segments whitish or greenish; lip blotched with violet-purple. Rain or cloud forests, (800–)1000–2100 m; Amazonas (Sierra de la Neblina), Bolívar (Cerro Uroi, El Paují, La Escalera to Cerro Venamo region, Macizo del Chimantá, Ptaritepui, Roraima-tepui). Aragua, Nueva Esparta, Sucre, Yaracuy; West Indies, Colombia, French Guiana. Dichaea picta Rchb. f. in Saunders, Refug. Bot. pl. 84. 1872 [1869]. Epiphyte, small to medium-sized, cespitose, erect, arching to pendulous, shadeloving; stems 5–20 cm long, unbranched or branched near base; flowers small, campanulate, white with red specks. Rain forests, 50– 1000 m; Delta Amacuro (Río Amacuro, Río Araguao, Río Grande), Bolívar (Altiplanicie de Nuria, Río Caroní, Río Caura, Río Parguaza), Amazonas (Cerro Yutajé, Río Casiquiare, Río Negro). Miranda, Nueva Esparta; Trinidad, Guyana, Ecuador. Dichaea robusta Schltr., Repert. Spec. Nov. Regni Veg. Beih. 27: 83. 1923. Dichaea morrisii auct. non Fawc. & Rendle 1910: sensu Dunst. and Garay, Venez. Orchid. Ill. 1: 97. 1959, pro parte; Foldats in Lasser, Fl. Venez. 15(5): 454. 1970.

Dichaea muricata auct. non (Sw.) Lindl 1833: sensu J. Folsom, A Systematic Monograph of Dichaea Sect. [Subg.] Dichaea, unpublished doctoral thesis, University of Texas, Austin. 1987, pro parte. Epiphyte, erect to pendulous, shade-loving; stems 10–55 cm long; flowers greenish or greenish yellow, sometimes streaked with brown or violet; sepals 12–14 mm long; lip violet or whitish with violet or brown blotches. Rain or cloud forests, (50–)400– 1500 m; Delta Amacuro (Agua Muerta), Bolívar (basin of Río Caroní), Amazonas (Sierra de la Neblina). Mérida, Miranda, Portuguesa, Zulia; Costa Rica, Panama, Colombia, Ecuador, Peru, Bolivia. Dichaea robusta is part of a complex of at least two species (R. Dressler, personal communication), both of which have been confused under one name. One of them, to which the name D. robusta applies, occurs from Costa Rica southward to at least Peru. It also occurs in the flora area. The second entity, D. morissii, occurs in the West Indies (type locality) and Central America. They are sympatric in Costa Rica. Dichaea robusta features broader sepals and petals that are acute to obtuse at apex, the claw of the lip is thick, the lateral lobes of the lip are long and reflexed, and the lip apex is more or less truncate to rounded. Dichaea morrisii, on the other hand, displays narrower, narrowly acute to acuminate petals and sepals, short, straight lateral lobes to the lip, and the lip apex is acute. Flower color is also different. Dichaea robusta usually has greenish or green-white flowers with a few red or purple spots, and D. morrisii is characterized by white flowers with pink-purple stripes. Both species have dark purple-red lips, these somewhat darker in D. robusta. Dichaea splitgerberi Rchb. f., Ned. Kruidk. Arch. 4: 327. 1859. Dichaea kegelii auct. non Rchb. f. 1877: sensu Dunst. and Garay, Venez. Orchid. Ill. 4: 67. 1966; Foldats in Lasser, Fl. Venez. 15(5): 454. 1970.

304

O RCHIDACEAE

Fig. 289. Dichaea venezuelensis

Fig. 290. Dichaea hystricina

Epiphyte, pendulous, shade-loving; stems to 50 cm long; leaves sometimes browntinged; flowers campanulate; perianth segments brownish green or dull cream-yellow with violet-purple spots within; sepals 7–99 mm long; lip white with zones of blue-violet or totally blue-violet. Rain forests, 50–1300 m; Delta Amacuro (Río Acure), Bolívar (Altiplanicie de Nuria, El Paují, La Escalera to Cerro Venamo region, Ptari-tepui, San Ignacio de Yuruaní), Amazonas (Cerro Aratitiyope, Cerro Huachamacari, Cerro Duida, Cerro Marahuaka, near San Carlos de Río Negro, northwest of Yutajé). Guyana, Suriname, French Guiana. Dichaea splitgerberi has been known until recently by the name D. kegelii, a name that actually applies to plants with leaves articulated to their sheaths.

Dichaea histrio auct. non Rchb. f. 1859: sensu Dunst. & Garay, Venez. Orchid. Ill. 5: 66. 1972. Epiphyte, pendulous, shade-loving; leaves marcescent; perianth segments widely spreading, green or greenish white; sepals 10–13 mm long; lip with violet-purple zones. Presumably cloud forests, 1000–2000 m; Amazonas (Cerro Marahuaka). Táchira; Colombia, Ecuador, Brazil. Dichaea tachirensis is included here based on its inclusion by Dunsterville and Garay in Orchids of Venezuela: An Illustrated Field Guide (1: 156. 1979). We have not seen any material of Dichaea histrio from the flora area. However, we have seen material apparently referable to it from French Guiana, which makes it likely it occurs in the flora area.

Dichaea tachirensis G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1130. 2000.

Dichaea trinitensis Gleason, Bull. Torrey Bot. Club 54: 605. 1927. —Dichaea

Dimerandra 305

rendlei var. trinitensis (Gleason) Foldats, Acta Biol. Venez. 2: 404. 1959. Dichaea rendlei var. rendlei auct. non Gleason 1927: sensu Foldats in Lasser, Fl. Venez. 15(5): 471. 1970. Epiphyte, shade-loving, cespitose, nearly erect, arched, or pendulous; leaves narrow, membranous, sometimes reddish purple when living; flowers whitish or greenish; sepals 4–9 mm long; lip white. Rain forests, 50–200 m; Amazonas (basin of Río Casiquiare, basin of Río Cataniapo, Río Sipapo). Trinidad-Tobago, Guyana, Suriname, French Guiana. Dichaea trulla Rchb. f., Beitr. Orchid.-K. C. Amer. 104. 1866 [1867]. Epiphyte, erect to subpendulous, cespitose, shade-loving; perianth segments not opening widely, white; sepals 7–11 mm long; lip red- or purple-specked. Rain forests, 100– 800(–1500) m; Bolívar (widespread), Ama-

zonas (Cerro Marahuaka, basin of Río Casiquiare, Río Cataniapo, Río Negro, Río Sipapo, Sierra de la Neblina). Táchira; Central America, Colombia, Ecuador, Peru, Guyana, Suriname, French Guiana, Brazil (widespread). Dichaea trulla is easily recognized by the long, narrow leaves. Dichaea venezuelensis Carnevali & I. Ramírez, Novon 3: 102. 1993. Epiphyte, shade-loving, pendulous, cespitose, rarely branching; flowers small, apparently autogamous; perianth segments and lip white; column with ventral face red-spotted. Rain forests, 400–900 m; Bolívar (Río Caroní basin), Amazonas (Río Ararí). Miranda, Táchira; Ecuador. ◆Fig. 289. Dichaea venezuelensis is related to a number of species from northwestern South America (particularly from the Pacific slopes of the Andes) and Central America.

40. DIMERANDRA Schltr., Repert. Spec. Nov. Regni Veg. 17: 43. 1922. [Subtribe Laeliinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, rarely lithophytes, cespitose. Roots flexuous, glabrous; rhizomes short. Stems erect to horizontal, fleshy, several-articulate, distichously leafy throughout, covered with thin, imbricate leaf sheaths, old stems striate-sulcate and sometimes hollow. Leaves subcoriaceous, conduplicate, articulate, usually concave, shiny when fresh, oblong-linear to ligulate, sessile. Inflorescences 1–3, terminal, without peduncle, 1–3 successively flowered. Flowers with spreading perianth segments, showy, short-lived, pink to purple, rarely white; floral bracts inconspicuous; pedicellate ovary terete, elongate. Sepals similar, free; lateral sepals oblique, all ± linear-lanceolate, acuminate; petals ± rhombic to elliptic, the margin often lobed along one side. Lip either free to base or laterally adnate to column basally, the blade cuneate-flabellate; disk under column with a callus of imbricate lamellae in 3 rows with an additional callose ridge on each side. Column short, somewhat arched, with 2 prominent lobes to the clinandrium, both of which are externally keeled; anther incumbent, operculate, longitudinally septate, small, completely hidden within the lobes of the clinandrium; pollinia 4, compressed; stigma ventral. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 8 species, 2 in Venezuela, both in the flora area. Dimerandra is related to Epidendrum but is distinguished by the lip ± free from the column and the lack of a viscidium.

306

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Key to the Species of Dimerandra 1.

1.

Callus with 3 rows of lamellae united into transverse plates at apex with a few free-standing plates in front of it; columnar lobes transversely oblong, acute to obtuse, margins noncellular ............................ D. elegans Callus with 3 rows of lamellae free at the tips, without accessory tubercles or plates in front of it; columnar lobes subquadrate with rounded angles, margins prominently cellular .................. D. emarginata

Dimerandra elegans (H. Focke) Siegerist, Bot. Mus. Leafl. 30: 207. 1986. —Isochilus elegans H. Focke, Tijdschr. WisNatuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 4: 68. 1851. Epiphyte. Semideciduous to rain forests, near sea level to 300 m; Delta Amacuro (Río Acure, Sacupana), northwestern Bolívar, Amazonas (near Samariapo). Apure, Aragua, Distrito Federal, Guárico, Merida, Miranda, Sucre, Táchira, Yaracuy, Zulia; Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 291. Dimerandra emarginata (G. Mey.) Hoehne, Bol. Agric. Estado São Paulo 34: 618. 1934. —Oncidium emarginatum G. Mey., Prim. Fl. Esseq. 259. 1818. Epiphyte. Rain forests, near sea level to 100 m; southeastern Delta Amacuro, northeastern Bolívar. Barinas, Táchira, Zulia; Mexico to Costa Rica, Trinidad-Tobago, Guyana, Suriname.

Fig. 291. Dimerandra elegans

Discyphus 307

41. DIPTERANTHUS Barb. Rodr., Gen. Spec. Orchid. 2: 232. 1882. [Subtribe Ornithocephalinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Twig epiphytes, small to minute, cespitose. Pseudobulbs small, oblong, ovoid, or spherical, 1-foliate, rarely absent, usually with 1–several leaf-bearing sheaths. Leaves conduplicate, coriaceous, flat, usually narrow-elliptic, acute. Inflorescences lateral, originating from the pseudobulb base, erect to arching, racemose, lax to dense, few- to many-flowered, as long or longer than leaves. Flowers small, not resupinate, usually yellow or greenish yellow; floral bracts inconspicuous; pedicellate ovary relatively long. Sepals free, similar or the lateral ones falcate and reflexed; petals similar to sepals or much broader. Lip fleshier than the other perianth segments, erect or decurved, unlobed, cymbiform (boat-shaped), symmetrical or oblique, usually pilose on inner face. Column ± erect or curved, terete, footless, wingless, basally with a pair of lobes or projections that are fleshy, linear, obtuse, rarely very reduced, with the basal portion of the column thickened; clinandrium shallow; anther terminal, operculate, incumbent, 1-locular, exappendiculate or with a short appendix; pollinia 4, cartilaginous, ovoid, slightly compressed and appendiculate, tegula long, filiform, viscidium small; rostellum elongate, usually S-shaped; stigma subventral. Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia; 10 or 11 species, 2 species in Venezuela, 1 of these in the flora area. Dipteranthus planifolius (Rchb. f.) Garay, Canad. J. Bot. 34: 758. 1956. —Ornithocephalus planifolius Rchb. f., Ned. Kruidk. Arch. 4: 315. 1859. Twig epiphyte, small, cespitose, sun-loving; inflorescences longer than to ± equaling the leaves; flowers small, greenish yellow or yellow. Rain or cloud forests, 400–1100 m; Bolívar (La Escalera to Cerro Venamo region). Aragua, Distrito Federal, Falcón. Ecuador, Peru. ◆Fig. 292. Fig. 292. Dipteranthus planifolius

42. DISCYPHUS Schltr., Repert. Spec. Nov. Regni Veg. 15: 417. 1919. [Subtribe Spiranthinae]. Spiranthes Rich., De Orchid. Eur. 20, 28, 36. Aug.–Sept. 1817, nom. cons., pro parte. Dikylikostigma Kraenzl., Notizbl. Bot. Gart. Berlin-Dahlem 7: 321. 1919. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial, shade-loving. Roots fasciculate, fusiform. Leaf 1, basal, broadly ovate or suborbicular, cordate, horizontal on the ground. Inflorescence a ± dense, many-flowered spike, erect, pubescent, terminal, flowers in the upper 1/3 of the inflorescence; peduncle terete. Flowers small, resupinate, glandulose-pubescent, green and white; ovary cylindric-arched, twisted. Sepals similar, spreading; dorsal sepal

308

O RCHIDACEAE

deeply concave, basally fused with lateral sepals for a short distance; lateral sepals connate at base, long-decurrent on column foot, together forming an internal cupshaped nectary; petals ± sinuous, free from dorsal sepal, decurrent at base. Lip longclawed, the claw fully adnate to connate part of lateral sepals, fleshy-sagittate at base. Column short, conduplicate-furrowed in front, at base with a long, decurrent, incurved foot; anther ovate, acute, dorsal; pollinia 4, soft, clavate, with narrowly elliptic viscidium; stigmas 2, terminal on truncate column, cup-shaped, well separated by the frontal furrow of column; rostellum triangular, notched at it apex. Panama, Venezuela, Trinidad-Tobago, Brazil; 1 species. Discyphus scopulariae (Rchb. f.) Schltr., Repert. Spec. Nov. Regni Veg. 15: 417. 1919. —Spiranthes scopulariae Rchb. f., Bonplandia (Hanover) 2: 11. 1854. —Borax. Dikylikostigma preusii Kraenzl., Notizbl. Bot. Gard. Berlin-Dahlem 7: 321. 1919. Terrestrial, shade-loving; leaf horizontal on soil; flowers small, green and white. Rain forests, ca. 200 m; Bolívar (Río Paragua). Aragua, Distrito Federal, Monagas; Panama, Venezuela, Trinidad-Tobago, Brazil. ◆Fig. 293.

Fig. 293. Discyphus scopulariae

Duckeella 309

43. DRYADELLA Luer, Selbyana 2: 207. 1978. [Subtribe Pleurothallidinae]. Masdevallia sect. Rhombopetalae Kraenzl., Repert. Spec. Nov. Regni Veg. 34: 188. 1925. Masdevallia sect. Saltatrices auct. non Rchb. f. 1877: sensu Woolward, Masdevallia part 12. 1896. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes or cespitose herbs, small to minute. Rhizomes short; ramicauls short, terete. Leaves fleshy, terete to deeply grooved, rarely flat, erect to perpendicular and longer than the ramicaul. Inflorescences racemose, 1- or successively few-flowered, originating from an annulus, shorter than the leaves. Flowers resupinate, small, fleshy. Sepals usually caudate at the apex, concolorous or variously punctate or banded; the laterals basally connate into a cup-like synsepal, with a thickened transverse callus, the free portions sharply decurved at the callus; dorsal sepal usually free or almost so; petals short, broad, multi-angled, originating from the column foot. Lip hinged to column foot by a long, slender claw, then expanded into a callose blade; blade subquadrate, usually rounded at apex, often sagittate basally. Column basally terete, apically winged; anther ventral, operculate; clinandrium 3-lobed; pollinia 2, viscidium tiny; stigma ventral. Southern Mexico, Central America, Colombia, Venezuela, Ecuador, Peru, southern Brazil; ca. 40 species, 1 in Venezuela. Dryadella is a recent segregate from Masdevallia, from which it is easy to distinguish by its ecallose petals and transversely callose synsepal. Dryadella lueriana Carnevali & G.A. Romero, Novon 1: 73. 1991. Minute epiphyte; inflorescence 1-flowered; flowers not spreading widely, pale brown overlaid by red. Dwarf, mossy cloud forests, 1600–1700 m; Bolívar (Cerro Guaiquinima). Amazonian Brazil. ◆Fig. 294. Fig. 294. Dryadella lueriana

44. DUCKEELLA Porto & Brade, Anais Reunião Sul-Amer. Bot. 3: 31. 1940. [Subtribe Pogoniinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial herbs, small to medium-sized, sun-loving; stem elongate with leaves along its length, or the stem ± abbreviate. Roots fasciculate, pubescent. Leaves coriaceous, convolute, few to several arranged basally in rosette fashion, others arranged distichously along the stem, the lowermost larger, the distal ones bract-like, linear, linear-lanceolate, or oblong, black in dried specimens. Inflorescences erect, racemose or paniculate, few- to many-flowered, much surpassing the leaves, with a small leaf-bearing sheath in basal 1/2; floral bracts shorter than the terete pedicellate ovary. Flowers medium-sized, campanulate, resupinate, shortlived, yellow or yellow-green, glabrous. Sepals ± equal, free, with several well-developed longitudinal veins, ± fleshy; petals of similar texture of sepals or more mem-

310

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branous, nearly equal to or broader than sepals. Lip fleshier than the other perianth segments, nearly sessile, erect, concave, basally 3-lobed, lateral lobes small, auriculate to elliptic, in natural position clasping the column; central lobe larger than laterals, oblong to narrowly obovate, disk with a much-crested callosity at base, crests apically fimbriate. Column short, footless, apically with a pair of erect wings; anther terminal, operculate, 2-locular; pollinia 4, granulose; rostellum transverse; stigma ventral. Colombia, Venezuela, Brazil; 3 species, 2 in Venezuela, both in the flora area. The architecture of the plants in this genus is somewhat difficult to interpret. There are several, basal leaves arranged in a rosette, and some additional distal ones along the stem. These distal leaves are conspicuously smaller and could be interpreted as being inflorescence bracts, the stem as being a thickened inflorescence, thus making the plant rosette-like. Key to the Species of Duckeella 1. 1.

Sepals fleshy; leaves 9–17 mm wide .............................................. D. alticola Sepals membranous; leaves 3–8 mm wide ................................ D. pauciflora

Duckeella alticola C. Schweinf., Bot. Mus. Leafl. 19: 195. 1961. Flowers dull yellow or yellow-green. Bogs, wet tepui associations, 1300–2000 m; Bolívar (widespread), Amazonas (Cerro Huachamacari). Endemic. ◆Fig. 296.

Fig. 295. Duckeella pauciflora

Fig. 296. Duckeella alticola

Elleanthus 311

Duckeella pauciflora Garay, Bot. Mus. Leafl. 18: 168, t. 34. 1958. Duckeella adolphii auct. non Porto et Brade 1940: sensu Foldats in Lasser, Fl. Venez. 15(1): 164. 1970. Flowers bright yellow, the callus with dull orange keels; Amazonian savannas, 50–200

(–500) m; Amazonas (widespread). Amazonian Colombia, Amazonian Brazil. ◆Fig. 295. This species is closely related to the generic type, Duckeella adolphii, but features much wider petals. Material reported from the Venezuelan Amazonas as being D. adolphii clearly belongs in D. pauciflora.

45. DUNSTERVILLEA Garay in Dunst. & Garay, Venez. Orchid. Ill. 5: 70. 1972. [Subtribe Oncidinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Twig epiphytes, minute, cespitose, pseudomonopodial. Roots many, thin, fleshy, pubescent. Stem short, ascendent. Leaves congested toward apex, fleshy, equitant, elliptic to oblong-elliptic, acute; sheaths basally imbricate, articulate. Inflorescences erect, lateral, ± equaling the leaves to slightly longer, few-flowered; floral bracts flattened, distichous, ± equaling the short, terete pedicellate ovary. Flowers minute, campanulate, white or yellow-green, erect. Petals and sepals similar, linear-oblong, acute, dorsally keeled, mucronate; lateral sepals slightly oblique. Lip fleshier than other perianth segments, ovate, unlobed, obtuse, at base with a conic, saccate, internally pubescent spur, disk 2-lamellate, transversely keeled between the lamellae. Column cylindric, footless; anther incumbent; clinandrium obliquely ascendent, slightly marginate; pollinia 4, globose, tegula linear, viscidium small, ovate-rhombic; rostellum elongate; stigma large, filling the whole ventral face of column. Capsule triquetrous. Venezuela, Ecuador; 1 species. Dunstervillea mirabilis Garay in Dunst. & Garay, Venez. Orchid. Ill. 5: 70. 1972. Twig epiphyte. Rain or cloud forests, 400– 1400 m; Bolívar (La Escalera to Cerro Venamo region, Río Carrao). Ecuador. ◆Fig. 297.

Fig. 297. Dunstervillea mirabilis

46. ELLEANTHUS C. Presl, Reliq. Haenk. 1: 97. 1827. [Subtribe Sobraliinae]. Evelyna Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 32, t. 56. 1835 [1836]. Adeneleutherophora Barb. Rodr., Gen. Spec. Orchid. 2: 170. 1882. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes or terrestrial or subterrestrial herbs, erect to pendulous, cespitose, often forming dense clumps. Roots relatively thick, glabrous; rhizomes short. Stems thin to relatively thick, woody, elongate, branched or unbranched, much-articulated, terete or ± laterally compressed, covered by sheaths, foliose in the apical 3/4. Leaves convolute, plicate or rarely subconduplicate, articulate, mostly distichous, soft or rigid, sessile; blade linear to lanceolate, usually glabrous; veins prominent, especially on the lower surface; sheaths amplexicaul to funnel-shaped, often pubescent or verruculose. Inflorescences terminal or (in some Andean species) originating directly from the rhizome, erect, racemose, usually with many spiraled or few distichous flowers, or capitate, always shorter than the stem, usually subtended by several foliaceous or scarious bracts; floral bracts usually conspicuous and brightly col-

312

O RCHIDACEAE

ored, rarely inconspicuous or scarious. Flowers resupinate or not, usually campanulate, mostly brightly colored, sometimes white; pedicellate ovary relatively thick and short, smooth or verruculose. Perianth segments membranous; sepals free, ± equal, mostly elliptic, lanceolate or narrowly ovate; petals similar to sepals but usually narrower and apically more obtuse. Lip free from column to base, erect, subequal or longer than the sepals, blade concave in the basal 1/2, funnelform and clasping the column, apically unlobed or inconspicuously 3-lobed, margins entire, denticulate, or fimbriate; disk basally with 1 or 2 prominent calli. Column erect, semiterete, sometimes winged, footless; anther terminal, operculate, ± incumbent or erect, 2-locular or imperfectly 4-locular; clinandrium small; pollinia 8, waxy or soft, ovoid, viscidium absent; rostellum transverse, stigma ventral, emergent. Capsule cylindric. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 80 species, ca. 17 species in Venezuela, 10 of these in the flora area. The genus Elleanthus is much in need of a revision. Key to the Species of Elleanthus 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4).

5.

6(5).

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

Leaves very narrow, linear, 1–3(–5) mm wide, ≥ 15 times longer than wide ................................................................................... E. graminifolius Leaves wider, at least 5 mm wide (usually ≥ 8 mm), < 8 times longer than wide ........................................................................................................ 2 Leaf sheaths inflated, not tightly clasping the stem ......................... E. norae Leaf sheaths not inflated, tightly clasping the stem ................................ 3 Leaf sheaths and floral bracts covered with long, deep red or purple hispid hairs ............................................................................... E. caravata Leaf sheaths and floral bracts glabrous, never hispid ............................. 4 Leaves 10 mm or less wide; stems thin, 2 mm thick, usually branching .... 5 Leaves 18 mm or more wide; stems thick to stout, 2.5 mm thick; stems unbranched ............................................................................................ 7 Stems unbranched or with 1 or 2 basal branches; inflorescences polystichously and densely several-flowered; rachis relatively thick, straight ........................................................................................ E. gracilis Stems strongly branched, with at least 1 lateral branch at each intermediate node; inflorescences distichously and laxly 2–4-flowered; rachis thin, zigzag ............................................................................................. 6 Floral bracts forming an acute angle with the rachis; lip apex deeply emarginate; petal margins entire, fimbriate or erose only in the apical 1/3 and not at apex ..................................................................... E. confusus Floral bracts ± perpendicular to rachis; lip apex entire; petal margins fimbriate at apex and upper 1/3 ..................................... E. malpighiiflorus Inflorescence dense, capitate to subcapitate ................... E. sphaerocephalus Inflorescence lax or dense but never capitate or subcapitate, always longer than wide .................................................................................... 8 Lip basally with a single callus, which may be apically emarginate ................................................................................................... E. wageneri

Elleanthus 313

8. 9(8).

9.

Lip basally bicallose ................................................................................... 9 Lip obovate-pandurate in general outline, not abruptly dilated above the middle into a suborbicular or transversely elliptic central lobe; dorsal sepal 7.5 mm long; central lobe of lip 0.7 cm wide ...... E. arpophyllostachys Lip when expanded abruptly dilated above middle into a suborbicular or transversely elliptic central lobe; dorsal sepal ca. 10 mm long; central lobe of lip 0.8–1 cm wide ...................................................... E. columnaris

Elleanthus arpophyllostachys (Rchb. f.) Rchb. f. in Walp., Ann. Bot. Syst. 6: 479. 1862. —Evelyna arpophyllostachys Rchb. f., Bonplandia (Hanover) 2: 21. 1854. Elleanthus attenuatus J.R. Johnst., Proc. Amer. Acad. Arts 40: 684. 1905. Epiphyte, sometimes terrestrial, 50–100 cm tall; bracts and flowers orange. Rain forests, 600–1500 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, La Escalera to Cerro Venamo region, Macizo del Chimantá [Apacará-tepui]), Amazonas (Cerro Aracamuni). Aragua, Carabobo, Falcón, Mérida, Sucre, Trujillo; Colombia, Ecuador. Elleanthus caravata (Aubl.) Rchb. f., Otia Bot. Hamburg. 2: 92. 1878. —Serapias caravata Aubl., Hist. Pl. Guiane 816, t. 320. 1775. Usually an epiphyte 30–100 cm tall; bracts pink or purplish; flowers yellowish. Rain forests, 300–1300 m; Bolívar (Altiplanicie de Nuria, Aparamán-tepui, JauaSarisariñama massif, La Escalera to Cerro Venamo region, Río Paragua), Amazonas (Cerro Marahuaka, Cerro Yapacana). Lesser Antilles, Guyana, French Guiana, Ecuador. Elleanthus columnaris (Lindl.) Rchb. f. in Walp., Ann. Bot. Syst. 6: 483. 1863. —Evelyna columnaris Lindl., Orchid. Linden. 62. 1846. Terrestrial or epiphyte 60–120 cm tall; bracts and flowers bright purple or magenta at anthesis. Rain or cloud forests, (700–)1200– 2200 m; Bolívar (Cerro Guaiquinima, La Escalera to Cerro Venamo region, Serranía Marutaní), Amazonas (Cerro Marahuaka, Sierra de la Neblina). Falcón, Mérida, Táchira, Trujillo; Colombia. Elleanthus confusus Garay, Bot. Mus. Leafl. 26: 13. 1978.

Elleanthus kermesinus auct. non (Lindl.) Rchb. f. 1862: sensu Foldats in Lasser, Fl. Venez. 15(1): 226. 1970. Elleanthus virgatus auct. non (Rchb. f.) C. Schweinf. 1938: sensu Dunst. & Garay, Venez. Orchid. Ill. 2: 101. 1961. Epiphyte, erect to pendulous, straggling; bracts and flowers pink-lavender to deep purple. Cloud forests, (700–)1400–2600 m; Bolívar (widespread), Amazonas (Cerro Marahuaka, Cerro Yaví, Sierra de la Neblina). Aragua, Zulia. ◆Fig. 298. Elleanthus gracilis (Rchb. f.) Rchb. f. in Walp., Ann. Bot. Syst. 6: 841. 1863. —Evelyna gracilis Rchb. f., Linnaea, 22: 843. 1849. Epiphyte, erect to pendulous, 20–50 cm tall; bracts and flowers pink or pale purple. Rain or cloud forests, (100–)1500–2200 m; Bolívar (Macizo del Chimantá [Toronó-tepui], Roraima-tepui), Amazonas (near San Carlos de Río Negro). Mérida, Portuguesa; Colombia, Ecuador. The San Carlos de Río Negro material from ca. 100 m elevation is included here with reservation since Elleanthus gracilis is a species from cloud forests. Furthermore, the material upon which this citation is based is not in good condition. Elleanthus graminifolius (Barb. Rodr.) Lojtn., Bot. Not. 129: 447. 1977. —Adeneleutherophora graminifolia Barb. Rodr., Gen. Spec. Orchid. 2: 171. 1882. Elleanthus pusillus Schltr., Notizbl. Bot. Gart. Berlin-Dahlem 8: 117. 1922. Epiphyte, usually erect but sometimes ascendent, 10–50 cm tall; bracts scarious; flowers cream-colored or white. Rain or cloud forests, (200–)500–1500 m; Bolívar (widespread), Amazonas (widespread). Widespread elsewhere in Venezuela; Greater Antilles, Honduras, Nicaragua, Costa Rica, Panama, Co-

314

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Fig. 298. Elleanthus confusus

Fig. 299. Elleanthus wageneri

Fig. 300. Elleanthus sphaerocephalus

Eloyella 315

lombia, Guyana, Suriname, Ecuador, Peru, Brazil. The Central American populations may represent a different species. Elleanthus malpighiiflorus Carnevali & G.A. Romero in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1131. 2000. Epiphyte, erect or ascending to drooping, 30–100 cm tall; flowers bright rose to lavender. Low, open cloud forests, usually in low positions, 2300–2400 m. Known only from the Brazilian side of Sierra de la Neblina. Elleanthus norae Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 110. 1976. Terrestrial herb, erect, to 1.5 m tall in open areas, to 3.5 m in forests; bracts greenish with lavender tinge toward apex; sepals pale pink or pale cream flushed with lavender at base; petals and lip pale pink, darker at margins and apex; disk bearing a patch of light orange with dull orange vein lines. Cloud forests or open associations on tepui summits, (1300–)1600–2600 m; Bolívar (Au-

yán-tepui, Macizo del Chimantá), Amazonas (Cerro Coro Coro, Cerro Marahuaka, Sierra Parima). Endemic. Elleanthus sphaerocephalus Schltr., Repert. Spec. Nov. Regni Veg. 27: 17. 1924. Epiphyte or terrestrial, erect to pendulous; flowers often embedded in a gelatinous material within the capitate inflorescence; bracts and perianth segments purple. Rain or cloud forests, 700–1300 m; Bolívar (La Escalera to Cerro Venamo region, Río Chicanán), Amazonas (Sierra Parima). Aragua, Distrito Federal, Mérida, Miranda, Nueva Esparta, Trujillo; Colombia, Ecuador, Peru, Bolivia. ◆Fig. 300. Elleanthus wageneri (Rchb. f.) Rchb. f. in Walp., Ann. Bot. Syst. 6: 474. 1862. —Evelyna wageneri Rchb. f., Bonplandia (Hanover) 2: 21. 1854. Terrestrial 1.5–2 m tall. Cloud forests, 1400–1800 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region). Distrito Federal, Mérida, Táchira, Trujillo. ◆ Fig. 299.

47. ELOYELLA P. Ortiz, Orquideología 13: 233. 1979. [Subtribe Ornithocephalinae]. Phymatidium Lindl., Gen. Sp. Orchid. Pl. 209. 1833, pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Sygmoid twig epiphytes, 1.5–5 cm tall. Rhizome abbreviate. Stems laterally compressed, abbreviate to almost absent. Leaves laterally compressed, articulate with their sheaths, imbricate, 3–7 per stem, linear to obliquely elliptic, acute. Inflorescences axillary from leaf bases, erect, spreading to pendulous, 2–few-flowered racemes longer than leaves; peduncle terete or laterally flattened to winged, shorter than or subequal to the leaves; bracts widely spaced, often perpendicular to peduncle, boat-shaped; rachis longer than peduncle, straight or zigzag. Flowers mostly resupinate, successive or several open simultaneously. Floral bracts similar to peduncle bracts but smaller. Perianth segments widely spreading, membranous to subfleshy, 1-veined, greenish, dark yellow, or whitish; sepals free, subequal; petals free, subequal to sepals or broader, sometimes of different color. Lip free to base, footless, unlobed, elliptic to ovate, margin entire to dentate; disk with a large, concave or convex, plate- or cushion-like callus, often with tuberosities or strap-like projections. Column straight or arched, subterete or dorsoventrally compressed, wingless or winged in the basal 1/3; anther apical, operculate, incumbent; clinandrium small; pollinia 4, ovoid or pear-shaped, tegula large, obtriangular or obtrullate, viscidium small; rostellum small, stigmatic surface ventral, slit-like, longitudinal, close to the base of the column. Panama, Colombia, Venezuela, Ecuador, Peru; 4 species, 1 in Venezuela.

316

O RCHIDACEAE

Eloyella panamensis (Dressler) Dodson, Icon. Pl. Trop. ser. 2, 5: pl. 455. 1989. —Phymatidium panamense Dressler, Orquideología 6: 42. 1971. Twig epiphyte to 2 cm tall including the inflorescence; sepals ca. 2 mm long, pale green; petals and lip white; lip denticulate, callus green, with several tuberosities. Rain forests, often high in trees or lower when close to water courses, 100–200 m; Bolívar (Río Nichare). Panama, Ecuador. ◆Fig. 301.

Fig. 301. Eloyella panamensis

48. ELTROPLECTRIS Raf., Fl. Tellur. 2: 51. 1836 [1837]. [Subtribe Spiranthinae]. Spiranthes Rich., De Orchid. Eur. 20, 28, 36. Aug.–Sep. 1817, nom. cons., pro parte. Centrogenium Schltr., Beih. Bot. Centralbl. 37(2): 451. 1920. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial or humicolous, medium-sized, erect herbs. Roots fleshy, fasciculate. Leaves convolute, 1–several, basal; petiole ± well developed; blades concolorous to variegated, deciduous, occasionally absent at anthesis. Inflorescences terminal, originating from the center of rosette, a few- to many-flowered, lax raceme or spike; peduncle thick to slender, erect. Flowers medium-sized to large, resupinate, frequently white or pink; floral bracts large; ovary cylindric, sessile or nearly so. Sepals free, ± spreading; lateral sepals longer than dorsal sepal, decurrent on column foot, free part of column foot forming a pendulous, spur-like process; petals adherent to dorsal sepal, forming a galea, dorsally ± decurrent on column, variable in shape. Lip membranous, distinctly clawed, arched, recurved, with basal marginal, glandular thickenings, somewhat longer than sepals. Column slender, ± short, erect, with a basal foot adnate to ovary and apically protruding from the ovarian tissue; anther ovate-hooded, persistent; pollinia 4, soft, clavate, viscidium oblong, sheath-like, covering the rostellum remnant; stigmas 2, free, terminal, horizontal, often appearing 2-lobed, ± separated from one another by terminal edge of a distinct fold running full length on face of column; rostellum rigid, ± cartilaginous, subulate to linear, sharply pointed. U.S.A. (Florida), Mexico, Central America, West Indies, Colombia, Venezuela, Suriname, Ecuador, Peru, southern Brazil, Bolivia, Paraguay, northern Argentina; 12 species, 2 in Venezuela, 1 of these in the flora area. Eltroplectris calcarata (Sw.) Garay & Sweet, J. Arnold Arbor. 53: 390, fig. 71. 1972. —Neottia calcarata Sw., Fl. Ind. Occid. 3: 1413, t. 28. 1804. —Centrogenium calcaratum (Sw.) Schltr., Beih. Bot. Centralbl. 37(2): 452. 1920. Pelexia setacea Lindl., Gen. Sp. Orchid. Pl. 482. 1840. —Centrogenium setaceum (Lindl.) Schltr., Beih. Bot. Centralbl. 37(2): 453. 1920.

Terrestrial or humicolous herb, erect; leaves rosulate, basal; inflorescence to 50 cm long; flowers white, basally greenish, with widely spreading lateral sepals; dorsal sepal and petals nearly parallel to column and forming a hood over it. Semideciduous forests, 400–500 m; Bolívar (Cerro Arimagua southeast of Báquiro). Falcón; U.S.A. (Florida), West Indies, Colombia, Ecuador, southern Brazil. ◆Fig. 302.

Encyclia 317

Fig. 302. Eltroplectris calcarata

49. ENCYCLIA Hook., Bot. Mag. 55: t. 2831. 1828. [Subtribe Laeliinae]. Epidendrum L., Gen. Pl. 272. 1737, pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, lithophytes, or sometimes subterrestrial herbs, usually erect, rarely subpendulous, mainly sun-loving but often growing in shade. Rhizome short, creeping to subscandent, terete, rooting at internodes. Stems pseudobulbose-thickened (except one Central American species); pseudobulbs heteroblastic (one internode), pear-shaped to subspheric, more rarely fusiform to cylindric, apically or subapically 1–3(4)-foliate, basally enveloped by tubular, applicate, eventually deciduous sheaths. Leaves subcoriaceous to rigid-fleshy, conduplicate, articulate, usually flat but sometimes concave, not petiolate, linear, oblong to elliptic or broadly elliptic, midvein not prominent. Inflorescences terminal, from the mature pseudobulb or rarely from the developing pseudobulb, racemes or panicles, rarely few-flowered; peduncle and rachis smooth to verruculose, peduncle few- to many-articulated; bracts inconspicuous; rachis straight to zigzag. Flowers usually resupinate, fragrant, small to large and showy, long-lived, sometimes cleistogamous; floral bracts shorter than pedicellate ovary, often inconspicuous, fleshy; pedicellate ovary cylindric, smooth or verruculose, longer than column. Perianth segments fleshy or fleshy-membranous, variously colored, widely spreading to subcampanulate; sepals

318

O RCHIDACEAE

free, similar or the laterals somewhat oblique; petals usually similar to sepals. Lip free to base from column or more rarely ventrally connate in its basal 1/3–1/4, fleshier than the other perianth segments; blade usually 3-lobed, rarely unlobed, lateral lobes narrower than central lobe; disk almost ecallose to variously callose, glabrous or pubescent; calli usually of several raised, longitudinal keels, or plate-like. Column erect, footless, straight or somewhat recurved, parallel to lip, often with a pair of auricles in the apical 1/3, flanking the stigma and usually embracing the claw between the midlobe and the lateral lobes of the lip; anther operculate, incumbent, 2locular; clinandrium 3-dentate, teeth conspicuous or not; pollinia 4, subreniform, with caudicles, rostellum transversely perpendicular to column; stigma ventral, semiorbicular, obcordate, or transversely fiddle-shaped. U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, northern Argentina; ca. 120 species, ca. 18 in Venezuela, 12 of these in the flora area. Key to the Species of Encyclia 1.

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

4(3).

4.

5(4).

5.

6(5).

Rhizome clearly creeping and elongate; pseudobulbs spaced 3–6 cm apart, fusiform or narrowly ovoid-ellipsoid; plants often lithophytic ................................................................................................... E. granitica Rhizome short; pseudobulbs spaced < 2 cm apart, usually pear-shaped or subspheric; plants usually epiphytic, rarely lithophytic ..................... 2 Column without a pair of auricles at each side of the stigmatic surface ................................................................................................................ 3 Column with a pair of auricles at each side of the stigmatic surface that embrace the claw at base of the central lobe of the lip ........................ 7 Central lobe of lip (2.5–)3–5 cm wide; plants from lowland dry or deciduous forests ................................................................................ E. cordigera Central lobe of lip < 1.5 cm wide; plants from a variety of environments but rarely from lowland dry or deciduous forests (except E. leucantha) ................................................................................................................ 4 Central lobe of lip longer than wide (width/length ratio 0.8–0.9), acute; callus glabrous; inflorescence a very lax raceme or a panicle with 1 lateral branch, usually to 2.5 times longer than the leaves ....................... ....................................................................................... E. auyantepuiensis Central lobe of lip wider than long (width/length ratio 1.2–1.4), obtuse to truncate; callus densely pubescent with long hairs; inflorescence a dense or somewhat dense panicle (sometimes racemose in very young plants), 1–2 times longer than the leaves ............................................ 5 Sepals and petals rounded to subacute, green to brownish green, not glossy; central lobe of lip 10–13 mm wide, white or pale cream-yellow with longitudinal purple streaks; plants from dry or seasonal vegetation area of the Llanos ............................................................ E. leucantha Sepals and petals usually acute to subacute, glossy dark brown; central lobe of lip 7–9 mm wide, bright yellow without purple streaks; plants from lowland rain forests or tepui slopes ............................................. 6 Inflorescence a dense panicle; sepals 9–11.5 mm long; petals 8.5–11 mm

Encyclia 319

long; central lobe width/length ratio 1.67–1.69; plants from tepui summits or slopes at 700–1500 m ............................................. E. conchaechila 6. Inflorescence a lax panicle; sepals 13–14 mm long; petals 13–15 mm long; central lobe width/length ratio 1.4–1.5; plants from lowland shrublands ..................................................................................... E. pilosa 7(2). Central lobe of lip with acute or cuspidate apex; mature leaves 5–12 cm long ......................................................................................... E. guianensis 7. Central lobe of lip with broadly obtuse to truncate or emarginate apex, sometimes with an apicule at lip apex; mature leaves usually > 15 cm long ......................................................................................................... 8 8(7). Total lip length/pedicellate ovary length ratio < 0.5(0.46–0.48); petals with acute apex; callus of lip very thick and raised, extending longitudinally throughout central lobe ........................................... E. chloroleuca 8. Total lip length/pedicellate ovary length ratio > 0.6; petals with obtuse or rounded apex; callus of lip not extending (or only slightly so) to the central lobe .................................................................................................. 9 9(8). Perianth segments microscopically papillose; central lobe oblong to broadly ovate-elliptic; lateral lobes truncate; inflorescence 1–6-flowered ....................................................................................................... 10 9. Perianth segments smooth; central lobe suborbicular to broadly ovate-elliptic; lateral lobes subacute to rounded, very rarely subtruncate; inflorescences many-flowered on adult plants ........................................... 11 10(9). Dorsal sepal 16–21 mm wide; width/total ratio of lip 1.22–1.27; lip length/pedicellate ovary length ratio 0.60–0.63; plants from southern Amazonas .............................................................................. E. remotiflora 10. Dorsal sepal 23–24 mm wide; width/length ratio of lip 0.9–1; lip length/ pedicellate ovary length ratio 0.79–0.95; plants from Delta Amacuro and easterm Bolívar ............................................................ E. pachyantha 11(9). Pedicellate ovary verruculose; petals and sepals marginally concolorous; petal width/length ratio 0.31–0.41; central lobe of lip broadly elliptic, suborbicular or transversely broadly elliptic, with width/length ratio of 1–1.1; plants from Serranía de Imataca ..................................... E. diurna 11. Pedicellate ovary smooth; petal and sepal with a narrow, hyaline, whitish, or yellowish margin; petal width/length ratio 0.5–0.59; central lobe of lip ovate, with width/length ratio of 0.93–0.95; plants from the Gran Sabana ................................................................................. E. ivonae Encyclia auyantepuiensis Carnevali & I. Ramírez, Lindleyana 9: 61. 1994. Epidendrum oncidioides auct. non Lindl. 1833: sensu Dunst. & Garay, Venez. Orchid. Ill. 2: 151. 1961, pro parte. Epidendrum diurnum auct. non (Jacq.) Rchb. f. 1866 [1867]: sensu Foldats in Lasser, Fl. Venez. 15(3): 233. 1970, pro parte. Lithophyte 40–60 cm tall including the inflorescence; sepals and petals yellow-brown, lip white with purple keels; pedicellate ovary verruculose; dorsal sepal 13–18(–24) mm

long. Open sandstone outcrops, 1000–1100 m; Bolívar (Auyán-tepui). Endemic. ◆Fig. 306. Encyclia auyantepuiensis is related to E. leucantha from the Llanos and the northernmost, driest portion of the flora area. It is rare and apparently always terrestrial. It has an acute central lobe of the lip and unbranched (or shortly 1-branched) inflorescences. Encyclia linearifolioides Kraenzl. from South America south of the Amazon basin has smaller flowers with narrower perianth segments on branching inflorescences.

320

O RCHIDACEAE

Encyclia chloroleuca (Hook.) Neumann, Rev. Hort. ser 2, 4: 138. 1846. —Epidendrum chloroleucum Hook., Bot. Mag. t. 3557. 1828. —Encyclia chloroleuca (Hook.) Schltr., Orchideen 208. 1914. Epidendrum chloranthum Lindl., Edwards’s Bot. Reg. 24: misc. 25. 1838. Epidendrum diurnum auct. non (Jacq.) Rchb. f. 1866 [1867]: sensu Foldats in Lasser, Fl. Venez. 15(3): 233. 1970, pro parte. Encyclia acuta auct. non Schltr. 1825: sensu Carnevali & I. Ramírez, Bol. Com. Orqui. Soc. Venez. Ci. Nat. 23: 34. 1988. Epiphyte 10–30(–40) cm tall including the inflorescence; petals and sepals greenish, often suffused with brown or purple; lip white or pale green with 3 or 4 purple streaks; dorsal sepal 10–14 mm long. Locally common in rain forests or dwarf open forests on white sand, often in well-illuminated positions or low on trees close to rivers, 100–600 m; Delta Amacuro (Río Amacuro, Serranía de Imataca), Bolívar (near Icabarú, Río Cuyuní basin), Amazonas (Río Atacavi, Río Cataniapo, Río Sipapo). Miranda, Monagas; Colombia, Guyana, Suriname, French Guiana, TrinidadTobago, Brazil (Amazonas, Roraima-tepui). The confusion between Encyclia chloroleuca and other related species in the literature and herbaria with E. gravida (Lindl.) Schltr., E. oncidioides (Lindl.) Schltr., and E. diurna (Jacq.) Schltr., makes it impossible at present to determine their true distributions and status. The recently described E. maravalensis Withner from Trinidad is almost certainly conspecific Encyclia chloroleuca is possibly widespread in the northeastern Amazon basin. Encyclia conchaechila (Barb. Rodr.) Porto & Brade, Rodriguésia 1: 28. 1935. —Epidendrum conchaechilum Barb. Rodr., Gen. Spec. Orchid. 1: 53. 1877. Encyclia amicta auct. non (Linden & Rchb.) Schltr. 1919: sensu Carnevali & I. Ramírez, Bol. Com. Orqui. Soc. Venez. Ci. Nat. 23: 36. 1988, pro parte. Epiphyte; leaves 10–20 cm long, 2 or 3 per pseudobulb; flowers with widely opening segments, petals and sepals deep brown-red with yellowish edge, lip pale yellow, anther deep wine purple; column green; dorsal sepal

9–1.5 mm long. Rain forests, close to granitic lajas and white-sand savannas, along rivers, ca. 100–200 m; northern Amazonas (Río Sipapo basin). Possibly Peru, Brazil. ◆Fig. 303. Locally common in restricted localities of the Venezuelan Amazonas, this showy species had not been collected in Venezuela until recently. Encyclia cordigera (H.B.K.) Dressler, Taxon 13: 247. 1964. —Cymbidium cordigerum H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 341. 1815 [1816]. —Epidendrum cordigerum (H.B.K.) Foldats, Bol. Soc. Ven. Ci. Nat. 28: 234. 1969. Epidendrum atropurpureum auct. non Willd. 1805: sensu Dunst. & Garay, Venez. Orchid. Ill. 2: 107. 1961. Epiphyte 30–100(–130) cm tall including the inflorescence; flowers showy, long-lasting, sepals and petals brown-purple, greenpurple, or green; lip white with a purple spot at base of central lobe. Locally common, dry or deciduous forests, ca. 50–400 m; northern Bolívar (Cerro Bolívar, lower Río Caroní and Río Paragua basins, near Upata and Tumeremo). Widespread in northern Venezuela; Mexico, Central America, Colombia, Trinidad-Tobago. Encyclia diurna (Jacq.) Schltr., Repert. Spec. Nov. Regni Veg. Beih. 6: 74. 1919. —Limodorum diurnum Jacq., Collectanea 4: 107. 1790 [1791]. —Cymbidium diurnum (Jacq.) Sw., J. Bot. (Schrader) 1: 221. 1799. —Epidendrum diurnum (Jacq.) Rchb. f., Beitr. Orchid.K. C. Amer. 81. 1866 [1867]. Epidendrum wageneri Klotzsch, Otto & Dietr., Allg. Gartenzeitung 19: 250. 1851. —Encyclia wageneri (Klotzsch, Otto & Dietr.) Schltr., Repert. Spec. Nov. Regni Veg. Beih. 6: 75. 1919. Epiphyte, or rarely a lithophyte, (30–)50– 100 cm tall including the inflorescence; sepals and petals yellow-brown with green areas; central lobe of lip yellowish cream with raised purple keels. Rain forests, 100–200 m; Bolívar (Serranía de Imataca). Aragua, Distrito Federal, Falcón, Lara, Mérida, Miranda; Colombia. Material of Encyclia diurna from the flora area has smaller flowers, a more elliptic cen-

Encyclia 321

tral lobe of the lip, and shorter lateral lobes than the populations from the Coastal Range of Venezuela and almost certainly belongs to a different species. The color description applies only to flora area material. Encyclia granitica (Bateman ex Lindl.) Schltr., Repert. Spec. Nov. Regni Veg. Beih. 6: 74. 1919. —Epidendrum graniticum Bateman ex Lindl., J. Bot. (Hooker) 3: 83. 1840. Epidendrum diurnum auct. non (Jacq.) Rchb. f. 1866 [1867]: sensu Foldats in Lasser, Fl. Venez. 15(3): 233. 1970, pro parte. Epidendrum selligerum auct. non Bateman ex Lindl. 1838: sensu Dunst. & Garay, Venez. Orchid. Ill. 2: 151. 1961. Encyclia oncidioides auct. non (Lindl.) Schltr. 1833: sensu Carnevali & I. Ramírez, Bol. Com. Orqui. Soc. Venez. Ci. Nat. 23: 67. 1988. Lithophyte or rarely an epiphyte; petals and sepals greenish suffused or spotted with brown or purple, lip white with few purple veins; dorsal sepal 13–16 mm long. Rain forests, lajas, near sea level to 400 m; southern Delta Amacuro, Bolívar (Altiplanicie de Nuria, Serranía de Imataca, near Tumeremo). Guyana, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 307. Encyclia granitica is very closely related to and perhaps conspecific with E. oncidioides (Lindl.) Schltr., which is apparently restricted to French Guiana and northeastern Brazil. Encyclia guianensis Carnevali & G.A. Romero, Lindleyana 9: 63. 1994. Epidendrum diurnum auct. non (Jacq.) Rchb. f. 1866 [1867]: sensu Foldats in Lasser, Fl. Venez. 15(3): 233. 1970, pro parte. Encyclia picta auct. non (Lindl.) Hoehne 1952: sensu Carnevali & I. Ramírez, Bol. Com. Orqui. Soc. Venez. Ci. Nat. 23: 70. 1988. Epiphyte to 40 cm tall; petals and sepals bright greenish yellow or brown, typically with red or purple longitudinal stripes; lip white or yellowish; dorsal sepal 10–12 mm long. Locally common in rain forests, 200– 500 m; Bolívar (Altiplanicie de Nuria, La

Escalera to Cerro Venamo region). Guyana, Suriname. Encyclia ivonae Carnevali & G.A. Romero, Lindleyana 9: 65. 1994. Encyclia recurvata auct. non Schltr. 1919: sensu Dunst. & Garay, Venez. Orchid. Ill. 6: 126. 1976, with Epidendrum halatum Garay & Dunst. as a nom. nov. for it. —Encyclia recurvata Carnevali & I. Ramírez, Bol. Com. Orqui. Soc. Venez. Ci. Nat. 23: 73. 1988. Epidendrum diurnum auct. non (Jacq.) Rchb. f. 1866 [1867]: sensu Foldats in Lasser, Fl. Venez. 15(3): 233. 1970, pro parte. Epiphyte 40–120 cm tall including the inflorescence; flowers opening widely; sepals and petals deep green to dark brown-maroon, lip white or yellowish with few purple stripes; dorsal sepal 14–16 mm long. Locally common in rain or riparian forests, 700–1300 m; Bolívar (Auyán-tepui, Gran Sabana, near Perai-tepui). Adjacent Brazil (Roraima). Encyclia leucantha Schltr., Repert. Spec. Nov. Regni Veg. Beih. 6: 40. 1919. —Epidendrum leucanthum (Schltr.) Schnee in Pittier et al., Cat. Fl. Venez. 1: 215. 1945. —Epidendrum leucanthum (Schltr.) C. Schweinf., Bot. Mus. Leafl. 20: 17. 1962. Epidendrum amictum auct. non Linden & Rchb. f. 1855: sensu Dunst. & Garay, Venez. Orchid. Ill. 1: 180. 1959. Epiphyte or lithophyte 20–60 cm tall including the inflorescence; petals and sepals greenish or yellowish, often with brown tinges; lip white, rarely yellowish, usually with few longitudinal purple stripes or veins on the central lobe. Locally common in dry or semideciduous forests, often growing on lajas, ca. 50–200 m; northern Bolívar, and northern half of Amazonas. Widespread elsewhere in Venezuela in the Llanos and the eastern Andean foothills; Colombia. ◆Fig. 305. Encyclia pachyantha (Lindl.) Hoehne, Arq. Bot. Estado São Paulo n.s. formato maior 2: 154. 1952. —Epidendrum pachyanthum Lindl., Edwards’s Bot. Reg. 24: misc. 31. 1838. Epidendrum latipetalum C. Schweinf., Bull. Torrey Bot. Club 72: 219. 1948.

322

O RCHIDACEAE

Fig. 303. Encyclia conchaechila

Fig. 304. Encyclia pilosa

Fig. 305. Encyclia leucantha

Encyclia 323

Fig. 306. Encyclia auyantepuiensis

Fig. 307. Encyclia granitica

324

O RCHIDACEAE

—Encyclia latipetala (C. Schweinf.) Pabst, Orquídea (Niteroi) 29: 9. 1967. Epidendrum diurnum auct. non (Jacq.) Rchb. f. 1866 [1867]: sensu Foldats in Lasser, Fl. Venez. 15(3): 233. 1970, pro parte. Epiphyte to 50 cm tall including the inflorescence; petals and sepals greenish; lip white, purple-striped; dorsal sepal 22–24 mm long. Rain forests, ca. 50 m; Delta Amacuro (Río Amacuro). Guyana, French Guiana, Amazonian Brazil. ◆Fig. 308. Encyclia pilosa (C. Schweinf.) Carnevali & I. Ramírez, Monogr. Syst. Bot. Missouri Bot. Gard. 45: 1257. 1993. —Epidendrum pilosum C. Schweinf., Amer. Orchid Soc. Bull. 11: 361. 1943. Encyclia amicta auct. non (Linden & Rchb.) Schltr. 1919: sensu Carnevali & I. Ramírez, Bol. Com. Orqui. Soc. Venez. Ci. Nat. 23: 36. 1988, pro parte. Epiphyte; flowers brown-purple with a yellow margin, lip yellow or yellow-green; dorsal sepal 13–14 mm long. Shrublands, low open forests, 700–1500 m; Amazonas (Cerro Duida, Cerro Parú). Amazonian Peru (Loreto). ◆Fig. 304. Encyclia pilosa is similar to E. conchaechila, but grows at higher elevations in the Venezuelan Guayana and has larger flowers with broader perianth segments.

Fig. 308. Encyclia pachyantha

Encyclia remotiflora (C. Schweinf.) Carnevali & I. Ramírez, Lindleyana 9: 66. 1994. —Epidendrum remotiflorum C. Schweinf., Bot. Mus. Leafl. 20: 18, pl. 7. 1962. Epidendrum diurnum auct. non (Jacq.) Rchb. f. 1866 [1867]: sensu Foldats in Lasser, Fl. Venez. 15(3): 233. 1970, pro parte. Epiphyte to 50 cm tall; inflorescence very lax, few-flowered; sepals and petals green suffused with dull brown toward the center of the segments; lip white-green with a few purple stripes; dorsal sepal 17–18 mm long. Riparian to rain forests, 100–200 m; Amazonas (Caño Guachapana, Caño Temi, Caño Yagua, Río Atacavi). Endemic.

Epidendrum 325

50. EPIDENDRUM L., Sp. Pl. ed. 2, 2: 1347. Jul.–Aug. 1763, nom. cons. [Subtribe Laeliinae]. Amphyglottis Salisb., Trans. Hort. Soc. London 1: 24. 1812, nom. nud. Auliza Salisb., Trans. Hort. Soc. London 1: 294. 1812, nom. nud. Lanium Lindl. ex Benth. in Hook., Icon. Pl. 14: t. 1334. 1881. Nanodes Lindl., Edwards’s Bot. Reg. 24: t. 1541. 1832. Physinga Lindl., Edwards’s Bot. Reg. 24: misc. 32. 1832. Neolehmannia Kraenzl., Bot. Jahrb. Syst. 26: 478. 1899. Epidendropsis Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 39. 1976. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, lithophytes, subterrestrial, terrestrial, or rarely subaquatic herbs, 0.25–1.5(–7) m tall, cespitose, creeping or subscandent, rarely the whole plant grayish or glaucous. Rhizome short to elongate, or absent. Stems mostly cane-like, or frequently pseudobulbose, sometimes almost absent, simple or variously branched, erect to pendulous, clothed by leaf sheaths to subnaked, distichously leaved throughout or 1–few-leaved apically; pseudobulbs, when present, homoblastic to heteroblastic. Leaves conduplicate, sometimes subterete or laterally compressed, thinly coriaceous to fleshy, usually articulate, sessile. Inflorescence usually terminal, in some cases axillary from leaf sheaths, from base of stem, or from the apex of the developing stem, 1-flowered, racemose, umbellate, capitate, or paniculate, frequently successively flowered, often subtended by 1–few basal spathes; peduncle naked or clothed by several sheaths or spathes, usually terete but sometimes flattened or 3-edged; floral bracts inconspicuous to large and showy; pedicellate ovary variable, mostly terete and smooth. Flowers resupinate or not, campanulate to widely open, very variable in shape and size (sepals 1–100 mm long), usually very fleshy to subfleshy, mostly greenish, yellowish, or whitish, but sometimes of bright colors. Sepals free or rarely the laterals basally connate, variable in shape and relative size, often dorsally carinate; petals free, very rarely basally decurrent-adnate to column, mostly narrower than or subequal to the sepals, very rarely broader. Lip adnate to column at the apex of the latter, very rarely adnate only in the basal 1/2, sometimes the clinandrium is very elongate and oblique and the column looks as if almost free; the adnate zone is often hollow and contains a nectary that frequently embeds itself into the pedicellate ovary, sometimes nectary swollen and noticeable from the outside; the free blade simple to variously lobed, callose or rarely smooth, margins entire or variously erose, dentate, fimbriate, or lacerate, erect on the column, or frequently concave or variously deflexed. Column short to long; often laterally compressed; clinandrium mostly small, sometimes inflated or elongate, margins entire, erose, or denticulate; anther dorsal or subterminal, operculate, incumbent, 2-locular or imperfectly 4-locular; pollinaria variable, pollinia 4 or rarely 2 or 6, caudicles short to elongate, viscidium mostly well developed, rarely almost absent; rostellum horizontal, slit-like, breaks longitudinally when pollinia are detached; stigma ventral. Capsules ellipsoid to 3-edged, smooth or verruculose to echinate.

326

O RCHIDACEAE

U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, northern Argentina; ca. 1500 species, ca. 150 in Venezuela, 65 of these plus 1 hybrid in the flora area. Key to the Species of Epidendrum 1. 1. 2(1).

Rachis and/or peduncle of the inflorescence pubescent ............................ 2 Rachis and/or peduncle of the inflorescence glabrous .............................. 3 Stems well developed, at least 20 cm long, leaves 15–25 cm long; peduncle glabrous ......................................................................... E. caurense 2. Stems abbreviated, 2–6 cm long; leaves 1–3 cm; peduncle pubescent ........................................................................................... E. microphyllum 3(1). Stems thickened into a pseudobulb, this orbicular to spindle-shaped; leaves 1–3(4), clustered at the apex of pseudobulb .............................. 4 3. Secondary stem not thickened into a pseudobulb; leaves usually distributed along stem, sometimes only at its apex ........................................ 9 4(3). Pseudobulb globose, 5–8 mm long; inflorescences very abbreviate, almost sessile ............................................................................................ E. stalkyi 4. Pseudobulbs 3–20 times longer than wide; inflorescences various ......... 5 5(4). Pseudobulbs ≥ 15 times longer than wide, 1-foliate; peduncle 7–20 times longer than the successively few-flowered rachis; sepals connate for about 1/3 of their length .................................................. E. magnicallosum 5. Pseudobulbs ≤ 10 times longer than wide, 1–4-foliate; peduncle shorter, ≤ 3 times longer than rachis and then rachis simultaneously flowered; sepals free .............................................................................................. 6 6(5). Pseudobulbs homoblastic, i.e., no abrupt change in size and thickness of their internodes ..................................................................................... 7 6. Pseudobulbs heteroblastic, i.e., a sharp differentiation in size and thickness at least at 1 or 2 centermost internodes ....................................... 8 7(6). Inflorescences originating from pseudobulb apex, racemose; lip 3-lobed, central lobe linear; dorsal sepal 3.2–9 cm long ........................... E. ciliare 7. Inflorescences originating from pseudobulb base, very rarely from its apex, paniculate; lip 4-lobed due to deeply emarginate central lobe; dorsal sepal 1–2.5 cm long ............................................ E. stamfordianum 8(6). Inflorescences originating from totally mature pseudobulbs, peduncle longer than to ± equal to rachis; lip ± simple to shallowly 3-lobed, central lobe gradually attenuating toward apex; plants from tepui cloud forests above 500 m ..............................................................E. attenuatum 8. Inflorescences originating from immature pseudobulbs, short-pedunculate (rachis much shorter than peduncle); lip sharply 3-lobed, central lobe basally attenuated; plants from lowland forests (80–300 m) ............................................................................................E. purpurascens 9(3). Inflorescences with strongly compressed peduncle ................................ 10 9. Inflorescences sessile, or, when pedunculate, with terete or subterete peduncles ................................................................................................. 11 10(9). Inflorescences in subcapitate racemes or with few subcapitate branches;

Epidendrum 327

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

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

16(15).

16.

17(13).

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

21(19). 21.

central lobe of lip transversely elliptic, truncate, retuse, or emarginate, sometimes conspicuously 2-lobed ......................................... E. anceps Inflorescences amply and laxly paniculate; lip with a fleshy, subconic central lobe ............................................................................... E. compressum At least some of the inflorescences lateral from stem ............................ 12 All inflorescences originating from stem apex ....................................... 13 Lip 1–1.4 times longer than wide, truncate or emarginate; leaves acute, falcate, often purple on lower surface; ovary with a conspicuous vesicle ............................................................................................... E. dichaeoides Lip 1.8–3 times longer than wide, acute; leaves apically obtuse or retuse, concolorous; ovary without a vesicle ........................................ E. sculptum Inflorescences pedunculate with basal part of the peduncle or its whole length covered by relatively large, spathaceous, imbricating bracts .... 14 Inflorescences sessile or if pedunculate, then peduncle not covered with imbricating spathaceous bracts .......................................................... 17 Lip simple or apically emarginate; the whole plant and flowers fleshycoriaceous .................................................................E. amazonicoriifolium Lip distinctly 3-lobed; plants and flowers subcoriaceous to coriaceous ..... 15 Plants to 31 cm tall, epiphytic; inflorescences erect; leaves 5–10 cm long; lateral lobes of the lip small, falcate, acute ......................... E. churunense Plants 70–200 cm, mostly terrestrial; inflorescences pendulous; leaves 9– 26 cm long; lateral lobes of the lip small or large, fimbriate to rounded or obtuse ............................................................................................... 16 Lip with a long, clawed, transverse, fimbriate central lobe; petals and sepals purple- or purple-brown-spotted; plants from elevations below 1000 m (usually 200–600 m) ................................................... E. cristatum Lip with an elliptic or oblong-elliptic, truncate, rounded to emarginate central lobe; petals and sepals not spotted; plants from elevations above 1800 m (usually higher) ....................................... E. klotzscheanum Floral bracts large, imbricating, longer than internodes; rachis totally clothed by floral bracts; floral bracts about as long as ovaries; plants branching freely ................................................................................... 18 Floral bracts inconspicuous to relatively large but never imbricating nor totally enclosing the rachis, or inflorescence 1-flowered ................... 22 Lateral sepals with a large, dorsal conspicuous wing, apically erose; leaves 8–12 mm wide ........................................................... E. urichianum Lateral sepal only carinated on dorsal face; leaves 5–8(–15) mm wide ..... 19 Leaves 3–5 times longer than wide ......................................................... 20 Leaves 7–15 times longer than wide ....................................................... 21 Leaves coriaceous, flat; petals broader toward apex, about 1/2 as wide as dorsal sepal; lip triangular, basally truncate .................... E. strobiliferum Leaves fleshy-coriaceous, cucullate-conduplicate; petals linear or broader toward base; 1/3 or less as wide as dorsal sepal; lip ovate-elliptic, basally cuneate or cuneate-rounded .................................... E. strobiloides Lip obtuse, broadly acute to ± rounded; dorsal sepal 3.5–4.5 mm long; leaves broadening toward base .................................... E. pseudoramosum Lip acuminate; dorsal sepal 6–7.5 mm long; leaves oblong, of uniform width ........................................................................................ E. ramosum

328

O RCHIDACEAE

22(17). Inflorescences 1-flowered, solitary or fasciculate, without evident rachis or peduncle ........................................................................................... 23 22. Inflorescence 1–several-flowered, with distinct peduncle and/or rachis ... 36 23(22). Lip distinctly 3-lobed ............................................................................... 24 23. Lip simple or slightly lobed ..................................................................... 29 24(23). Leaves 2–4, clustered at the apical 1/3 of the stem, very broadly ovate to elliptic, rounded or broadly obtuse; plants usually growing in cloud forests above 1000 m ......................................................... E. carpophorum 24. Leaves 2–10, elliptic, usually evenly distributed along stem, oblong or linear, apex obtuse, acute, or acuminate; plant usually growing in rain forests or open rocky places as epiphytes or terrestrials at various altitudes ..................................................................................................... 25 25(24). Leaves linear or linear-elliptic, acute to acuminate; epiphytic in rain forests, 80–900 m ..................................................................................... 26 25. Leaves elliptic, oblong, or ovate elliptic, obtuse or emarginate, very rarely subacute ............................................................................................... 27 26(25). Plants usually pendulous; leaves at least 80 × 5 mm; stems usually > 10 cm; dorsal sepal 20–30 mm long ..................................... E. longicolle 26. Plants erect; leaves 35–65 × 2–4.5 mm; stems 6–9 cm long; dorsal sepal 15–20 mm long ............................................................. E. micronocturnum 27(25). Plants (30–)50–100 cm long, lithophytic or subterrestrial, rarely low epiphytes on shrublands at elevations of (500–)700–1800 m; petals and sepals 8–12 cm long .................................................. E. tumuc-humaciense 27. Plants 10–50 cm long, epiphytic in tropical rain forests at elevations from near sea level to 700 m ........................................................................ 28 28(27). Petals and sepals (30–)35–80 mm long; plants growing in rainforests ................................................................................................ E. nocturnum 28. Petals 19–25 mm long; plants usually growing in riparian associations in areas of deciduous or semideciduous forests ................................. E. norae 29(23). Pedicellate ovary longer than the leaves ................................ E. apuahuense 29. Pedicellate ovary shorter than the leaves ............................................... 30 30(29). Leaves very fleshy, either semiterete or thickly V-shaped in cross section; plants thickly succulent when fresh, very shriveled in herbarium specimens ..................................................................................................... 31 30. Leaves coriaceous, flat in cross section; plants coriaceous when fresh, chartaceous in herbarium specimens ................................................. 32 31(30). Ventral face and sheaths of leaves smooth; plants erect or prostrate; stems never branching; lip slightly convex broadly cordate, apex acute or obtuse; dorsal sepal acuminate ...................................... E. congestoides 31. Ventral face and sheaths of leaves rugulose; plants usually pendulous; stems usually branching; lip shallowly concave, transversely oblong, apex emarginate; dorsal sepal broadly obtuse, apiculate .......... E. pachyphyton 32(30). Pedicellate ovary with a conspicuous vesicle on ventral face; plants usually purple-tinged, especially on lower surface of leaves ................... 33 32. Pedicellate ovary without a vesicle in the ventral face; plants usually concolorous ........................................................................................... 34 33(32). Plants with extremely abbreviated inflorescences .................. E. dichaeoides 33. Plants with an elongate inflorescence ..................................... E. prostratum

Epidendrum 329

34(32). Plants prostrate or pendulous, usually branching; leaves to 32 mm long; flowers solitary; blade of lip triangular to ovate, acute to obtuse ....................................................................................................... E. repens 34. Plants erect to arching; stems simple; leaves ≤ 5 cm long; flowers solitary or fasciculate; blade of lip transversely widely ovate or reniform, apiculate to shallowly lobed ......................................................................... 35 35(34). Flowers solitary or few (1–3); lip 2.5–3.5 cm wide, subreniform, apically shallowly 4-lobed .................................................................. E. althausenii 35. Flowers fasciculate; lip to 1.5 cm wide, transversely widely ovate, apiculate ............................................................................................. E. sertorum 36(22). Peduncle and rachis thin, wiry, < 0.3 mm thick, inflorescence a laxly flowered panicle, longer than subtending leaves; plants small with leaves < 2.5 cm long, 5 mm wide; sepals < 5.5 mm long, 1-veined; pollinia 2 ............................................................................................... 37 36. Peduncle and rachis thicker, at least 0.5 mm thick, usually much thicker, inflorescence a raceme or a panicle, longer or shorter than subtending leaves; plants small to large but leaves at least 1 cm long and 0.8 mm wide; sepals usually > 6 mm long, 3-veined; pollinia 4 ...................... 38 37(36). Plants from high-tepui associations at elevations usually above 1800– 2000 m, mostly lithophytic or subterrestrial; stems rigidly erect; leaves succulent, often red-purple-tinged; lip transversely oblong or subquadrate, upper surface smooth, margins smooth or coarsely erose ................................................................................................. E. violascens 37. Plants from cloud forests at elevations of 600–1300 m, always epiphytic; stems horizontal to suberect, softer; leaves thinly coriaceous, rarely if ever red-tinged; lip ovate to broadly ovate, upper surface coarsely papillose-verruculose, margins finely cellular-fimbriate ........ E. vincentinum 38(36). Pedicellate ovary with a conspicuous vesicle on ventral face, the vesicle about as long as wide or slightly longer than wide ............................ 39 38. Pedicellate ovary without a conspicuous vesicle on ventral face, if vesicle is present, not prominent and several times longer than thick ........ 40 39(38). Stems 1-leaved; plants cespitose; inflorescence racemose, successively few-flowered; sepals basally connate; plants growing in rain forests between 80–120(–500) m; lip simple; vesicle covered by sepal bases ........................................................................................ E. magnicallosum 39. Stems 2–many-leaved; plants creeping or scandent; inflorescences panicles; sepals free; plants from intermediate to high elevations, 1000–2500 m; lip sharply lobed; vesicle naked ......................... E. saxatile 40(38). Rhizomes distinctly creeping; floral bracts large, navicular and enclosing pedicellate ovary and part of the rachis ................................... E. rigidum 40. Rhizomes very abbreviated; plants cespitose; floral bracts usually small and inconspicuous, when ± conspicuous and navicular, not enclosing the rachis ............................................................................................. 41 41(40). Lip distinctly 3-lobed, the central lobe linear to linear-elliptic or narrowly triangular, at least 3 times longer than wide, simple, acute ............. 42 41. Lip simple or lobed, when lobed, the central lobe not linear or narrowly triangular, not more than 2 times longer than wide, usually 2-lobed or emarginate ........................................................................................... 43

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42(41). Inflorescences usually 1-flowered, rarely 2-flowered, very short to inconspicuous, not subtended by a conspicuous spathe .................................. ............................................... E. nocturnum complex (see couplets 24–28) 42. Inflorescences multiflowered, subtended by a conspicuous spathe ....... 44 43(41). Stems thin, < 1 m long; plants usually epiphytic; inflorescences usually with > 20 flowers; pedicellate ovary < 10 mm long; dorsal sepal 5– 10 mm long ...................................................................... E. klotzscheanum 43. Stems stout, usually > 1.5 m at maturity; plants often terrestrial; inflorescences 3–15-flowered; pedicellate ovary 50–120 mm long; dorsal sepal 30–50 cm long ............................................................. E. leucochilum 44(42). Plants small to 13 cm high (without inflorescences); leaves to 6 mm wide, at least 10 times longer than wide; stems simple; lip simple or emarginate apically ...................................................................................... 45 44. Plants usually > 20 cm tall (without inflorescences); leaves usually > 8 mm wide, when < 8 mm wide then < 7 times longer than wide; stems simple or branched; lip simple or lobed .................................... 48 45(44). Lip wider than long .................................................................................. 46 45. Lip longer than wide ................................................................................ 47 46(45). Lip deeply emarginate apically; inflorescences subcapitate ................. ............................................................................................. E. chimantense 46. Lip truncate to shallowly emarginate; inflorescences laxly racemose .................................................................................................... E. schlimii 47(45). Flowers very fleshy; lip broadly acute to obtuse; inflorescences nodding, sometimes branching; column at least 2.5 times longer than thick; plants growing in open associations on tepui summits ......... E. imthurnii 47. Flowers subfleshy; lip acuminate; inflorescences erect, always simple; column about as long as thick; plants epiphytic in cloud forests ............................................................................................. E. miserrimum 48(44). Inflorescences racemose or paniculate, with flowers distributed along a well-developed rachis, never capitate; pedicellate ovary shorter than the flower-bearing zone of rachis; the flowers developing ± simultaneously so there are always several to many open at a time ............ 49 48. Inflorescences 1–many-flowered, capitate or subcapitate; pedicellate ovary longer to slightly shorter than flower-bearing portion of rachis; flowers may open 7–many simultaneously but are always successive (look for scars on rachis as evidence for old, fallen flowers) .............. 60 49(48). Lip lobed; plants terrestrial or more frequently epiphytic in forests of low to medium elevations; lip flat or convex ............................................. 50 49. Lip simple or nearly so, without sharp sinuses or breaks in lip outline; plants usually terrestrial or subterrestrial on tepui summits in open associations; lip concave (E. dendrobioides-durum complex sensu lato) .............................................................................................................. 53 50(49). Central lobe of lip separated from main blade by a deep, conspicuous constriction (claw) ..................................................................................... 51 50. Central lobe of lip not separated from main blade by a deep, conspicuous constriction (therefore sessile) ............................................................ 52 51(50). Disk with 1 fleshy, longitudinal ridge; petal narrowly obovate; lobes of central lobe parallel to slightly divergent; inflorescences racemose,

Epidendrum 331

51.

52(50). 52. 53(49). 53. 54(53). 54.

55(54).

55. 56(55). 56. 57(56). 57. 58(56). 58. 59(58).

59.

60(48).

60. 61(60). 61. 62(61). 62. 63(62).

pendulous; flowers fleshy ...................................................... E. coronatum Disk with 3 fleshy, longitudinal ridges; petals linear ovate; lobes of central lobe strongly divergent; inflorescences usually paniculate; flowers subfleshy, erect to arching ............................................... E. unguiculatum Disk of lip without fleshy longitudinal ridges; lobes of central lobe rounded ............................................................................ E. agathosmicum Disk of lip with 3 fleshy, longitudinal ridges; lobes of central lobe divergent, acute ............................................................................ E. densiflorum Inflorescences shorter than leaves; lip longer than wide ........... E. imthurnii Inflorescences longer than the leaves; lip as broad or broader than long .............................................................................................................. 54 Flowers pale to deep pink; lip fleshy-coriaceous; spreadable without breaking it, thicker toward base ........................................ E. montigenum Flowers green, yellowish, or cream-colored; lip very fleshy, unspreadable without breaking it, thicker toward the apex (E. dendrobioides-durum complex sensu stricto) .......................................................................... 55 Plants prostrate, sprawling and creeping over boggy substrate; plants apparently restricted to Ilú-tepui, Kukenán-tepui, and Roraima-tepui .......................................................................................................... E. sp. A Plants erect, rarely weakly prostrate over shrubs on tepui shrublands; plants of many places in the Guayana Highlands ............................. 56 Floral bracts 3–7(–8) mm long ................................................................ 57 Floral bracts 9–13 mm wide (since they are conduplicate, the width of one half can be doubled for easy measuring) ............................................ 58 Dorsal sepal 5–7 × 1.9–2.1 mm; lip 4–4.5 mm long, 6–7 mm wide ...................................................................................................... E. durum Dorsal sepal 8–9 × 4–4.5 mm; lip 5–5.5 mm long, 7.5-9 mm wide ...... E. ulei Floral bracts 3–4 times longer than the pedicellate ovary ... E. urbanianum Floral bracts shorter to 2 times longer than the pedicellate ovary ....... 59 Inflorescences strictly erect, unbranched; rachis straight; leaves acute; floral bracts with straight apices; dorsal sepal 11–12 mm long; petals 10–12 mm long; plants from Sierra de la Neblina ... E. commelinispathum Inflorescences arched, decurved, or pendent, often branching; rachis zigzag; leaves obtuse; dorsal sepal 6–7 mm long; petals 4.5–5.5 mm long; plants from Roraima-tepui and neighboring tepuis ..................... E. alsum Lip simple or apically emarginate to ± 4-lobed, never with sharp, deep indentations, reaching at least 1/3 of lip length or width, when sublobed, the lobes originate from apical margin of lip blade ........................... 61 Lip distinctly 3-lobed, with sharp, deep indentations reaching at least 1/3 or more or lip length or width ......................................................... 67 Lip not emarginate ........................................................................ E. nuriense Lip emarginate ......................................................................................... 62 Lip without lobes at each side of emargination ...................................... 63 Lip either with 2 small lobes at each side of the emargination or with a small 3-lobed apical lobe, or altogether without apical lobes ............ 65 Stems usually branched above their base, each branch 3–7 cm long, bearing 2–4 leaves; plants of cloud forests (400–)1500–2500 m; floral bracts at least 1/3 the length of the pedicellate ovary ................... E. chimantense

332

O RCHIDACEAE

63.

64(63). 64. 65(62). 65.

66(65).

66.

67(60).

67.

68(67).

68.

69(68).

69.

70(69). 70. 71(70).

Stems simple, clustered on a short rhizome, at least 20 cm long, bearing 5–25 leaves; plants of rain forests usually below 300 m (rarely to 600 m), often growing in association with ant nests; floral bracts inconspicuous, < 1/5 the length of the pedicellate ovary .............................. 64 Lip transversely broadly ovate to subquadrate, to 1.5 times broader than long ...................................................................................... E. hombersleyii Lip transversely oblong to transversely oblong-elliptic, 2.5–3 times broader than long ........................................................ E. myrmecophorum Flowers pink, membranous, not resupinate; lip longer than broad with lateral indentations or lacerations in the basal 1/3 ................ E. flexuosum Flowers green or greenish with maroon-brown or purple dots or blotches, subfleshy to fleshy, resupinate; lip broader than long, without basal lacerations or indentations ................................................................. 66 Plants epiphytic, often growing in association with ant nests; inflorescence relatively short, usually shorter than 1/3 the length of secondary stem; lateral margins of lip (or lobes) rounded, not projecting forward ...................................................................................... E. myrmecophorum Plants terrestrial growing on sand or sandstone outcrops; inflorescences elongate, longer than 1/2 the length of the secondary stem, often about as long to longer than it; lateral margins (or lobes) of lip somewhat porrect or reniform ........................................................... E. orchidiflorum Flowers green, fleshy; inflorescences shorter than to about as long as the subtending leaf, horizontal or nodding; petals 1/3 as wide as lateral sepals; lateral lobes of lip with entire margin ................ E. smaragdinum Flowers variously colored, never green, subfleshy to membranous; inflorescences longer than subtending leaf (in cases of depauperation it can be equal to subtending leaf); petals broader to about as wide as the lateral sepals; lateral lobes of lip with dentate, laciniate, or fimbriate margins ................................................................................................ 68 Callus complex consisting of several plates and ridges, sometimes at various levels, these placed at lip isthmus; flowers not resupinate ............................................................................. E. secundum (sensu lato) Callus simpler, consisting of 2 hemiellipsoid to globular calli and a narrow central keel which sometimes reaches the apex of central lobe; flowers resupinate or not .................................................................................. 69 Petals distinctly broader than lateral sepals; central lobe of lip rounded or truncate, apiculate, with entire margins; inflorescence relatively short, about as long as to 3 times as long as subtending leaf, very rarely somewhat longer .......................................................... E. flexuosum Petals about as broad as to narrower than lateral sepals; central lobe of lip usually emarginate with a dentate to laciniate margin; inflorescences elongate, 3–10 times longer than subtending leaf (E. ibaguense complex sensu lato) .............................................................................. 70 Central lobe of lip not separated from disk by a conspicuous isthmus or claw (therefore sessile), emarginate ................................................... 71 Central lobe of lip separated from main body of lip by a conspicuous isthmus (hence clawed), emarginate to acuminate .................................. 72 Flowers resupinate; dorsal sepal 6–9 mm long, obtuse; plants terrestrial

Epidendrum 333

on road cuttings or granitic outcrops, not on sandy soil or sandstone outcrops .................................................................................. E. calanthum 71. Flowers not resupinate; dorsal sepal 10–15 mm long, acute to acuminate; plants growing on sandy soil or sandstone outcrops .............................. ........................................................................ E. ibaguense × E. secundum 72(70). Central lobe of lip without a conspicuous apicule or acumen in middle emargination, emargination always evident; plants usually growing as terrestrials or lithophytic; flowers variously colored; dorsal sepal 10– 20 mm long .............................................................................. E. ibaguense 72. Central lobe of lip with a conspicuous apicule or acumen in middle emargination, emargination sometimes not apparent; plants generally growing in ant nests; flowers orange, red, or orange-red; dorsal sepal 19–24 mm long ..................................................................................... 73 73(72). Sepals 15–20 mm long; flowers usually reddish or orange-red ................. ........................................................................................... E. macrocarpum 73. Sepals 25–35(–40) mm long; flowers usually bright orange or orange-red .................................................................................................. E. splendens Epidendrum agathosmicum Rchb. f., Linnaea 22: 841. 1849. Epiphyte to 1 m tall; flowers white, fragrant; dorsal sepal 1–13 mm long. Rain forests, 100–200 m; Amazonas (between Samariapo and San Fernando de Atabapo). Distrito Federal, Lara, Miranda, Trujillo. Epidendrum alsum Ridl. ex Thurn, Timehri 5: 202. 1886. —Epidendrum alsum Ridl., Trans. Linn. Soc. London, Bot. 2: 281. 1887, nom. superfl. Epidendrum durum auct. non Lindl. 1841: sensu Foldats in Lasser, Fl. Venez. 15(3): 240, fig. 466. 1970, pro parte. Terrestrial 25–40 cm tall, erect, sun-loving; leaves with strongly revolute margins, 25–35 × 14–18 mm; inflorescences abruptly deflexed, 3–7 cm long, with 1–3 branches; flowers yellow, campanulate; dorsal sepal 6– 7 × 3.2–3.3 mm. Tepui meadows and rocky outcrops, 2700–2900 m; Bolívar (summit of Roraima-tepui and neighboring tepuis). Probably adjacent Guyana. Epidendrum alsum is one of several members of the Epidendrum dendrobioides-durum complex, a group of mainly terrestrial taxa usually restricted to sandstone-derived or boggy substrates. They are characterized by flowers with extremely fleshy, simple or nearly so, deeply concave lips. Members of this complex are often the most common orchids on the tops of the tepuis and may even

become dominant in some particular habitats. The group, although ranging into some of the West Indies and to southeastern Brazil, seems most diverse in the Guayana Highlands and the resolution of its taxonomy awaits further study, preferably involving field work. In general, plants of lower (800– 1400 m or rarely to 1900 m) elevations are both vegetatively and florally smaller and seem to fit comfortably into one single entity, E. durum. At higher elevations, the plants become stouter with larger flowers, and are more variable in lip and leaf shape, inflorescence architecture, and floral bract size. Frequently, two or more entities are apparently sympatric on most tepuis and the lack of careful observations renders it impossible to assess whether there are some microecological patterns in their local distributions and habitat partitions. The present treatment of the complex, although recognizing far more taxa than previous literature, is probably still very conservative and underestimates the real number of taxa since only the most distinctive populations have been chosen for recognition. True Epidendrum dendrobioides is probably restricted to southeastern Brazil. Epidendrum althausenii A.D. Hawkes, Orquídea (Niteroi) 18: 168. 1957, “althauseni.” —Epidendrum uniflorum Barb. Rodr., Gen. Spec. Orchid. 1: 61. 1877, non Vell. 1831, non Lindl. 1833.

334

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Neolehmannia barbeyana auct. non (Kraenzl.) Garay & Dunst. 1976: sensu Dunst. & Garay, Venez. Orchid. Ill. 6: 38. 1976. Erect to subpendulous epiphyte; flowers solitary, opening well, greenish; dorsal sepal 4–5 cm long. Rain forests, 50–200 m; Delta Amacuro (Río Acure), Amazonas (between La Esmeralda and Ocamo). Amazonian Colombia, Guyana, Suriname, Amazonian Ecuador, Peru, and Brazil. Epidendrum amazonicoriifolium Hágsater, Icon. Orch. 4: 409. 2001. Epidendrum imitans auct. non Schltr. 1921: sensu G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 247. 2000. Epidendrum coriifolium auct. non Lindl. 1851: sensu Dunst. & Garay, Venez. Orchid. Ill. 2: 113. 1961; Foldats in Lasser, Fl. Venez. 15(3): 214. 1970. Epiphyte, rarely a lithophyte, erect or horizontally spreading, to 60 cm long; flowers nonresupinate with widely spreading or reflexed perianth segments, these green or yellow green often with purplish tinges, very fleshy; dorsal sepal 2–2.6 cm long. Cloud or rain forests, 400–1700(–2000) m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Cerro Uaipán, Gran Sabana, La Escalera to Cerro Venamo region, Macizo del Chimantá, Serranía Marutaní, Sierra Pakaraima, Sororopán-tepui), Amazonas (Cerro Duida, Cerro Marahuaka, Sierra Parima). Aragua; Colombia, Guyana, Ecuador, Peru, Bolivia. ◆Fig. 318. Epidendrum amazonicoriifolium is a segregate of E. coriifolium Lindl., which is known from Central America. Epidendrum amazonicoriifolium has among the largest plants and flowers in the E. coriifolium complex, and it is one of the few members of the complex occurring in South America. Epidendrum anceps Jacq., Select. Stirp. Amer. Hist. 224, t. 138. 1763. Epidendrum secundum auct. non Jacq. 1760: sensu Dunst. & Garay, Orchids Venez. Ill. Field Guide 282. 1979. Epiphyte, usually growing in shady spots in forests, to 100 cm tall including the inflorescences, often the plant partially or totally purple-tinged; inflorescences capituliform with 5–15 flowers open at a time; flowers with widely spreading perianth segments,

these fleshy, brownish green to purplish; dorsal sepal 5–10 mm long. Rain forests, 100– 600 m; Bolívar (Altiplanicie de Nuria, La Escalera to Cerro Venamo region, lower Río Caura). Aragua, Barinas, Distrito Federal, Falcón, Monagas, Nueva Esparta, Sucre; West Indies, Guyana, Ecuador, Peru, Brazil. This species is replaced in Mexico and Central America by a closely related species, Epidenrum galeottianum A. Rich. & Galeotti, which is a larger, always dark purple-tinged plant. Epidendrum apuahuense Mansf., Repert. Spec. Nov. Regni Veg. 28: 93. 1930. Epiphyte to 5 cm tall; flowers large for the plant, greenish, whitish, or flesh-colored, cleistogamous; dorsal sepal ca. 2.7–3.8 cm long. Rain forests, 100–200 m; Amazonas (Laja Budare on Río Temi, Río Autana, San Felipe, Yavita to Maroa road). Amazonian Brazil (Amazonas). The Venezuelan populations of Epidendrum apuahuense produce flowers with deformed, asymmetric lips, a frequent condition in cleistogamous plants. At least some Brazilian populations of this taxon produce allogamous, widely spreading flowers. Epidendrum attenuatum Lindl., Fol. Orchid., Epidendrum 41. 1853. Erect epiphyte to 25 cm tall; leaves 4–12 × 3–7 mm; flowers greenish with reflexed petals and sepals; dorsal sepal 4.7–8 mm long. Cloud forests, 1900–2300 m; Bolívar (Macizo del Chimantá). Distrito Federal, Mérida, Trujillo. Ecuador. Epidendrum calanthum Rchb. f. & Warsz., Bonplandia (Hanover) 2: 111. 1854. Epidendrum schomburgkii var. confluens Lindl., Fol. Orchid., Epidendrum 70. 1853. —Epidendrum ibaguense var. confluens (Lindl.) C. Schweinf., Bot. Mus. Leafl. 11: 235. 1944. Terrestrial, lithophyte, or rarely epiphyte, 25–150 cm tall, sun-loving; flowers resupinate; sepals and petals widely spreading, usually pink but often purple or white; dorsal sepal 9–12 mm long. Open roadsides or savanna-like associations, 400–1600 m; Bolívar (Gran Sabana, La Escalera to Cerro Venamo region). Aragua, Sucre, Táchira, Yaracuy; West Indies, Colombia, Ecuador, Peru, Bolivia. ◆Fig. 312.

Epidendrum 335

Epidendrum calanthum is often confused with, or placed in the synonymy of, E. ibaguense, but E. calanthum is ecologically different and the morphology of the smaller flowers is consistently distinct. Leaves of E. calanthum are also thicker than leaves of E. ibaguense. Guayanan populations of E. calanthum differ from Andean populations of the species by having the lip slightly lobed as opposed to entire. Epidendrum carpophorum Barb. Rodr., Gen. Spec. Orchid. 2: 148. 1882. Epidendrum nocturnum var. latifolium Lindl., Edwards’s Bot. Reg. 23: t. 1961. 1837. —Epidendrum latifolium (Lindl.) Garay & Sweet, J. Arnold Arbor. 53: 392. 1972. Epiphyte or lithophyte, rarely terrestrial, shade-loving, erect, arched-ascending, or subpendent, 35–100 cm tall; flowers showy with widely spreading segments but often cleistogamous; sepals and petals greenish with a purplish tint, lip white; dorsal sepal 6–8 cm long. Cloud or rain forests, 700–1500 m; Bolívar (Altiplanicie de Nuria, Amaruaytepui, Cerro Guaiquinima, Gran Sabana, La Escalera to Cerro Venamo region), Amazonas (Cerro Aracamuni). Aragua, Nueva Esparta, Yaracuy; Salvador to Panama, West Indies, Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 320. Epidendrum carpophorum is frequently included in the synonymy of E. nocturnum, but it is found at higher elevations, often from cloud forests, and has 1-flowered inflorescences with very short peduncles hidden inside the sheaths of the upper 1 or 2 leaves. Typically, the leaves number 2–4(5) and increase in size upward, whereas in E. nocturnum leaves are ± the same size along the stem. Epidendrum caurense Carnevali & G.A. Romero, Novon 2: 312. 1992. Erect epiphyte; stems 30–50 cm long; leaves 15–25 cm long; flowers resupinate or not, with widely spreading segments, light yellowish green; dorsal sepal ca. 7 mm long. Rain forests, ca. 300 m; Bolívar (Las Pavas falls at Río Caura). Endemic. Epidendrum caurense is closely related to the Andean E. lanipes Lindl., but has smaller flowers with a differently shaped midlobe to the lip and a different callus. The Andean

species also comes from higher elevations. Some material from Western Colombia could represent E. caurense or a taxon closely related to it. Epidendrum chimantense Hágsater & Carnevali, Icon. Orch. 2: 121. 1993. Epidendrum lechleri auct. non Rchb. f. 1876: sensu Foldats in Lasser, Fl. Venez. 15(3): 297. 1970. Erect epiphyte to 40 cm tall; leaf sheaths usually purple-tinged; flowers resupinate, subcampanulate or with widely spreading segments, greenish, cream-green, or pale lavender, lip yellowish; dorsal sepal 9–10 mm long. Cloud forests, (400–)1500–2300 m; Bolívar (La Escalera to Cerro Venamo region, Macizo del Chimantá, Urimán). Endemic. Epidendrum churunense Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 3: 92. 1965. Erect epiphyte to 33 cm tall; flowers greenish with widely spreading perianth segments; dorsal sepal 8–9 mm long. Rain forests, 400–500 m; Bolívar (Río Chicanán, Río Churún near Guarimba). Endemic. Epidendrum ciliare L., Syst. Nat. ed. 10, 2: 1246. 1759. Herb. Mexico, Central America, West Indies, Colombia, Venezuela, Ecuador, Peru, Brazil; 2 varieties, both in Venezuela, 1 of these in the flora Area. E. ciliare var. squamatum Schnee, Revista Fac. Agron. (Maracay) 1: 206. 1953. Epiphyte or lithophyte, erect; leaves 8–15 cm long; flowers showy, white, with spreading perianth segments; dorsal sepal 3–4.2 cm long; peduncle of the inflorescence covered by imbricate, spathaceous sheaths. Dry forests or (rarely) rain forests, 50–400 m; Bolívar (Altiplanicie de Nuria, La Paragua). Anzoátegui, Monagas; Trinidad-Tobago. Typical Epidendrum ciliare is a plant from rain to cloud forests at low to intermediate elevations, with 1-leaved pseudobulbs and peduncle covered by few tubular sheaths. The variety squamatum is an entity from dry forests or coastal scrub characterized by thicker leaves borne in pairs at the apex of the pseudobulbs, the peduncles of the inflorescences covered by imbricate spathaceous sheaths, and smaller flowers. The con-

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fused nomenclature of the group prevents assigning specific rank to this entity, but it may eventually prove to be one of several specific or subspecific taxa in this complex. Material similar to this coastal Venezuelan entity, but with laxer, longer inflorescences is found in some islands of the West Indies. A thorough revision of the Epidendrum ciliare complex is required before the number, circumscriptions, and distributions of the entities included in it are fully understood. Epidendrum commelinispathum Carnevali & I. Ramírez in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1132. 2000. Terrestrial herb to 1 m tall, sun-loving; leaves 30–45 × 7–9 mm; inflorescences erect, 3–8-flowered; flowers nonresupinate, cream or cream-yellow; dorsal sepal 9–11 × 5–6 mm. Tepui meadows, 1700–2400 m; Amazonas (expected on Sierra de la Neblina). Brazil (known only from the Brazilian side of the summit of Serra da Neblina). The rigidly erect inflorescences with broad floral bracts resembling the spathes of some Commelinaceae are features that distinguish this taxon among other members of the Epidendrum dendrobioides complex. Epidendrum compressum Griseb., Fl. Brit. W. I. 617. 1864. Epidendrum laxum Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 2. 1838, non Sw. 1788. Epidendrum yatapuense Barb. Rodr., Vellosia ed. 2, 1: 123. 1891. Epiphyte to 90 cm tall, including the inflorescences; leaves 5–10(–14) cm long; flowers greenish or yellowish, often tinged with purplish or maroon or entirely maroon, with widely spreading perianth segments; dorsal sepal 5–10 mm long. Rain forests, 50–600 m; Delta Amacuro (Río Amacuro), Bolívar (Río Churún near Guarimba, Río Toro, base of Roraima-tepui, Serranía de Imataca), Amazonas (Río Padamo, Río Siapa, Río Sipapo, San Fernando de Atabapo, Sierra de la Neblina). Colombia, Trinidad-Tobago, Guyana, Ecuador, Peru, Amazonian Brazil, Bolivia. Epidendrum congestoides Ames & C. Schweinf., Sched. Orchid. 10: 61. 1930. Epidendrum schlechterianum auct. non

Ames 1924: sensu Foldats in Lasser, Fl. Venez. 15(3): 381. 1970; Dunst. & Garay, Venez. Orchid. Ill. 2: 147. 1961. Epiphyte, rarely a lithophyte, erect or prostrate; stems 2–5(–7) cm long; flowers very fleshy, widely spreading, segments greenish or whitish with purple or rose tinges, sometimes all floral parts dull maroon; dorsal sepal 8–18 mm long. Rain forests, mangrove forests, near sea level to 50 m; Delta Amacuro (Caño Grande, Caño Simoina west of Isla Cocuina). Aragua, Miranda, Sucre; Mexico, Central America, West Indies, Colombia, Peru, Brazil. Epidendrum coronatum Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 1: 242. 1798. Epidendrum moyobambae Kraenzl., Repert. Spec. Nov. Regni Veg. 1: 185. 1905. Epidendrum amazonicum Schltr., Beih. Bot. Centralbl. 42(2): 78. 1925. Epiphyte, 40–80(–150) cm long, erect to subpendent, usually shade-loving; inflorescence pendent or arched-pendent; flowers resupinate, with widely spreading perianth segments, these cream-white or greenish white, often tinged with purplish; dorsal sepal 16–25 mm long. Rain forests, 100–300 m; Bolívar (La Escalera to Cerro Venamo region, Río Caura), Amazonas (upper Río Ventuari). Anzoátegui, Aragua, Barinas, Distrito Federal, Monagas, Portuguesa, Zulia; Mexico, Central America, Colombia, Trinidad-Tobago, Ecuador, Peru, Amazonian Brazil, Bolivia. Epidendrum cristatum Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 1: 243. 1798. Epidendrum bathyschistum Schltr., Repert. Spec. Nov. Regni Veg. Beih. 6: 36. 1919. Epiphyte or terrestrial, 50–200 cm tall (rarely to 500 cm), erect or horizontally spreading; inflorescence arched-pendent to pendent, all flowers in simultaneous anthesis; peduncle covered with spathaceous sheaths; flowers resupinate, with widely spreading perianth segments, these yellowish or greenish with dark red, purple, or maroon spots or streaks; lip white or yellowish; dorsal sepal 15–30 mm long. Rain forests, 200–600 m; Bolívar (Altiplanicie de Nuria,

Epidendrum 337

La Escalera to Cerro Venamo region, Río Paragua, Río Paramichi). Anzoátegui, Distrito Federal, Monagas, Sucre; tropical South America. ◆Fig. 315. The Central American Epidendrum raniferum Lindl. has often been considered conspecific with E. cristatum, but the midlobe of the lip is much shorter in the former species. The western Amazonian species, Epidendrum criniferum Rchb. f., has also been included in the synonymy, but it is a much smaller plant with narrower leaves, shorter inflorescences, and tendril-like laciniae to the lip. Epidendrum densiflorum Hook., Bot. Mag. 67: t. 3791. 1840. Epidendrum paniculatum auct. non Ruiz & Pav. 1798: sensu Foldats in Lasser, Fl. Venez. 15(3): 332. 1970, pro parte. Terrestrial, lithophyte, or epiphyte, to 100 cm tall, erect; inflorescences in dense panicles; flowers with widely spreading perianth segments, these green with a white lip; dorsal sepal 8–12 mm long. Rain forests; 500– 1100 m; Bolívar (Altiplanicie de Nuria, Santa Elena de Uairén). Aragua, Falcón, Lara, Mérida, Miranda, Táchira; Colombia, Ecuador. A member of the Epidendrum paniculatum complex, E. densiflorum is distinguished by the green flowers with sessile, diverging lobes of the midlobe. Epidendrum dichaeoides Carnevali & G.A. Romero, Novon 2: 314. 1992. —Neolehmannia pabstii Braga, Bradea 3: 171. 1981, non Epidendrum pabstii A.D. Hawkes 1957. Epiphyte, pendent, shade-loving, (5–)8– 19 cm long; leaves usually purplish-tinged or only the abaxial surface so; inflorescences terminal or lateral, 1-flowered; flowers with widely spreading perianth segments, translucent green to dull pale or dark purple, the petals usually paler, lip green or greenish pink with a pink or purple margin; dorsal sepal 10–13 mm long. Rain forests, growing low on moss-covered trunks; 100–200(–600) m; Amazonas (Brazo Casiquiare, Caño Yagua, Río Ararí, Río Negro, near San Carlos de Río Negro, Sierra de la Neblina). Amazonian Brazil. ◆Fig. 313. Epidendrum dichaeoides is locally com-

mon in restricted habitats (white-sand scrub, i.e., banas and surrounding forests in the Río Negro basin), but this distinctive species had remained uncollected in Venezuela until recently. Epidendrum durum Lindl., J. Bot. (Hooker) 3: 87. 1840. Epidendrum carnosum Lindl., J. Bot. (Hooker) 3: 87. 1840. Terrestrial or lithophyte, to 120 cm tall, usually growing in full sun on sandstone outcrops or boggy conditions; leaves 15–30 × 7– 12 mm; inflorescences 5–10 cm long, fewbranched, nodding or arching; flowers white, yellowish, or greenish; dorsal sepal 4–7 mm long. Open tepui associations, 800–1400 (–1900) m; Bolívar and Amazonas (locally common in the Gran Sabana area and the lower and intermediate slopes of the tepuis in all of the flora area). Widespread elsewhere in the Guayana Highlands. See comments under Epidendrum alsum. Epidendrum flexuosum G. Mey., Prim. Fl. Esseq. 260. 1818. Epidendrum imatophyllum Lindl., Gen. Sp. Orchid. Pl. 106. 1831. Epiphyte to 150 cm tall, growing on ant nests, very rarely subterrestrial, usually erect and in full sun; flowers nonresupinate, showy with widely spreading, usually pink or sometimes pale purple perianth segments, lip often of a darker hue, callus yellow; dorsal sepal 10–22 mm long. Rain forests, near sea level to 200(–500) m; Delta Amacuro (Río Amacuro, Río Cuyubini), Bolívar (Río Acanán, Río Carapo, Río Cuyuní, Río Venamo), Amazonas (Río Cataniapo, Río Cuao, Río Siapa, Río Ventuari, San Carlos de Río Negro). Carabobo, Falcón, Miranda, Portuguesa, Yaracuy, Zulia; Mexico, Central America, Colombia, Trinidad-Tobago, Guyana, Ecuador, Peru, Amazonian Brazil, Bolivia, Paraguay. The shape of the lip is variable in Epidendrum flexuosum. Epidendrum hombersleyi Summerh., Bull. Misc. Inform. Kew 1934: 105. 1934. Epidendrum duckei Porto & Brade, Anais Reunião Sul-Amer. Bot. 3: 40, t. 6. 1940. Epiphyte to 1 m tall, growing in ant nests, cespitose, erect to subpendulous; inflores-

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cence with a short peduncle to 40 cm long; flowers opening well or cleistogamous, perianth segments fleshy, greenish, with purple or brownish spots; dorsal sepal 6–6.7 mm long. Rain forests, often at river edges, 50– 200 m; Amazonas (Pimichín, Río Cataniapo, San Carlos de Río Negro). Trinidad, Brazil (Amazonas). ◆Fig. 323. Epidendrum ibaguense H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 352. 1815 [1816]. Terrestrial or lithophyte, 30–150 cm tall, rarely a low epiphyte in shrublands, erect, sun-loving; flowers resupinate with widely spreading perianth segments, these pink, purple, or white, rarely yellow or orange, callus white or yellow, apex of column often dark purple; dorsal sepal (9–)12–18(–20) mm long. Open vegetation over sandy soils or on sandstone or granitic outcrops, 50–1500 m; Bolívar (widespread, particularly in the Gran Sabana area), Amazonas (Cerro Aratitiyope, near Puerto Ayacucho, Río Negro, upper Río Orinoco). Falcón, Lara, Táchira, Trujillo, Zulia; Colombia, Amazonian Brazil, Guyana. ◆Fig. 325. As here understood, Epidendrum ibaguense is a complex of several related entities, some of which might deserve specific recognition. Most local populations tend to be constant in floral morphology, color, and size, but in some others colors are extremely variable. One of the most distinctive populations in the Venezuelan Guayana occurs in the granitic outcrops of the northern Amazonas state and Distrito Cedeño of Bolívar, where flowers are consistently larger (sepals 18–20 mm) and orange or yellow in color while most of the populations of Bolívar are pink, purple, or white. Florally, these populations resemble E. baummanianum Schltr., a species from Costa Rica, Panama, Colombia, and Ecuador that usually grows in ant nests. A thorough revision of the Epidendrum ibaguense complex, which ranges from Mexico to southeastern Brazil and includes entities such as E. laetum Schltr., E. decipiens Lindl., E. filomenoi Schltr., and others is needed. Such a revision can only be completed successfully with an understanding of population variation in the field. Epidendrum ibaguense H.B.K. × Epidendrum secundum Jacq. Terrestrial in open places, usually growing in sandy soils, vegetatively indistinguish-

able and intermediate between the parents; flowers nonresupinate, mostly purplish or white with a yellow callus, size similar or intermediate between the parents. Locally common in shrublands or savanna, 1200– 1500 m; Bolívar (near El Soldado Pionero, Río Tarotá, and elsewhere in the Gran Sabana). Endemic. ◆Fig. 327. The plants of this hybrid grow intermixed with the parents in several places along the road from El Soldado Pionero to Santa Elena de Uairén. They can be recognized by the nonresupinate flowers (as in Epidenderum secundum) with a simple (3-keeled) callus, as found in E. ibaguense. Colors and shapes of the floral segments are intermediate between the parents. Epidendrum imthurnii Ridl. ex Thurn, Timehri 5: 203. 1886. Epidendrum frigidum var. stenophyton auct. non (Schltr.) C. Schweinf. 1944: sensu Foldats in Lasser, Fl. Venez. 15(3): 257. 1970. Terrestrial or lithophyte, to 50 cm tall, in open places, often in boggy conditions; inflorescence nodding; flowers campanulate, greenish or yellow-green, fleshy; dorsal sepal 6–9 mm long. Locally common in tepui tops or slopes, (1400–)1800–2900 m; Bolívar (Auyán-tepui, Cerro Guanacoco, Ilú-tepui, Kamarkawarai-tepui, Kukenán-tepui, Macizo del Chimantá, Murisipán-tepui, Ptari-tepui, Roraima-tepui), Amazonas (Cerro Duida, Cerro Marahuaka, Serranía de la Neblina). Guyana, Amazonian Brazil. Epidendrum imthurnii is one of the most distinctive members of the E. dendrobioides complex due to its very narrow leaves and extremely small flowers. Epidendrum klotzscheanum Rchb. f., Linnaea 22: 838. 1849. Epidendrum ernstii Schltr., Repert. Spec. Nov. Regni Veg. Beih. 6: 37. 1919. Terrestrial to 1.8(–3) m tall, sun-loving; inflorescence nodding; flowers resupinate, perianth segments widely spreading, greenish or yellow-green, often flushed with pink or purple outside; dorsal sepal 11–15 mm long. Open tepui associations, 2000–2500 m; Amazonas (Cerro Marahuaka, Cerro Yaví, Sierra de la Neblina). Anzoátegui, Aragua, Distrito Federal, Mérida, Táchira, Trujillo; Colombia. ◆Fig. 316.

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A specimen of Epidendrum klotzscheanum from Cerro Yaví, K.D. Phelps & Hitchcock 74 (NY), had been reported in the literature as E. alpicolum Lindl. Epidendrum leucochilum Klotzsch,, Allg. Gartenzeitung 11: 145. 1843. Terrestrial or epiphyte to 130 cm tall. Flowers showy, resupinate, with widely spreading, yellow-green perianth segments and a white lip and column; dorsal sepal 4–6 cm long. Rain forests, 400–700 m; Bolívar (Río Carrao). Aragua, Barinas, Distrito Federal, Mérida, Trujillo; Colombia, Ecuador. Epidendrum leucochilum grows at higher elevations in the Andes and Coastal Range. It is known from the flora area from a single specimen cultivated in Mexico and said to have been collected in Río Carrao. Its occurrence in the flora area will remain doubtful until it is collected again. Epidendrum longicolle Lindl., Edwards’s Bot. Reg. 24: misc. 34. 1838. Epiphyte, generally low on tree trunks, pendent or horizontally spreading, shadeloving; flowers showy, resupinate; perianth segments green, cream, or yellow, sometimes purple-tinged, lip white; dorsal sepal 2–3 cm long. Rain forests, 50–1000 m; Delta Amacuro (Río Amacuro), Bolívar (basin of Río Caroní and Río Caura, Serranía de Imataca), Amazonas (basin of Río Casiquiare and Río Negro, Sierra Parima, Yavita–Maroa road). Colombia, Trinidad-Tobago, Guyana, Amazonian Ecuador, Peru, and Brazil. Epidendrum longicolle is very near E. nocturnum but has much narrower leaves and rounded lateral lobes on the lip. Epidendrum macrocarpum Rich., Actes Soc. Hist. Nat. Paris 1: 112. 1792. Epidendrum incisum Vell., Fl. Flumin. 9, t. 18. 1825 [1829]. Epidendrum schomburgkii Lindl., Edwards’s Bot. Reg. 24: misc. 15. 1838. —Epidendrum ibaguense var. schomburgkii (Lindl.) C. Schweinf., Bot. Mus. Leafl. 11: 235. 1944. Epiphyte, medium-sized to usually large, erect, growing in ant nests, sun-loving; peduncles bearing a terminal subumbel of large, showy orange-red, red, or orange flowers. Humid forests, frequently riparian forests, 100–700(–1000) m; Bolívar (Río Caroní, Río Caura, Río Cuyuní), Amazonas (wide-

spread). Venezuelan Coastal Range (Aragua, Miranda), Andes (Táchira); Colombia, Guyana, Suriname, Ecuador, Peru, Brazil. Material from the Venezuelan Coastal Range, previously referred to this concept, belongs in Epidendrum mimeticum Carnevali & G.A. Romero. There seems to be at least two entities within a broad, commonly used concept of E. macrocarpum, easily distinguishable by the size of the flowers. One, with sepals 15–20 mm long and flowers tending to be reddish, apparently fits the E. macrocarpum concept; this entity seems to be restricted to the Guianas and the northeastern Amazon basin. Larger flowered (sepals 25–35 mm long) bright orange-red or orange forms are referable to the Peruvian E. splendens Schltr. and are more common in Amazonian Ecuador, Peru, and Bolivia. Epidendrum magnicallosum C. Schweinf., Bot. Mus. Leafl. 11: 96. 1943. Epiphyte, pendent or horizontally spreading, often the whole plant purple-tinged, shade-loving; flowers inconspicuous, resupinate, with widely spreading perianth segments, sepals white, petals and lip very pale green-white, all segments tinged with pale pink at apex, column pale green-white; dorsal sepal 8–12 mm long. White-sand scrub (bana), shrub savannas (caatinga), 50–200 m; Amazonas (near San Carlos de Río Negro, road between Yavita and Maroa). Amazonian Peru. Epidendrum micronocturnum Carnevali & G.A. Romero, Lindleyana 11: 241. 1996. Epiphyte, shade-loving; stems 6–9 cm long; leaves 35–65 mm long, 2–4.5 mm wide; flowers with pale green or pale yellowish green perianth and white lip; dorsal sepal 15–20 mm long. Rain forests, 300–900 m; Bolívar (El Paují, La Escalera to Cerro Venamo region, Río Aonda on Auyán-tepui). Brazil (Amazonas). Epidendrum micronocturnum looks like a depauperate, miniature E. nocturnum but occurs in populations. The plants remain small in cultivation. The narrow, more succulent leaves are also distinctive. Epidendrum microphyllum Lindl., J. Bot. (Hooker) 3: 85. 1841. —Lanium microphyllum (Lindl.) Benth., J. Bot. (Hooker) 3: 85. 1841.

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Epiphyte, creeping; stems 2–6 cm long; leaves 1–3 cm long; inflorescence a raceme or panicle; flowers nonresupinate with widely spreading perianth segments, greenish or greenish with purple tinges; dorsal sepal 4–8 mm long. Rain forests, Delta Amacuro (Sacupana), Bolívar (La Escalera to Cerro Venamo region, Roraima-tepui), Amazonas (Río Ventuari, Yavita–Maroa road). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil, Bolivia. ◆Fig. 309. Epidendrum miserrimum Rchb. f., Bonplandia (Hanover) 3: 220. 1855. Epiphyte, small to minute, erect, cespitose; inflorescence a lax-flowered raceme; flowers minute with subparallel green or green-purple-tinged perianth segments, conspicuous vesicle under the ovary. Lowland to cloud forests, 900–1400 m; Bolívar (Altiplanicie de Nuria, La Escalera to Cerro Venamo region, Macizo del Chimantá, Río Cuyuní basin), Amazonas (Cerro Avispa, Cerro Marahuaka, Sierra Parima). Sucre; Costa Rica, West Indies, Ecuador, Brazil. ◆Fig. 310. Epidendrum montigenum Ridl. ex Thurn, Timehri 5: 203. 1886. Terrestrial or lithophyte, medium-sized to large; stems erect, cane-like; inflorescences short, pendulous, or arched raceme or panicles; flowers rose. Open savanna-like tepui formations, 2400–2800 m; Bolívar (Ilú-tepui, Kukenán-tepui Macizo del Chimantá, Roraima-tepui). Guyana. Epidendrum montigenum is another of the more distinctive taxa of the E. dendrobioides complex. Epidendrum montigenum is easy to recognize by its pink flowers, a description of which is unfortunately often overlooked by collectors. Epidendrum myrmecophorum Barb. Rodr., Vellosia ed. 2, 1: 173. 1891. Epidendrum huebneri Schltr., Beih. Bot. Centralbl. 42(2): 99. 1925. Epidendrum rectopedunculatum C. Schweinf., Bot. Mus. Leafl. 11: 110. 1943. Epidendrum spilotum Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 144. 1976. Epiphyte, medium-sized (to 80 cm tall), cespitose, erect to pendulous, often shadeloving; inflorescences with a peduncle to 30 cm; flowers small to medium; sepals to 8 mm

long; perianth segments greenish with purple or brownish spots. Rain forests, 100– 900 m; Bolívar (Río Caura, San Ignacio de Yuruani), Amazonas (Río Cataniapo, Río Cuntinamo, near San Carlos de Río Negro). Venezuelan Andes foothills (Táchira); Amazonian Ecuador, Peru (Loreto), and Brazil (Amazonas). ◆Fig. 324. Epidendrum myrmecophorum is somewhat variable in lip shape and inflorescence length, but the variations of the different structures do not appear to be correlated. The original description based on the type of E. spilotum was inaccurate in that the lip looks much more tranversely oblong than it really is. Epidendrum nocturnum Jacq., Enum. Syst. Pl. 29. 1760. Terrestrial, lithophyte, or epiphyte, small to large, erect to subpendulous; inflorescence one-flowered; medium-sized to very large flowers, green, yellow, or white, often tinged with purple or red; lip white. Lowland rain forests to tepui cloud forests, savanna-like tepui formations or sandstone exposures, near sea level to 1000 m; widespread in lowlands of Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela in humid or semideciduous forests; U.S.A. (Florida), Mexico, Central America, West Indies, south to Brazil. Epidendrum nocturnum is one of the most common and widespread of the genus. It is usually epiphytic and flowers are large on a medium-sized plant (see key). Epidendrum norae Carnevali & G.A. Romero, Lindleyana 11: 245. 1996. Epiphyte 20–40 cm long; leaves 8–13 × 1– 2.5 cm; flowers light brownish green or totally green, usually brownish-tinged, lip white; dorsal sepal 19–25 mm long. Riparian forests, in a mosaic of Trachypogon savannas and forests, 50–200 m; Bolívar (Cabruta in Río Parguaza area), Amazonas (Caño Majagua tributary of Río Parucito, Río Yureba, Río Yutajé). Apure; probably in adjacent Colombia. Epidendrum norae is similar to and closely related to E. nocturnum, but the much smaller flowers on a relatively large plant are distinctive. Epidendrum norae usually grows in seasonally dryer places than E. nocturnum.

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Epidendrum nuriense Carnevali & Hágsater, Novon 2: 316. 1992. Epiphyte 7–40 cm tall, erect to arching, shade-loving; flowers nonresupinate, segments widely spreading, sepals and petals bronze-green, veins somewhat darker, lip shiny, bronze-colored with green calli, column green with pink anther; dorsal sepal 17–18 mm long. Cloud forests or rain forests, 500–700 m; Bolívar (Altiplanicie de Nuria, La Escalera to Cerro Venamo region). Endemic. Epidendrum orchidiflorum Salzm., Gen. Sp. Orch. 103. 1831. Epidendrum rectopedunculatum f. denticulatum C. Schweinf., Bot. Mus. Leafl. 20: 18. 1962. Epidendrum huebneri auct. non Schltr. 1925: sensu Dunst. & Garay, Venez. Orchid. Ill. 2: 127. 1961; Foldats in Lasser, Fl. Venez. 15(3): 273. 1970. Terrestrial or lithophyte, 100–250 cm tall, erect, often brownish purple-tinged, sun-loving; inflorescence very elongate, with a small, apical subumbel; flowers fleshy, shortlived, sepals widely spreading, petals subparallel to column; all perianth segments greenish with purple or brown dots. Open places such as sandstone outcrops, whitesand savanna-like formations, 50–1400 m; Bolívar (widespread), Amazonas (widespread). Colombia, Guyana, Suriname, French Guiana, Brazil (Amazon basin, Bahia). ◆Fig. 321. Epidendrum pachyphyton Garay, Orquideología 8: 182. 1973. Epiphyte, erect to pendulous; stems simple or branched; leaves short, fleshy; flowers small, opening well in short bracteate racemes; perianth segments and lip green or green purple-tinged. Cloud forests on tepui slopes or summits, 1400–1800 m; Bolívar (Auyán-tepui, Jaua-Sarisariñama massif, Kamarkawarai-tepui, La Escalera to Cerro Venamo region, Macizo del Chimantá), Amazonas (Cerro Aracamuni, Cerro Marahuaka. Cerro Sipapo). Colombia, Ecuador. Epidendrum prostratum (Lindl.) Cogn. in Mart., Fl. Bras. 3(5): 112. 1898. —Physinga prostrata Lindl., Edwards’s Bot. Reg. 24: misc. 45. 1838. Epiphyte 5–17 cm tall, shade-loving; inflorescences 5–10(–18)cm long; flowers sub-

campanulate, successive, lilac-rose, purplish, or greenish with reddish tinges; dorsal sepal 5–7 mm long. Rain forests, 100–300 m; Bolívar (between El Dorado and Kilómetro 88). Amazonian parts of Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia. Epidendrum pseudoramosum Schltr., Repert. Spec. Nov. Regni Veg. 10: 361. 1912. Epiphyte, small to medium-sized, branching like a small subfrutex; flowers small, green, yellow-green, or green-purple-tinged, not opening widely. Cloud forests, (600–)1000– 1800(–2500) m; Bolívar (Altiplanicie de Nuria, Cerro Venamo, Macizo del Chimantá, Ptari-tepui). Aragua, Miranda, Monagas, Sucre, Yaracuy; Guatemala, Honduras, Nicaragua, Costa Rica. Epidendrum purpurascens H. Focke, Tijdschr. Wis- Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 4: 64. 1851. Epidendrum clavatum Lindl., Edwards’s Bot. Reg. 22: t. 1870. 1836. Lithophyte or rarely an epiphyte, small to medium-sized, erect to ascendent, conspicuously rhizomatous; pseudobulbs stipitate, apically 1- or 2-foliate; inflorescences originating from the new growth, conspicuously bracteate, few-flowered racemes; flowers medium-sized, perianth segments spreading, green, greenish gray, or green-purple-tinged; lip white. Semideciduous or evergreen forests, shaded rocky outcrops; Bolívar (El Pao, Serranía de Imataca), Amazonas (Puerto Ayacucho, basin of Río Manapiare and Río Parucito). Sucre; Costa Rica, Colombia, Guyana, Suriname, Brazil (Pará). ◆Fig. 314. Epidendrum ramosum Jacq., Enum. Syst. Pl. 29. 1760. Epiphyte, small to medium-sized, with subfrutex-like branching; flowers small, green or yellowish green, open widely. Humid to cloud forests, (200–)600–1600(–2500) m; Bolívar (Altiplanicie de Nuria, Auyán-tepui, Cerro Venamo area, Jaua-Sarisariñama massif, Macizo del Chimantá, Roraimatepui, Serranía de Imataca), Amazonas (Río Siapa, Sierra Parima). Aragua, Falcón, Lara, Miranda, Táchira, Zulia; Mexico, Central America, West Indies, Ecuador, Peru, Brazil. Epidendrum ramosum is one of the most widespread and common of all neotropical orchids.

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O RCHIDACEAE

Epidendrum repens Cogn., Repert. Spec. Nov. Regni Veg. 7: 12. 1909. Epiphytes, small to medium-sized, pendent, prostrate or subcreeping, branched; flowers solitary, small, widely opening, green or yellowish green. Cloud forest on tepui slopes or summits, 1300–2800 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region, Macizo del Chimantá, Ptari-tepui, Roraima-tepui), Amazonas (Sierra de la Neblina). Aragua, Distrito Federal, Lara, Mérida, Portuguesa, Táchira; Mexico, Central America, West Indies, Colombia, Guyana, Ecuador, Brazil. Epidendrum rigidum Jacq., Enum. Syst. Pl. 29. 1760. —Spathiger rigidus (Jacq.) Small, Fl. Miami 55. 1913. Epiphyte, rarely a lithophyte, erect to horizontally spreading, creeping and sometimes covering large areas over the substrate; leaves 3–10 cm long, 4–20 mm wide; inflorescences a short raceme (3–8-flowered); flowers opening widely, rigid, all segments green or yellow-green; dorsal sepal 4.5–10 mm long. Rain forests, less commonly in semideciduous forests or cloud forests, 50– 1000(–1500) m; Delta Amacuro (Río Amacuro), Bolívar (throughout), Amazonas (Caño Manapiare, San Juan de Ucata south of Río Guayapo, Sierra Parima). Venezuelan Coastal Range, Barinas, Falcón, Portuguesa, Zulia; West Indies, tropical South America. Epidendrum rigidum is one of the most common and widespread of all neotropical orchids. Epidendrum saxatile Lindl., J. Bot. (Hooker) 3: 84. 1841. Epidendrum miersii Lindl., Fol. Orchid., Epidendrum 59. 1853. Epidendrum weddellii Lindl., Fol. Orchid., Epidendrum 67. 1853. Subterrestrial, lithophyte, or epiphyte; stems proliferous, sprawling, often creeping; flowers in large, lax panicles, white, yellow, pink, or purple, concolorous or with white perianth segments and a pink or purple lip. Low open forests to savanna-like tepui associations or cloud forests, 900–2000 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region, Roraima-tepui, Sororopán-tepui), Amazonas (Cerro Parú, Cerro Sipapo). Guyana, southern Brazil, Paraguay.

The presence of Epidendrum saxatile in the flora area is an interesting phytogeographical disjunction. Epidendrum schlimii Rchb. f., Linnaea 22: 838. 1850. Epiphyte, cespitose; stems 5–15 cm tall; leaves 15–45 mm long, 1.8–4 mm wide; inflorescences racemes or panicles; flowers small, green or green-purple-tinged. Cloud forests, 1000–1400 m; Bolívar (La Escalera to Cerro Venamo region). Distrito Federal, Lara, Mérida, Táchira; Colombia, Ecuador, Peru. Epidendrum sculptum Rchb. f., Bonplandia (Hanover) 2: 89. 1854. Huebneria yauaperyensis auct. non Schltr. 1925: sensu Hoehne, Icon. Orch. Bras., t. 99. 1949. Epiphyte, pendent or prostrate; leaves 25–55 mm long, 9–15 mm wide; flowers not spreading widely, usually 2 open at a time, greenish or greenish yellow; dorsal sepal 9– 15 mm long. Rain forests, 300–400 m; Delta Amacuro (east of Río Grande). Belize, Costa Rica, Panama, Colombia, Guyana, Suriname, Ecuador, Amazonian Brazil. Epidendrum secundum Jacq., Enum. Syst. Pl. 29. 1760. Epidendrum elongatum Jacq., Collectanea 3: 260. 1789 [1791]. Epidendrum ellipticum Graham ex Hook., Exot. Fl. 3: pl. 207. 1826. Epidendrum dichotomum C. Presl, Reliq. Haenk. 1: 101. 1827. Epidendrum crassifolium Lindl., Gen. Sp. Orchid. Pl. 107. 1831. Epidendrum xanthinum Lindl., Edwards’s Bot. Reg. 30: misc. 18. 1844. Epidendrum inconstans Ames, Bull. Torrey Bot. Club 58: 350. 1931. —Epidendrum lindenii Lindl., Edwards’s Bot. Reg. 31: misc. 48. 1845, non Lindl. 1843. Usually terrestrial or lithophyte, rarely epiphyte, 30–150(–250) cm tall, sun-loving; inflorescences erect; flowers nonresupinate, with widely spreading perianth segments, these white, pink, purple, red, yellow, orange, or combinations of these colors, callus usually yellow or white; dorsal sepal 6–17 mm long. Open vegetation, roadsides, or rocky outcrops, sometimes inside or at the edges of rain or cloud forests, 700–2600 m; Bolívar

Epidendrum 343

(widespread), Amazonas (widespread). Widespread elsewhere in Venezuela and the Neotropics. ◆Fig. 326. Epidendrum secundum is related to E. ibaguense and associated taxa, but in the E. secundum complex the flowers are nonresupinate and the callus is composed of several, complexly arranged tubercles. The perianth segments are usually broader and shorter than in the former group of taxa. As treated here, this concept probably includes several species that can be distinguished by various combinations of color, flower size, and shape of the callus and midlobe of the lip. There are several distinctive entities in the Venezuelan Guayana, one with small, mostly pink, flowers and another fairly common entity with larger yellow flowers. A population from Sierra de la Neblina has a small lip and a relatively long column, resembling the Andean E. catillus Linden & Rchb. f., but with smaller flowers. Another group of populations from Roraima-tepui and Auyán-tepui has flowers with a simplified form of lip, but the group grades locally into forms with more typical, complex calli on the lip. The wide geographical and ecological range of the complex, and the subtle degree of variation of these characters among and between the countless populations, renders impractical the recognition of the segregates until a comprehensive revision of the group is completed. The synonymy here is not at all inclusive, but indicates some of the oldest names, some of the concepts that at one time or another have been accepted as distinct species, or that have been reported from Venezuela. Hybrids of one of the members of this complex and E. ibaguense have been collected in the flora area along the northern rim of the Gran Sabana area. Epidendrum sertorum Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 5: 102. 1972. Epiphyte, erect or ascending; leaves 55– 80 mm long, 12–18 mm wide; inflorescence umbellate; flowers light greenish yellow; dorsal sepal 15–17 mm long. Rain forests, 100– 200 m; Amazonas (Río Orinoco at Munduapo). Amazonian Colombia. Epidendrum sertorum is a member of the E. difforme complex. It is easily distin-

guished in the Venezuelan Guayana by its simple, ovate lip with raised veins. Epidendrum smaragdinum Lindl., Edwards’s Bot. Reg. 24: misc. 32. 1838. Epiphyte, usually growing in ant nests, or terrestrial; stems straight, 23–60 cm long, erect to subpendent; leaves 45–65 mm long, 5–12 mm wide; flowers usually spotless green, open one at a time; dorsal sepal 6–7 mm long. Rain forests, 100–400 m; Bolívar (Cerro Jaua), Amazonas (Brazo Casiquiare, Río Cataniapo, near San Carlos de Río Negro). Zulia; Guyana, Ecuador, Peru, Brazil (Amazonian states, Bahia, Mato Grosso), Bolivia. ◆Fig. 322. The Central American entity from Mexico and Guatemala, described as Epidendrum pachyrrhachys Ames, is closely related to E. smaragdinum, but the leaves are broader and the lip is transversely oblong. Epidendrum splendens Schltr., Repert. Spec. Nov. Regni. Veg. Beih. 9: 93. 1921. Epiphyte, growing in ant nests, very similar and closely related to Epidendrum macrocarpum; peduncles bearing a terminal subumbel of large, showy bright orange-red, red, or orange flowers; sepals 25–35 mm long. Rainforests, 500–1500 m; Bolívar (Auyán-tepui). Amazonian Colombia, Ecuador, Peru, and Bolivia. See comments under Epidendrum macrocarpum. Epidendrum stalkyi Carnevali & G.A. Romero in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1134. 2000. Epiphyte with small pseudobulbs on a creeping rhizome; flowers white; lip basally cordate or subcordate, with some pinkish flecking; dorsal sepal 3–4 mm long. Riparian shrublands on tepui summits, 1700–1800 m; Bolívar (Cerro Jaua). Endemic. ◆Fig. 311. Epidendrum manarae Foldats is the closest relative to E. stalkyi. It is known from the Venezuelan Coastal Range and is easily distinguished by its basally cuneate lip. Epidendrum stamfordianum Bateman, Orchid. Mexico Guatemala t. 11. 1838. Epiphyte or lithophyte; pseudobulbs 10– 30 cm long; leaves 10–27 cm long; flowers with widely spreading segments, pale yellow

344

O RCHIDACEAE

or greenish with red or purple dots; dorsal sepal 12–25 mm long. Deciduous or rain forests, or over granitic outcrops (lajas), 100– 200 m; Bolívar (Río Parguaza, near Tumeremo), Amazonas (near Puerto Ayacucho, Río Sipapo). Aragua, Carabobo, Distrito Federal, Guárico, Lara, Miranda, Yaracuy; Mexico, Central America, Colombia. Epidendrum strobiliferum Rchb. f., Ned. Kruidk. Arch. 4: 333. 1858. Epiphyte to 30 cm long; leaves 8–50 × 2– 10 mm; flowers not widely spreading, greenish, cream, or white; dorsal sepal 2.5–5.6 mm long. Rain forests, rarely in cloud forests, near sea level to 1400 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela in humid forests; widespread in the tropics and subtropics of the New World. Epidendrum strobiliferum is often overlooked by collectors because of its small vegetative size and inconspicuous flowers. Epidendrum strobiloides Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 4: 104. 1966. Epiphyte, erect or arching; leaves 5–15 mm long, 2.5–4 mm wide; flowers greenish, not widely spreading and almost completely enclosed by the floral bracts; dorsal sepal 1.8–2 mm long, 1–1.3 mm wide. Rain forests, 100–1000 m; Bolívar (northeasternmost part). Guyana, Suriname, French Guiana, Ecuador. Epidendrum tumuc-humaciense (Veyret) Carnevali & G.A. Romero, Lindleyana 11: 246. 1996. —Epidendrum nocturnum var. tumuc-humacience Veyret, Adansonia 3/4: 186. 1982. Terrestrial, lithophyte, or very rarely a low epiphyte, 50–150(–200) cm tall, stiffly erect, sun-loving; leaves 5–7 × 1.8–2.7 cm; pedicel 5–11 cm long, much surpassing the final leaves of the stem, flowers with white lip, petals and sepals yellowish suffused with purplish; dorsal sepal 4.5–6.5 cm long. Locally common in rocky outcrops, particularly over sandstone, 700–2300 m; Bolívar (widespread), Amazonas (widespread). Amazonian Colombia (Vaupés), Guyana, Suriname,

French Guiana, Brazil (Amazonas, Roraimatepui). Epidendrum tumuc-humaciense is easy to recognize among the Guayanan members of the E. nocturnum complex by its stiffly erect, sun-loving habit, its preference for rocky outcrops, and its large, long-pedicellate flowers. Epidendrum ulei Schltr., Notizbl. Königl. Bot. Gart. Berlin 6: 124. 1914. Epidendrum durum auct. non Lindl.: sensu Foldats in Lasser, Fl. Venez. 15(3): 240, fig. 466. 1970., pro parte. Epidendrum dendrobioides auct. non Thunb. 1818: sensu Dunst. & Garay, Orchids Venez. Ill. Field Guide 220. 1979. Terrestrial or lithophyte, 50–150 cm tall; flowers green or greenish yellow; dorsal sepal 8–9 mm long; petals 7.5–8 mm long, 2 mm wide; lip 5–5.5 mm long, 7.5–9 mm wide. Rocky outcrops, sandy or boggy soils, (1000–) 1500–2700 m; Bolívar (widespread), Amazonas (widespread). Guyana, Brazil (Amazonas, Roraima, Rio Branco). ◆Fig. 317. Most of the populations of the Epidendrum dendrobioides-durum complex from intermediate or higher elevations (1500–2500 m) have been assigned to this species. Epidendrum unguiculatum (C. Schweinf.) Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 38. 1976. —Epidendrum paniculatum var. unguiculatum C. Schweinf., Bot. Mus. Leafl. 11: 107. 1943. Epidendrum fastigiatum auct. non. Lindl. 1853: sensu Dunst. & Garay, Venez. Orchid. Ill. 2: 119. 1976. Epidendrum paniculatum auct. non Ruiz & Pav. 1798: sensu Foldats in Lasser, Fl. Venez. 15(3): 332. 1970, pro parte. Lithophyte or epiphyte to 1 m tall; leaves 10–14 × 3–4.5 cm; flowers with widely spreading perianth segments, these deep green, lip white basally, green apically and at the lateral lobes, calli and periferal area purple; dorsal sepal 12–15 mm long. Rain forests, 100–300 m; Amazonas (Raudal Monserrat). Falcón, Mérida, Miranda, Monagas, Táchira; Colombia, French Guiana, Ecuador, Peru, Brazil (Amazonas). Epidendrum unguiculatum is one of two members of the E. paniculatum complex in

Epidendrum 345

the flora area; the other is E. densiflorum. Epidendrum unguiculatum can easily be recognized by the claw that separates the two divergent apical lobes from the main body of the lip. Epidendrum urbanianum Cogn. in Urb., Symb. Antill. 6: 531. 1910. Terrestrial to 2.6 m tall; leaves 7–9 × 2.5– 2.9 cm; inflorescence erect; flowers olive green to pale yellow, lip olive green above, pale green below; dorsal sepal 9.5–10 mm long, 4.5 mm wide. Cloud forests dominated by Bonnetia, 2000–2200 m; Bolívar (Macizo del Chimantá), Amazonas (Cerro Yaví). Guadalupe, Martinique. Epidendrum urbanianum is a member of the E. dendrobioides-durum complex. It is easily recognized by the very long (8–10 mm) floral bracts that are 3–4 times longer than the pedicellate ovary. Epidendrum urichianum Carnevali, Foldats & I. Ramírez, Orquídea (Mexico) 12: 151. 1992. Epiphyte, branching, erect, prostrate to subpendent; inflorescences terminal at the apex of main secondary stem or lateral branches; leaves 3.8–4.7 cm long; flowers pale green or suffused with pale lavenderbuff, tip of column creamy white; perianth segments fleshy, sepals not spreading, petal reflexed; dorsal sepal 7–7.5 mm long. Cloud forests, 1400–2000 m; Bolívar (Cerro Venamo, Ptari-tepui), Amazonas (Cerro Yutajé). Adjacent Guyana. ◆Fig. 328. Epidendrum urichianum is closely related to the widespread E. ramosum that tends to grow at lower elevations. In E. urichianum the perianth segments, including the lip, are broader and obtuse, and the lateral sepals have broad, high, wing-like keels on the dorsal side. Epidendrum vincentinum Lindl., J. Bot. (Hooker) 3: 88. 1840. —Epidendropsis vincentina (Lindl.) Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 114. 1976. Epidendrum serricardium Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19: 218. 1923. Epiphyte 5–10 cm tall, often growing on thin branches in the margin of the forests,

arching to ascendent; leaves 15–30 × 3–4 mm, often purple-tinged; inflorescences purple-tinged, 4–12 cm long, with 2–5 branches; flowers pink-purple to greenish purple or brownish, petals often hyaline; dorsal sepal 3–4 mm long. Rain or cloud forests, 600–1300 m; Bolívar (Altiplanicie de Nuria, La Escalera to Cerro Venamo region), Amazonas (Cerro Aracamuni, Cerro Duida). Aragua; Costa Rica, Panama, West Indies, Colombia, Ecuador, Peru. Epidendrum vincentinum is a common, widespread little species, often overlooked by collectors. Its distribution within Venezuelan borders is probably broader than the collection record indicates. Epidendrum violascens Ridl. ex Thurn, Timehri 5: 203. 1886. —Epidendropsis violascens (Ridl. ex Thurn) Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 39. 1976. Lithophyte or rarely an erect epiphyte 3– 15 cm tall not including the inflorescence; leaves usually tinged purple, 9–21 × 3–5 mm; flowers nonresupinate, with widely spreading perianth segments, these red-purple, column and lip yellowish; dorsal sepal 3.9–6 mm long. Locally common in rocky outcrops and tepui meadows, 1500–2600 m; Bolívar (widespread on tepui tops), Amazonas (Cerro Sipapo, Cerro Yutajé). Adjacent Guyana, Brazil (Roraima-tepui). ◆Fig. 319. Epidendrum violascens is closely related to E. vincentinum, but E. violascens is more robust and more succulent and has longer inflorescences and narrower lateral sepals. While E. vincentinum always grows as an epiphyte in cloud forests, E. violascens is restricted to tepui shrublands and meadows, where it is terrestrial or a lithophyte. Epidendrum sp. A Terrestrial, prostrate; leaves 3.5–5 × 0.5–1 cm; inflorescences 4.5–7 cm long; flowers white or yellowish, dorsal sepal 7–8 mm long. Boggy conditions on tepui meadows, 2000– 2800 m; Bolívar (Ilú-tepui, Kukenán-tepui, Roraima-tepui). Guyana (Monte Roraima complex). This species is very similar to some forms currently referred to Epidendrum alsum, but is less robust and is prostrate, sprawling over the boggy substrate of tepui meadows.

346

O RCHIDACEAE

Fig. 309. Epidendrum microphyllum

Fig. 310. Epidendrum miserrimum

Fig. 311. Epidendrum stalkyi

Fig. 312. Epidendrum calanthum

Epidendrum 347

Fig. 313. Epidendrum dichaeoides

Fig. 314. Epidendrum purpurascens

348

O RCHIDACEAE

Fig. 315. Epidendrum cristatum

Fig. 316. Epidendrum klotzscheanum

Epidendrum 349

Fig. 317. Epidendrum ulei

Fig. 318. Epidendrum amazonicoriifolium

350

O RCHIDACEAE

Fig. 319. Epidendrum violascens

Fig. 320. Epidendrum carpophorum

Epidendrum 351

Fig. 321. Epidendrum orchidiflorum

Fig. 322. Epidendrum smaragdinum

Fig. 323. Epidendrum hombersleyi

Fig. 324. Epidendrum myrmecophorum

352

O RCHIDACEAE

Fig. 325. Epidendrum ibaguense

Fig. 326. Epidendrum secundum

Fig. 327. Epidendrum ibaguense × E. secundum

Fig. 328. Epidendrum urichianum

Epistephium 353

51. EPISTEPHIUM Kunth, Syn. Pl. 1: 340. 1822. [Subtribe Vanillinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial herbs, erect, sometimes climbing or prostrate. Roots many, fibrous or ± fleshy; rhizomes short, thick, subterranean. Stems ± woody, unbranched or branched in the apical 1/2, terete. Leaves not articulate, convolute, distichous or spiraled, rigid or coriaceous, in dried condition brown or blackish and shiny to matte; blades ovate to lanceolate, rarely suborbicular, sessile and basally cordateamplexicaul or narrowly petiolate; venation reticulate, usually conspicuous. Racemes terminal on the main stem or on the branches, rarely axillary, successively flowered and progressively lengthening, lax to ± dense, frequently purple-tinged; peduncle terete, elongate to almost absent; bracts tubular, usually widely spaced and leaving most of the peduncle naked; rachis straight or zigzag. Flowers resupinate, spiraled, fugacious, usually large and showy, mostly pink or purple, rarely white, bearing a conspicuous entire to dentate calyculus below the perianth segments; floral bracts fleshy, the basal one often foliaceous and then the basal flowers appearing axillary; pedicellate ovary elongate, with an abscission layer between ovary and perianth. Perianth segments membranous, rarely ± fleshy, spreading in anthesis; sepals free, subequal or the laterals ± oblique; petals usually similar to the sepals or broader. Lip clawed, the basal margins fused with the ventral face of the column, the blade unlobed or slightly lobed, broadly ovate to obovate, funnel-shaped and surrounding the column, margins wavy or ruffled; disk with 1–several ± hairy keels or series of transverse lamellae. Column elongate, semiterete, clavate; anther terminal or ± ventral, incumbent, operculate, 2-locular, articulate with clinandrium; clinandrium membranous, broadly 3-lobed; pollinia 4, soft and mealy; stigma entire, emergent, viscidium absent. Capsules 1- or 3-locular, oblong, rigid. Seeds small, prominently winged, seed coat hard. Central America, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina; ca. 25 species, 8 in Venezuela, 7 of these in the flora area. Epistephium is poorly understood because flowers in most herbarium specimens are difficult to re-hydrate and examine. Additional species will probably be detected in the flora area when a comprehensive revision of the genus is conducted. Key to the Species of Epistephium 1. 1. 2(1). 2. 3(2).

3.

Leaves petiolate or with pseudopetiole ..................................................... 2 Leaves sessile ............................................................................................. 5 Plants to 40 cm tall; leaves membranous to subcoriaceous ....... E. ellipticum Plants > 50 cm tall; leaves rigid-coriaceous to subcoriaceous .................. 3 Leaves yellowish to pale brown when dry, always pseudopetiolate, 1.5– 2.5 times longer than wide on mature basal leaves; lip disk longbarbate at apex, short-barbate at middle, glabrous at base; dorsal sepal 4–5.5 cm long .................................................................... E. sclerophyllum Leaves deep brown to blackish when dry, always conspicuously petiolate,

354

4(3).

4.

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

O RCHIDACEAE

3.5–7 times longer than wide on mature basal leaves; lip disk longbarbate at middle and apex; dorsal sepal 5.5–7 cm long ..................... 4 Inflorescence only terminal, rarely 1 axillary near the top of the stem at the same time, (3–)5–13 times longer than the subtending leaf, manyflowered; plants unbranched or rarely with 1–few apical branches; leaves coriaceous; petals broadly obovate ............................. E. subrepens Inflorescences terminal and axillary (rarely only terminal), 1.5–4 times longer than subtending leaf, generally few-flowered, plants usually several- to many-branched in apical 1/2; leaves subcoriaceous; petals spatulate .............................................................................. E. parviflorum Leaves coriaceous, yellowish or pale brown when dry, 1.5–2.5 times longer than wide ............................................................... E. sclerophyllum Leaves subcoriaceous to membranous, blackish when dry, usually > 3 times longer than wide .................................................................... 6 Lip unlobed .................................................................................. E. hernandii Lip slightly but distinctly 3-lobed .............................................................. 7 Lip emarginate, apical margin undulate; disk 3-keeled, hairs only covering the central area ...................................................................... E. duckei Lip at apex conspicuously 2-lobed; disk of lip totally covered with hairs that are much shorter toward apex ............................................ E. elatum

Epistephium duckei Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 7: 287. 1913. Plant 1–2.5(–4) m tall, erect, ± prostrate in forests; leaves shiny; flowers pale purple to deep purple, lip darker and with whitish or yellowish central area with dark purple veins. Common in open, rocky places or sometimes in semiaquatic areas, (400–)700– 1500 m; Bolívar (widespread but more common in Río Caroní basin), Amazonas (Sierra Parima). Guyana, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 329. Epistephium elatum H.B.K., Nov. Gen. Sp. (quarto ed.) 7: 159. 1825. Plant 1–2.5 m tall, erect or ± prostrate; leaves shiny; flowers purple-rose with a darker purple-striped lip. Open, humid places, 1100–1300 m; Bolívar (La Escalera to Cerro Venamo region, Río Uarama). Táchira, Trujillo; Colombia, Ecuador. The flora area material is here considered Epistephium elatum with some reservation, because we are uncertain if it is conspecific with the Andean populations. Epistephium ellipticum L.O. Williams & Summerh., Bull. Misc. Inform. Kew 1928: 145. 1928.

Plant 15–40 cm tall, erect, usually growing in shady, humid places; sepals 2–3 cm long, petals and sepals white with lilac or purple midvein, rarely entirely pale lilac; lip lilac or purple; column white. Rain or cloud forests, 100–1100(–1700) m; Bolívar (Macizo del Chimantá, Uonquén), Amazonas (30 km north of Puerto Ayacucho, Rincones de Chacorro, near Yavita). Trinidad-Tobago, Amazonian Brazil. Epistephium hernandii Garay, Amer. Orchid Soc. Bull. 30: 498. 1961. Plant 1.5–3.5 m tall, erect; leaves shiny; flowers purple; lip deep purple with a white throat, with deeper purple veins. Open rocky places, tepui meadows, 800–1700 m; Amazonas (Cerro Aracamuni, Cerro Aratitiyope, Cerro Autana, Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Sipapo). Amazonian Colombia. Epistephium parviflorum Lindl., Gen. Sp. Orchid. Pl. 433. 1840. Epistephium petiolatum Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 7: 288. 1910. Plant (0.5–)1–2(–4) m tall, erect when growing in open places to subprostrate when growing in forests; flowers pink to purple; lip

Epistephium 355

darker with white lamellae. Rain forests or open, humid places, 100–800(–1400) m; Bolívar (Cerro Guaiquinima, Cerro Pitón, Río Chibau in Río Chicanán basin, Serranía Caráruban near Guarimba, Serranía Marutaní), Amazonas (widespread but more abundant in the southern half). Amazonian Colombia, Guyana, Peru, Amazonian Brazil. ◆Fig. 331. Epistephium sclerophyllum Lindl., Gen. Sp. Orchid. Pl. 433. 1840. Plant 1–2 m tall, erect; leaves drying matte brown, rigid; flowers magenta or reddish purple; lip with yellow in center, sometimes white with purple veins. Open places in rain forests, 100–200(–900) m; Amazonas (Brazo Casiquiare, Cerro Yapacana, Río Pasimoni, Río Yatúa). Brazil, Bolivia, Paraguay. Epistephium subrepens Hoehne, Arq. Bot. Estado São Paulo n.s. 1, 6: 127, t. 143. 1944. Epistephium cruegeri Rchb. f. ex Griseb., Fl. Brit. W. I. 637. 1864, nom. nud. Plant 1–2 m tall, erect; flowers pale purple to magenta; lip darker apically. Open places, usually on sandstone outcrops, 100– 1500 m; Bolívar (widespread), Amazonas (widespread). Amazonian Colombia, TrinidadTobago, Guyana, Amazonian Brazil. ◆Fig. 330. Epistephium subrepens tends to grow at higher elevations and in more open places than the closely related E. parviflorum.

Fig. 329. Epistephium duckei

356

O RCHIDACEAE

Fig. 330. Epistephium subrepens

Fig. 331. Epistephium parviflorum

Eriopsis 357

52. ERIOPSIS Lindl., Edwards’s Bot. Reg. 33: sub t. 9. 1847. [Subtribe Cyrtopodiinae]. Pseudoeriopsis Rchb. f., Linnaea 22: 852. 1849. by Gustavo A. Romero-González Cespitose epiphytes, terrestrial, or sometimes lithophytic herbs. Stem modified into aggregate pseudobulbs, pear-shaped to elongate, cylindric, of several internodes, the basal-most internode elongate, the apical ones short to inconspicuous, when young entirely concealed by distichous, scarious sheaths. Leaves 2–4, coriaceous, narrowly elliptic or oblong-ovate to narrowly obovate, acute to acuminate, plicate, subpetiolate, articulate. Inflorescences lateral, arising from the base of the pseudobulbs, erect, racemose, few- to many-flowered, usually longer than the leaf. Flowers resupinate, showy; floral bracts linear-ovate, much shorter than the pedicellate ovary, often inconspicuous. Sepals spreading, fleshy, concave, oblong-elliptic to broadly elliptic, obtuse, the lateral ones often slightly oblique, basally connate, the dorsal one often mucronate; petals membranous, spreading, somewhat concave, oblong to oblong-elliptic, sometimes slightly oblique. Lip sessile to shortly clawed, articulate with the column foot, concave, conspicuously 3-lobed near the apex, lateral lobes broadly semicircular to semiovate, midlobe smaller, sessile to clawed, reniform to transversely oblong, apically obtuse, retuse to 2-lobed; disk at the base lamellate and/or verrucose. Column erect, slightly arched, semiterete to trigonous, clavate, with a short foot; anther terminal, operculate, incumbent, 1-locular; pollinia 2, yellow, cartilaginous, sulcate, ovate, semiterete in cross section, attached to a short, trullate tegula. Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 4 species, 3 in Venezuela, all in the flora area. Key to the Species of Eriopsis 1.

1.

2(1).

2.

Plants with fine, erect secondary roots, often concealing the pseudobulbs; epiphytic, most often in ant gardens, rarely terrestrial; callus of the lip with a pair of erect, parallel, plate-like, falcate to lanceolate horns ................................................................................................... E. sceptrum Plants without secondary roots; terrestrial, lithophytic, or epiphytic, if epiphytic never in ant gardens; callus of the lip with 1 or 2 pairs of erect, parallel, plate-like, triangular horns and a pair of parallel, verrucose processes extending toward the isthmus .................................. 2 Plants terrestrial or lithophytic, callus of the lip with 1 pair of erect, parallel, plate-like, triangular horns and a pair of parallel, verrucose processes extending toward but never reaching the isthmus .......... E. biloba Plants epiphytic or terrestrial, callus of the lip with 2 pairs of erect, parallel, plate-like, triangular horns and a pair of parallel, verrucose processes extending toward and always reaching the isthmus .................. ............................................................................................ E. rutidobulbon

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Fig. 332. Eriopsis biloba

Eriopsis biloba Lindl., Edwards’s Bot. Reg. 33: sub t. 9. 1847. Pseudoeriopsis schomburgkii Rchb. f., Linnaea 22: 853. 1849. Eriopsis grandibulbosa Ames & C. Schweinf., Bull. Torrey Bot. Club 58: 350. 1931. Variable terrestrial or lithophyte; pseudobulbs generally 3-foliate; inflorescence erect; flowers brownish yellow; sepals and petals yellowish brown, or magenta basally and yellow apically; lip base and disk brownish yellow or brownish gold, the lip midlobe white with brown spots; column yellow. Sandy savannas, open dwarf forests on sandy soils, sandy and rocky savannas on tepui summits, 100–2300 m; Bolívar (Gran Sabana), Amazonas (base of Cerro Autana and Cerro Sipapo, near Maroa). Colombia, Guyana, Ecuador, Peru, Brazil (Amazonas). ◆Fig. 332. Eriopsis rutidobulbon Hook., Bot. Mag. 75: t. 4437. 1849. Eriopsis colombiana Schltr., Repert. Spec. Nov. Regni Veg. Beih. 27: 172. 1924. Epiphyte or rarely terrestrial; pseudo-

bulbs generally 3-foliate; inflorescence erect; sepals orange-yellow apically, dull magenta in the middle suffused with yellow, greenish yellow basally; petals orange-yellow apically, magenta toward the base; lip base brownred, creamy white spotted with purple toward the apex; column orange-yellow apically, brown-red basally. Wet forests, 1000–1200 m; Bolívar (La Escalera). Táchira; Colombia, Ecuador. Eriopsis sceptrum Rchb. f., Bonplandia (Hanover) 2: 98. 1854. Eriopsis sprucei Rchb. f. in Walp., Ann. Bot. Syst. 6: 663. 1863. Epiphyte, often in large clumps, rarely terrestrial; pseudobulbs generally 3-foliate; inflorescence erect; flowers yellowish brown; sepals and petals yellow with brownish maroon margins; lip base and blade yellowish cream, the midlobe white with deep purple spots; column yellow. Wet forests bordering rivers, 100–600(–1800) m; Bolívar (Río Acanán, Río Icabarú), Amazonas (Río Atacavi, Río Autana, Río Sipapo, Sierra de la Neblina). Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Amazonas).

Erycina 359

53. ERYCINA Lindl., Fol. Orchid. Erycina 2. 1853. [Subtribe Oncidiinae]. Oncidium sect. Aphanobulbia subsect. Iridifolia Kraenzl. in Engl., Pflanzenr. IV. 50(Heft 80): 96. 1922. —Oncidium sect. Iridifolia (Kraenzl.) Garay, Taxon 19: 449. 1970. Psygmorchis Dodson & Dressler, Phytologia 24: 288. 1972. by Germán Carnevali and Ivón M. Ramírez-Morillo Twig epiphytes, small, cespitose, often short-lived. Rhizomes absent or very short. Pseudobulbs absent or present; if present, apically leafless (or bearing an abortive, scale-like leaf) and basally enveloped by 1 or 2 sheaths bearing leaf blades, or topped by 1 or 2 leaves; if pseudobulbs absent, then leaves laterally compressed, and plants psygmoid (fan-like with laterally compressed leaves). Leaves and the blades of the leaf sheaths dorsoventrally flattened and articulate with their sheaths (in pseudobulbous taxa), or laterally conduplicate and not articulate to their sheaths (in pseudobulbless taxa), flattened, when laterally conduplicate arranged in the manner of a fan and dorsally ancipitous (flattened with sharp edges), obliquely narrow-elliptic to narrowly oblong-elliptic, acute to obtuse, subfleshy to thinly coriaceous, in one species red-purple-striated. Inflorescences lateral, originating from the axils of the leaf sheaths, shorter, ± equal to much longer than leaves; peduncle subterete to ancipitous, few-bracted, successively 1–5-flowered or a simultaneously few- to multiflowered raceme or panicle. Flowers medium-sized to relatively large but always large for plant size, short to long-lasting, resupinate; floral bracts concave, elliptic to ovate-elliptic, or inconspicuous, acute to obtuse; pedicellate ovary slender, clavate, ridged. Perianth segments widely spreading, sepals yellow, small with respect to petals and lip, basally connate; dorsal sepal larger, forming a hood over column; lateral sepals completely hidden by the lateral lobes of lip; petals completely yellow or with red-purple or maroon blotches or streaks, usually similar to dorsal sepal, free. Lip completely yellow or with red, purple, or maroon blotches or markings at base, rigidly attached to column base, much larger than other perianth segments, flat or somewhat concave or convex, often ± pandurate, with 2 variously developed basal lateral lobes and a 2–4-lobed central lobe separated from basal lobes by a variously developed isthmus, the taxa with psygmoid habit have much larger midlobes of the lip compared to the lateral lobes; callosities on lip base and above the isthmus usually formed by a ± flattened, horizontal blade with entire to fimbriate apical edges, usually of a clearer hue than rest of lip. Column erect to ascendent, rather short, apically with well-developed wings that have erose or fimbriate margins and form a hood over the clinandrium and stigma; anther terminal, incumbent, operculate, shortly rostrate; pollinia 2, globose or somewhat clavate, with a long tegula and a small, flattened viscidium; stigma ventral. Capsule ellipsoid, relatively large, ridged. Tropical Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 9 or 10 species, 2 in Venezuela, both in the flora area. This genus is comprised of two sharply distinct clades, one entirely Mexican, the other spanning the whole distributional range of the genus. The Mexican clade includes two species and features pseudobulbous plants and dorsoventrally compressed leaves that are articulated with their sheaths. This clade comprises the type of the genus, Erycina echinata (Kunth) Lindl. The second clade comprises twig epi-

360

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phytic, psygmoid plants that were until recently referred to as the genus Psygmorchis. The relationships of these two morphologically distinct clades were not hinted at until recently with DNA sequence studies. Key to the Species of Erycina 1. 1.

Peduncle apically terete; callus of the lip digitate-fimbriate ..................... ............................................................................................ E. glossomystax Peduncle and rachis flattened; callus with entire margins ........... E. pusilla

Erycina glossomystax (Rchb. f.) N.H. Williams & M.W. Chase, Lindleyana 16(2): 136. 2001. —Oncidium glossomystax Rchb. f., Bot. Zeitung (Berlin) 10: 696. 1852. —Psygmorchis glossomystax (Rchb. f.) Dodson & Dressler, Phytologia 24: 289. 1972. Twig epiphyte, erect to pendulous, sun- or shade-loving; flowers widely open, lasting 3 or 4 days; perianth segments yellow, lip yellow with some purple blotches around callus; callus white or clear yellow. Rain forests, 50– 500 m; Bolívar (basins of Río Caroní and Río Caura), Amazonas (basins of Río Cataniapo and Río Gavilán). Anzoátegui, Aragua, Carabobo, Lara, Yaracuy; southern Mexico, Guatemala, French Guiana, Ecuador, Peru, Brazil. Erycina pusilla (L.) N.H. Williams & M.W. Chase, Lindleyana 16(2): 136. 2001. —Epidendrum pusillum L., Sp. Pl. ed. 2, 2: 1352. 1763. —Psygmorchis pusilla (L.) Dodson & Dressler, Phytologia 24: 288. 1972. —Oncidium pusillum (L.) Rchb. f. in Walp., Ann. Bot. Syst. 6: 714. 1863.

Twig epiphyte, cespitose, erect to pendulous, shade- or sun-loving; inflorescences shorter to slightly longer than leaves; flowers widely open, sepals greenish yellow, petals and lip bright yellow, sometimes with purple or maroon blotches on petal base. Rain forests, near sea level to 700 m; Delta Amacuro (Caño Atoiba, Río Amacuro), Bolívar (Río Cuyuní and Río Paragua basins), Amazonas (Brazo Casiquiare, Río Yatúa. Anzoátegui, Aragua, Barinas, Carabobo, Lara, Miranda, Monagas, Sucre, Yaracuy, Zulia; southern Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 333.

Fig. 333. Erycina pusilla

54. ERYTHRODES Blume, Bijdr. 410, t. 72. 1825. [Subtribe Goodyerinae]. Physurus Rich., Mém. Mus. Hist. Nat. 4: 55. 1818, nom. nud. Microchilus C. Presl, Reliq. Haenk. 1: 94. 1827. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial, humicolous, rarely ± epiphytic, creeping herbs, small to mediumsized, usually shade-loving. Stems terete, basally decumbent, rooting at internodes, foliose in the upper 1/3–1/2. Leaves convolute, ± fleshy, usually elliptic, acute, acuminate to obtuse, usually petiolate, with a funnel-shaped sheath. Inflorescence a spike, terminal, with a ± well-developed peduncle, few- to many-flowered, usually glandular-pubescent. Flowers small, generally of rather thin texture, resupinate; floral bracts usually elliptic to ovate-elliptic; sepals free, similar; petals connivent with dorsal sepal to form a 3-lobed hood or galea, lip without a pair of fleshy plates or ridges at base leading to the entrance to the spur, with a cylindric to sacciform spur at lip base, the apical lobes perpendicular to lip axis or reflexed. Column very short,

Erythrodes 361

sessile or subsessile, without distinct stalk; anther dorsal, erect; pollinia 4, sectile, caudicles attached to a well-defined viscidium, approximate, horizontal; rostellum neither articulate nor triggered, deeply 2-parted with clearly defined, elongate, tapering segments after viscidium has been removed; stigmas 2. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, northern Argentina, tropical Asia, Malaysia, Oceania; ca. 60 species, 8 in Venezuela, 2 of these in the flora area. Aspidogyne Garay and Ligeophila Garay are recent segregates from Erythrodes. Key to the Species of Erythrodes 1. 1.

Blade of lip abruptly dilated below its middle; leaves narrowly elliptic, lateral sepals ca. 1 mm wide ...................................................... E. arietina Blade of lip not dilated below its middle; leaves elliptic to widely elliptic; lateral sepals 1.7–2.3 mm wide ................................................. E. paleacea

Erythrodes arietina (Rchb. f. & Warm.) Ames, Orchidaceae 7: 66. 1922. —Physurus arietinus Rchb. f. & Warm. in Rchb. f., Otia. Bot. Hamburg. 2: 52. 1881. Small to medium-sized herb; leaves sometimes white-variegated; inflorescences longer than leaves; flowers small, white, with ± parallel perianth segments. Rain or cloud forests, 400–1500 m; Bolívar (upper Río Caroní basin), Amazonas (Cerro Huachamacari). Ecuador, Amazonian Brazil, Bolivia, northern Argentina. ◆Fig. 334. Erythrodes paleacea (Schltr.) Ames, Orchidaceae 7: 75. 1922. —Physurus paleaceus Schltr., Repert. Spec. Nov. Regni Veg. 7: 72. 1920. Medium-sized to relatively large herb, shade-loving, growing in humid places; inflorescence spiciform, erect, much longer than upper leaves; flowers small with perianth segments ± parallel to column; sepals greenish yellow or green; lip and petals pure white or whitish with a greenish spur. Rain or cloud forests, 600–1600 m; Bolívar (La Escalera to Cerro Venamo region, Macizo del Chimantá), Amazonas (Sierra de la Neblina). Aragua, Carabobo, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Sucre, Yaracuy, Zulia; Hispaniola, Trinidad-Tobago, Colombia, Guyana, Ecuador.

Fig. 334. Erythrodes arietina

362

O RCHIDACEAE

55. EULOPHIA R. Br., Bot. Reg. 7: t. 573 (“578”). Oct. 1821, nom. cons., “Eulophus,” ort. cons. [Subtribe Cyrtopodiinae]. Cyrtopera Lindl., Gen. Sp. Orchid. Pl. 189. 1833. by Gustavo A. Romero-González Cespitose, terrestrial herbs or rarely epiphytes. Stem modified into corm-like pseudobulbs, subterranean or above-ground, entirely concealed by leaf sheaths. Leaves 3–5, distichous, imbricating near the base, coriaceous, elongate, linear-elliptic to narrowly ovate, acuminate, plicate, subpetiolate, not articulate with the leaf sheaths, usually caducous. Inflorescences lateral, arising from the base of the pseudobulbs, erect, racemose or rarely paniculate, many-flowered, much longer than the leaf. Flowers resupinate, often showy; floral bracts narrowly ovate to filiform, much shorter than the pedicellate ovary, often inconspicuous. Sepals erectspreading, very narrowly elliptic to narrowly obovate, acuminate to apiculate, the lateral ones often slightly oblique, adnate to the column foot; petals spreading, broader and shorter to longer than the sepals. Lip sessile, adnate to or articulate with the column foot, concave, sacciform or sometimes produced into a conspicuous spur, entire to conspicuously 3-lobed, the lateral lobes partially enclosing the column, the midlobe apically obtuse to 2-lobed, the margins often undulate; disk sometimes with erect lamellae. Column short to somewhat elongate, erect, slightly arched, semiterete, sometimes shallowly winged and/or produced into a short foot; anther terminal, operculate, incumbent, 1-locular; pollinia 2, yellow, cartilaginous, sulcate, subtriangular in outline, attached to a short, ligulate tegula. U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina, pantropics (most diverse in Africa); 200 species, 1 in Venezuela. Eulophia alta (L.) Fawc. & Rendle, Fl. Jamaica 1: 112, t. 22. 1910. —Limodorum altum L., Syst. Nat. ed. 12, 2: 594. 1767. Terrestrial; pseudobulbs underground, tuber-like; inflorescence erect, many-flowered, racemose, to 1.5 m tall; sepals light green; petals green or green flushed with pink; lip red-maroon. Open, often disturbed, seasonally moist grasslands at 50–1500 m; Delta Amacuro (Caño Araguao, Río Cuyubini), Bolívar (Gran Sabana), Amazonas (around Puerto Ayacucho). Apure, Aragua, Mérida, Miranda; U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina, Gabon (tropical Africa). ◆Fig. 335. Fig. 335. Eulophia alta

Eurystyles 363

56. EURYSTYLES Wawra, Oesterr. Bot. Z. 13: 223. 1863. [Subtribe Spiranthinae]. Spiranthes Rich., De Orchid. Eur. 20, 28, 36. Aug.–Sept. 1817, nom. cons., pro parte. Stenorrhynchos Rich. ex Spreng., Syst. Veg. 3: 709. 1826, pro parte. Stenoptera C. Presl, Reliq. Haenk. 1: 95. 1827, pro parte. Trachelosiphon Schltr., Beih. Bot. Centralbl. 7(2): 423. 1920. Pseudoeurystyles Hoehne, Arq. Bot. Estado São Paulo 1: 129. 1944. Synanthes Burns-Bal., H. Rob. & M.S. Foster, Brittonia 37: 158. 1985. by Germán Carnevali, Ivón M. Ramírez-Morillo, and Carlos A. Vargas Small plants, commonly growing on small branches or boughs of trees. Roots slender, fleshy. Leaves basal, rosulate, shiny; margins ciliolate. Scape short, erect or more commonly arched-pendent, pubescent, few-sheathed, terminated by a densely, compact, capitate spike, densely several-flowered. Flowers small to minute, pendent, well-hidden by large, foliaceous bracts with densely, prominently ciliolate margins. Sepals basally connate, forming a short, inflated tube; dorsal sepal adnate basally to column; lateral sepals basally connate, didymous or ventricose, never forming a mentum. Petals connivent with a dorsal sepal, decurrent basally. Lip canaliculate, parallel with column, clawed; claw adnate to the inflated part of lateral sepals, nectar glands forming free fleshy auricles at base of lip or nectar glands adnate to the lip. Column slender, elongate, dorsally adnate to dorsal sepal for the most part, produced into a long, decurved foot; stigmas 2, approximate; rostellum erect, membranaceous, triangular or almost absent (in autogamous species), viscidium very small or absent (in autogamous species). Pollinia 4, soft, clavate. Ovary sessile. Southern Mexico, Central America, West Indies, Colombia, Venezuela, southern Brazil, Bolivia, northern Argentina; about 15 species; 2 or 3 species in Venezuela, 1 of these in flora area. Eurystyles cotyledon Wawra, Oesterr. Bot. Z. 13: 223. 1863. Stenoptera roehlii Schnee, Bol. Soc. Venez. Ci. Nat. 9: 249. 1946. Small, delicate plant to 5 cm tall; leaves elliptic or obovate-elliptic, acute, margins conspicuously ciliolate; scape slender but short, nodding; flowers greenish white; dorsal sepal to 3 × 0.8 mm; lip obscurely 3-lobed, 3.5(–4) mm long; ovary cylindric, glabrous. Cloud forests, ca. 1200 m; Bolívar (La Escalera to Cerro Venamo region). Aragua, Distrito Federal, Falcón, Mérida, Miranda; Colombia, Ecuador, Peru, southern Brazil, Bolivia. ◆Fig. 336. Eurystyles cotyledon is the first record of this genus in the Guayana Highlands. The Guayanan material has a narrower, more acute central lobe than typical E. cotyledon and more material is required to ascertain the taxonomic position of these plants.

Fig. 336. Eurystyles cotyledon

364

O RCHIDACEAE

57. GALEANDRA Lindl. in F.A. Bauer & Lindl., Ill. Orch. Pl. t. 8. 1838. [Subtribe Cyrtopodiinae]. Corydandra Rchb., Deut. Bot. Herb.-Buch 53. 1841. by Gustavo A. Romero-González Cespitose epiphytes or terrestrial herbs. Stem not modified into pseudobulbs, reed-like, fleshy when fresh, or most often modified into pseudobulbs of several internodes, ovoid or fusiform to cylindric, when young entirely concealed by distichous, scarious sheaths, eventually naked. Leaves several, distichous, coriaceous, mostly linear, acuminate, plicate, articulate or not, deciduous at the end of the growing season. Inflorescences terminal, arising from the apex of the pseudobulbs, erect or slightly arching to pendent, racemose, few-flowered. Flowers resupinate, often showy; floral bracts slightly concave to cymbiform, linear to narrowly ovate, acuminate, < 1/2 as long to almost as long as the pedicellate ovary. Sepals and petals membranous, erect to spreading; sepals narrowly elliptic to narrowly obovate, the lateral ones sometimes strongly oblique; petals similar to sepals, slightly to strongly oblique. Lip sessile, basally produced into a usually conspicuous spur continuous with the column foot, the spur elongate, funnelform, sometimes apically recurved, rarely shortly conical, blade simple or obscurely 2- or 3-lobed; disk basally 2–4keeled, often apically pubescent. Column short to somewhat elongate, slightly arched, sometimes subclavate and/or shallowly winged, semiterete, produced into a conspicuous foot; anther terminal, operculate, incumbent, imperfectly 2-locular or 1-locular; pollinia 2, yellow, cartilaginous, ovoid to subspherical, subcircular in cross section, sulcate, attached to a short, semilunate or narrowly oblong tegula. U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina; 20–25 species, 10 in Venezuela, all in the flora area. Key to the Species of Galeandra 1. 1. 2(1). 2. 3(2). 3. 4(1). 4. 5(4). 5.

Plants terrestrial, with small (to 3 cm), roundish underground corms, leafless at anthesis or with nonarticulate leaves ................................. 2 Plants epiphytic, with aerial, elongate, fusiform pseudobulbs, usually > 5 cm long, and articulate leaves ......................................................... 4 Plants saprophytic, leafless at anthesis; dorsal sepal at least 2 cm long; lip entire ................................................................................... G. beyrichii Plants autotrophic, with leaves at anthesis; dorsal sepal < 2 cm long, lip entire or conspicuously 3-lobed ............................................................. 3 Lip obscurely to conspicuouly 3-lobed, spur deep, narrow, strongly tapering ............................................................................ G. stylomisantha Lip entire, spur shallow, wide, not strongly tapering .......... G. carnevaliana Lip spur broadly conical or sacciform, much shorter than the pedicellate ovary, conspicuously under-curled ........................................ G. devoniana Lip spur narrowly conical, as long or longer than the pedicellate ovary, not under-curled .................................................................................... 5 Lip blade of equal length or much shorter than the spur; disk with 2 or 3 axial ridges terminating in a tuft of white hair in front ................... 6 Lip blade longer than the spur; disk with 2–4 axial ridges not terminating in a tuft of white hairs (if the blade is pubescent the hairs are aligned in axial rows) ......................................................................................... 8

Galeandra 365

6(5). 6. 7(6). 7. 8(5). 8. 9(8). 9.

Lip blade equal in length to the spur ......................................... G. stangeana Lip blade much shorter than the spur ...................................................... 7 Column, including clinandrium, < 1 cm long ........................ G. macroplectra Column, including clinandrium, > 1 cm long ..................... G. magnicolumna Lip conspicuously emarginate at the apex, thus slightly 2-lobed; disk with 2 axial ridges ......................................................................... G. badia Lip with 2 indentations alongside the apex, thus slightly 3-lobed, midlobe emarginate or not, or the disk with 4 axial ridges ............................... 9 Lip midlobe margin dark reddish purple, conspicuously emarginate .................................................................................................. G. duidensis Lip midlobe margin flushed with pale pink, entire ......................... G. minax

Galeandra badia Garay & G.A. Romero, Harvard Pap. Bot. 3: 58. 1998. Epiphyte; pseudobulbs fusiform; flowers greenish white; sepals and petals golden brown; lip base, blade, and margins white to yellowish white, the spur greenish brown with maroon-brown stripes. Wet forests, 100–200 m; Bolívar (tributary of Río Carrao). Guyana, Suriname, French Guiana. ◆Fig. 337. Galeandra beyrichii Rchb. f., Linnaea 22: 854. 1849. Terrestrial with small, roundish, underground corm; stems reed-like, fleshy, leafless; inflorescence terminal; flowers greenish white; sepals and petals pale green with white margins; lip greenish white, the margins white, striped with dull purple. Open wet forests, 1100–1300 m; Bolívar (tributary of Río Kukenán, base of Roraima-tepui). Coastal Cordillera around Caracas, Mérida, Portuguesa; U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina. Galeandra carnevaliana G.A. Romero & Warford, Lindleyana 10: 75. 1995. Terrestrial, presumably with a small underground corm; lip with a short, shallow spur. Gallery forests, ca. 200 m; Bolívar (road between Ciudad Bolívar and Ciudad Piar). Endemic. Galeandra devoniana R.H. Schomb. ex Lindl., Sert. Orchid. t. 37. 1840. Epiphyte, most often on Leopoldinia pulchra Mart.; pseudobulbs fusiform; flowers purple-brown; sepals and petals pale greenish brown; lip base and blade white, the spur white with maroon veining, margins white with maroon or pink veining; column cream-

colored. Wet forests along rivers, 50–200 m; Amazonas (Brazo Casiquiare, Río Negro, Río Sipapo). Brazil (Amazonas, Pará). ◆Fig. 338. Galeandra duidensis Garay & G.A. Romero, Harvard Pap. Bot. 3: 58. 1998. Epiphyte; pseudobulbs fusiform; flowers reddish bronze; sepals and petals reddish brown; lip base and blade white, spur reddish bronze, margins dark red-purple; column white with purple spots at base. Low tepui forests, ca. 1200 m; Amazonas (Cerro Duida). Endemic. ◆Fig. 342. Galeandra macroplectra G.A. Romero & Warford, Lindleyana 10: 77. 1995. Epiphyte; pseudobulbs fusiform; sepals and petals light green with reddish overlay; lip greenish yellow, white toward the apex, spur light green. Wet forests, 50–200 m; Amazonas (Río Cataniapo, Río Ventuari). Apure; Colombia. ◆Fig. 341. Galeandra magnicolumna G.A. Romero & Warford, Lindleyana 10: 81. 1995. Epiphyte; pseudobulbs fusiform; sepals and petals light green with bronze overlay; lip greenish yellow, reddish toward the apex, spur light green. Wet forests, 100–200 m; Amazonas (Río Pavones). Colombia. Galeandra minax Rchb. f., Gard. Chron. n.s. 1: 786. 1874. Galeandra lacustris Barb. Rodr., Gen. Spec. Orchid. 1: 86. 1877. Epiphyte; psuedobulbs fusiform; flowers greenish white; sepals and petals light green with bronze overlay; lip base white, the blade and margins white flushed with pale pink, spur light greenish yellow. Wet forests, 50– 200 m; Amazonas (Cerro Moriche, Río Cataniapo). Brazil (Amazonas, Pará, Mato Grosso). ◆Fig. 339.

366

O RCHIDACEAE

Fig. 337. Galeandra badia

Fig. 338. Galeandra devoniana

Fig. 339. Galeandra minax

Fig. 340. Galeandra stylomisantha

Fig. 341. Galeandra macroplectra

Fig. 342. Galeandra duidensis

Galeottia 367

Galeandra stangeana Rchb. f., Bonplandia (Hanover) 4: 323. 1856. Epiphyte; pseudobulbs fusiform; flowers greenish white; sepals and petals light green with bronze overlay; lip, including the spur, reddish white. Moist lowland forests, 50–200 m; Amazonas (San Fernando de Atabapo). Colombia, Brazil (Amazonas, Espírito Santo, Mato Grosso). Galeandra stylomisantha (Vell.) Hoehne, Arq. Bot. Estado São Paulo n.s. 2: 146. 1952. —Orchis stylomisantha Vell., Fl. Flumin. 9: t. 46. 1831. Galeandra juncea Lindl., Sert. Orchid. sub t. 37. 1840.

Terrestrial, with a small underground corm; flowers greenish pink; sepals and petals rose; lip base and blade white-pink, spur pinkish green, margins of the midlobe pink overlaid with dark rose-purple veins. Gallery forests, open savannas, 1200–1400 m; Bolívar (Kamarata, Roraima-tepui, Santa Elena de Uairén), Amazonas (Sierra Parima). Carabobo, Trujillo; Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Espírito Santo, Mato Grosso, Minas Gerais, Paraná, Río Grande do Sul, Rio de Janeiro, Santa Catarina, São Paulo), Bolivia, Paraguay, northern Argentina. ◆Fig. 340.

58. GALEOTTIA A. Rich., Ann. Sci. Nat. Bot. sér. 3, 3: 25. 1845. [Subtribe Zygopetalinae]. Mendoncella A.D. Hawkes, Orquídea (Niteroi) 25: 7. 1963. by Gustavo A. Romero-González Cespitose or rhizomatous epiphytes or sometimes terrestrial herbs. Stem modified into narrowly oblong to oblong-ovate, sometimes obtuse, tetragonal, straight or slightly arched pseudobulbs of one internode, aggregate or distant on the rhizome, when young concealed by distichous, scarious sheaths. Leaves most often 2, sometimes 3, coriaceous, narrowly linear-ovate to obovate, acute to acuminate, attenuate toward the base, subpetiolate or petiolate, plicate, articulate. Inflorescences lateral, arising from the base of the pseudobulbs, erect to slightly arching, racemose, few-flowered, < 1/2 as long as to longer than the leaves. Flowers resupinate, showy; floral bracts concave, ovate or narrowly ovate, shorter than to longer than the pedicellate ovary. Sepals and petals membranous, spreading to erectspreading; dorsal sepal linear-oblong, narrowly ovate, oblong-ovate, or ovate, acute to acuminate; lateral sepals laterally adnate to column foot, basally saccate, the inner basal margins sometimes strongly inrolled; petals similar to lateral sepals, sometimes slightly to strongly oblique. Lip fleshy, clawed, the claw with or without a retrorse horn; lip divided into a short, cup-shaped hypochile, rhomboid to subtriangular, narrower than the epichile, with an axial, lamelliform callus, and a narrowly ovate or ovate, acuminate epichile, or lip somewhat 3-lobed to conspicuously 3-lobed, lateral lobes dentate to lacerate, midlobe narrowly oblong, ovate, obovate, elliptic, or subcircular, obtuse, acute, or acuminate, apiculate or not, entire, finely dentate or lacerate, glabrous, sparsely puberulent, or verrucose, the disk with a lamellate callus. Column short to elongate, slightly arched, semiterete, basally produced into a conspicuous foot, apically auriculate, the auricles entire, denticulate, or lacerate; anther terminal; pollinia 4, yellow, cartilaginous, dorsoventrally flattened, obovate to clavate, attached in 2 pairs to a subtriangular or rhombiform tegula. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, Peru, Brazil, Bolivia; 11 species, 3 in Venezuela, 2 of these in the flora area.

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Key to the Species of Galeottia 1. 1.

Lip widest at the midlobe, lateral lobes falcate, midlobe with finely dentate margins .................................................................................. G. burkei Lip widest at the lateral lobes, lateral lobes round, midlobe deeply fimbriate .................................................................................... G. jorisiana

Galeottia burkei (Rchb. f.) Dressler & Christenson, Lindleyana 3: 221. 1989. —Zygopetalum burkei Rchb. f., Gard. Chron. n.s. 20: 684. 1883. —Mendoncella burkei (Rchb. f.) Garay, Orquideología 8: 28. 1973. Terrestrial (rarely epiphytic); pseudobulbs 2- or 3-foliate; inflorescence erect; sepals and petals yellow-green with purplebrown mottling; lip white with a large purple crest; column yellow with purple streaks. Growing on low shrubs or on sandstone boulders in dwarf, swampy tepui forests, 1000– 2200 m; Bolívar (Auyán-tepui, Cerro El Sol, Cerro Guaiquinima, Cerro Guanay, JauaSarisariñama massif, Macizo del Chimantá [Acopán-tepui], Ptari-tepui, base of Roraimatepui, Sierra Pakaraima, Uaipán-tepui), Amazonas (Cerro Duida, Cerro Sipapo, Cerro Yapacana, Río Atabapo basin, Río Iguapo, Río Matacuni, Sierra Parima). Guyana. ◆Fig. 344. Galeottia jorisiana (Rolfe) Schltr., Repert. Spec. Nov. Regni Veg. Beih. 6: 86. 1919. —Zygopetalum jorisianum Rolfe, Gard. Chron. ser. 3, 7: 704. 1890. —Mendoncella jorisiana (Rolfe) A.D. Hawkes, Orquídea (Niteroi) 25: 8. 1963.

Fig. 343. Galeottia jorisiana

Fig. 344. Galeottia burkei

Gomphichis 369

Epiphyte (sometimes terrestrial); pseudobulb 2-foliate; inflorescence erect to slightly arching, arising from the new growth; flowers showy; sepals and petals light green with dark green or brown markings; lip mostly white, deeply laciniate, the crested callus yellow; column cream. Tall forests in savannas,

dwarf forests, forested slopes, sandstone substrate on tepuis or high plateaus, 1000–2000 m; Bolívar (Cerro Guaiquinima, Macizo del Chimantá [Amurí-tepui], Serranía Marutaní, Uaipán-tepui, Uei-tepui), Amazonas (Cerro Marahuaka, Sierra de la Neblina). Endemic. ◆Fig. 343.

59. GOMPHICHIS Lindl., Gen. Sp. Orchid. Pl. 446. 1840. [Subtribe Prescottiinae]. Stenoptera C. Presl, Reliq. Haenk. 1: 95. 1827, pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial or subterrestial herbs, or very rarely subepiphytes, erect, usually shade-loving. Roots fleshy, fasciculate, pubescent. Stems unbranched, terete, densely foliose basally, laxly foliose toward apex. Leaves not articulate, convolute, the basal ones larger, often basally attenuate into a pseudopetiole and then into a sheathing base; blade narrowly elliptic to rarely obovate. Inflorescence terminal, spiciform, lax to densely many-flowered, erect, glabrous or variously pubescent; peduncle longer than the rachis, enveloped by several tubular to foliaceous sheaths; floral bracts conspicuous, membranous. Flowers nonresupinate, spreading to nodding. Perianth segments variously pubescent or ciliate, greenish or yellowish, lip sometimes white. Sepals free, subequal, erect or spreading in the apical 1/2; petals linear to obovate. Lip fleshier than the other perianth segments, free from column, mostly rhombic-ovate or suborbicular, unlobed or 3-lobed, usually pubescent; often divided in a saccate hypochile and a very fleshy, sulcate epichile. Column often clavate, bent backward or S-shaped, mostly pilose, wingless, footless, apically blunt; anther dorsal, erect, subequal to rostellum; clinandrium well developed, membranous; pollinia 4, soft, viscidium terminal; rostellum erect, broad, laminar, emarginate; stigma entire. Costa Rica, Panama, Colombia, Venezuela, Guyana, Ecuador, Peru, Bolivia; ca. 25 species, 7 in Venezuela, 2 of these in the flora area. Key to the Species of Gomphichis 1. 1.

Lateral sepals narrower than dorsal one; column apex not conspicuously dilated, column geniculate only once .......................................... G. adnata Lateral sepals as wide as dorsal sepal; column apex conspicuously dilated, column geniculate twice ........................................... G. costaricensis

Gomphichis adnata (Ridl. ex Thurn) Schltr., Repert. Spec. Nov. Regni Veg. Beih. 6: 51. 1919. —Stenoptera adnata Ridl. ex Thurn, Timehri 5: 205. 1886. —Stenoptera adnata Ridl. ex Oliv., Trans. Linn. Soc. London, Bot. 2: 284, t. 47a, figs. 1–6. 1887, nom. superfl. Terrestrial herb to 55 cm tall at flowering. Cloud forests on tepui summits, ca. 1800 m; Bolívar (Roraima-tepui). Mérida; Guyana.

Gomphichis costaricensis (Schltr.) Ames, Hubb. & C. Schweinf., Bot. Mus. Leafl. 3: 37. 1934. —Stenoptera costaricensis Schltr., Beih. Bot. Centralbl. 36(2): 375. 1918. Terrestrial or subterrestrial herb to 70 cm tall in flower. Cloud forests on tepui summits, 1500–2100 m; Bolívar (Auyán-tepui, Macizo del Chimantá, Roraima-tepui, Sororopántepui). Anzoátegui, Lara, Mérida, Monagas, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Costa Rica, Panama, Colombia, Ecuador. ◆Fig. 345.

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Fig. 345. Gomphichis costaricensis

Fig. 346. Gongora atropurpurea

60. GONGORA Ruiz & Pav., Fl. Peruv. Prodr. 117, t. 25. 1794. [Subtribe Stanhopeinae]. by Gustavo A. Romero-González Cespitose epiphytes, often in ant gardens. Stem modified into ovoid to ellipticsubcylindric, sulcate pseudobulbs of 1 internode. Leaves 1–3, most often 2, narrowly to broadly elliptic, acute to acuminate, plicate, articulate. Inflorescences 1–several, lateral, arising from the base of the pseudobulbs, pendent, racemose, many-flowered. Flowers nonresupinate, showy; floral bracts inconspicuous, triangular-ovate, shorter than the elongate, straight to strongly recurved pedicellate ovary. Sepals and petals membranous, basally connivent or the petals and the dorsal sepal adnate to the column basal 1/2; sepals spreading to sharply reflexed, ovate to subtriangular, apically abruptly apiculate; petals broadly to narrowly ovate, oblique, acuminate. Lip fleshy, waxy, sessile to conspicuously clawed, divided into 2 complex portions, the hypochile, conspicuously lobed or sacciform, with horn-like projections or auricles or not, and the elongate, narrowly ovate, or gibbous epichile. Column slender, elongate, semiterete, glabrous, slightly clavate, apically winged or not, basally produced into a conspicuous foot; anther terminal, operculate, incumbent, 1-locular; pollinarium with 2 yellow pollinia, narrowly obovate, cleft, cartilaginous, and dorsoventrally flattened, the tegula short, oblong-linear, the viscidium subtriangular.

Guanchezia 371

Southern Mexico, Central America, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 40 species, 3–5 in Venezuela, 2 of these in the flora area. Key to the Species of Gongora 1. 1.

Lip conspicuously clawed, hypochile with 2 baso-lateral, linear-oblong appendices at least 5 mm long .......................................... G. atropurpurea Lip shortly or not clawed at all, the hypochile appendices to 3 mm long ............................................................................................. G. pleiochroma

Gongora atropurpurea Hook., Exot. Fl. pl. 178. 1825. Epiphyte; inflorescence long, pendent, to 15-flowered; flowers dark purple. Wet forests, 50–2000 m; Delta Amacuro (locality not given), Bolívar (Río Antavari), Amazonas (San Carlos). Coastal Cordillera; Trinidad-Tobago, Guyana, Ecuador, Peru, Brazil. ◆Fig. 346. Gongora pleiochroma Rchb. f., Hamburg. Gart. Blumenzeit. 16: 421. 1860. Variable epiphyte; inflorescence long, pendent, to 25-flowered; flowers purple or

yellowish white spotted with maroon. Locally frequent in wet forests, 100–200 m; Delta Amacuro (Río Acure), Bolívar (Río Cachimi, Río Erebato), Amazonas (Río Cataniapo, Río Cuao, Río Siapa, Sierra de La Neblina). Coastal Cordillera, Monagas; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. Gongora pleiochroma is part of a difficult species group and until the species boundaries within the group are better understood, the determination should be regarded as tentative.

61. GUANCHEZIA G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1135. 2000. [Subtribe Bifrenariinae]. by Gustavo A. Romero Cespitose, terrestrial or rarely lithophytic herbs. Stem modified into ovoid pseudobulbs of 1 internode, when young entirely concealed by distichous, scarious sheaths, eventually naked. Leaf 1, lanceolate, acute, plicate, articulate, long-petiolate. Inflorescences lateral, arising from the base of the pseudobulbs, erect, racemose, 1- or 2(3)-flowered. Flowers fleshy, resupinate, showy; floral bracts concave, narrowly elliptic, acute, shorter than the pedicellate ovary. Sepals oblong to oblonglanceolate, the lateral ones slightly oblique and at the base adnate to the column foot; petals oblong-lanceolate, smaller than the sepals. Lip fleshy, sessile, spatulateobovate when flattened, pilose toward the base but otherwise without ornamentation, basally produced into a spur continuous with the column foot, the spur short, conical, apically slightly recurved. Column erect, recurved, semiterete, produced into a foot, apically with a conspicuous clinandrial hood; anther terminal, operculate, incumbent, 1-locular; pollinarium with 2 yellow pollinia, obovate, cleft, cartilaginous, and dorsoventrally slightly flattened, the tegula U-shaped, with the viscidium placed on the very base of the “U.” Endemic to the Guayana Shield in Colombia (Guánia), Venezuela, and Brazil (Amazonas); 1 species. Guanchezia differs from Bifrenaria in its ovoid, non-angulate pseudobulbs, with cataphylls covering up to 1/4 of the petiole, the terete petiole as long or longer than the leaf blade, the erect inflorescence, and the flowers lacking any ornamentation on the lip. Another unique character of the flowers of this species is the clinandrium hood that stands over the entire anther and beyond.

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Guanchezia maguirei (C. Schweinf.) G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1136. 2000. —Bifrenaria maguirei C. Schweinf., Amer. Orchid Soc. Bull. 28: 199, fig. 1. 1959. Terrestrial or rarely rupicolous herb to 1.2 m tall; leaves long-petiolate; flowers showy, cream or yellowish, externally with greenish or purplish spots. Open tepui associations, often in bogs, 1200–1600 m; Amazonas (Cerro Aracamuni, Cerro Autana, Cerro Duida, Cerro Parú, Cerro Sipapo, Cerro Yapacana, Sierra de la Neblina). Colombia, northern Amazonian Brazil. ◆Fig. 347.

Fig. 347. Guanchezia maguirei

62. HABENARIA Willd., Sp. Pl. 4: 44. 1805. [Subtribe Habenariinae]. by Ernesto Foldats, Germán Carnevali, and Ivón M. Ramírez-Morillo Terrestrial, sometimes aquatic or semiaquatic, rarely subepiphytes, erect, mostly sun-loving, with 1–3 underground tubers. Leaves not articulate, convolute, membranaceous to thinly fleshy, usually cauline, in some species basal, sometimes almost absent or the blades reduced and sheath-like; sheaths funnelform. Inflorescences terminal racemes, lax or dense, mostly few- to many-flowered, rarely 1-flowered, erect; floral bracts usually conspicuous, sometimes spathaceous. Flowers inconspicuous to relatively large and showy, resupinate, usually white, green, or dull yellow, rarely orange, bright yellow, pink, or purple; mostly fragrant at night. Perianth segments fleshy, widely spreading or the dorsal sepal and petals hooded over the column; sepals free, usually broader than the petals; petals usually 2-lobed, sometimes simple or 3-lobed. Lip usually 3-lobed, sometimes simple, basally with a conspicuous spur, rarely absent; lateral lobes usually narrower than central lobe, often linear or filiform, rarely broader than central lobe. Column short and thick; anther erect, basally firmly united with the column, 2-locular, each locule bulging out on an elongated channel; pollinia 2, sectile, with 2 basal, interlocular caudicles; stigmas convex, 2-lobed, the lobes often stalked. Pantropics and subtropics, U.S.A. (Florida, Georgia, North Carolina, South Carolina, Texas, Virginia), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina, Uruguay; ca. 600 species, 40 in Venezuela, 30 of these in the flora area. Most of the species with green, dull yellow, or white flowers are pollinated by moths of different sizes; pollination by butterflies has been reported for temperate species with bright yellow, orange, or purple flowers. The few tropical species with

Habenaria 373

these bright colors are likely pollinated by butterflies or hummingbirds. Many species are autogamous. Portions of this treatment have been extracted and adapted from a recent compendium of the genus for the Guianas (J. Renz. The genus Habenaria (Orchidaceae) in the Guianas. Candollea 47: 483–513. 1992). This paper has been particularly important in elucidating the identities of the several species with setaceous, grass-like leaves (here called the Habenaria leprieuri complex). However, the paper only occasionally indicates localities or vouchers for the Venezuelan plants included (or mentioned in passing). Thus, a few taxa (e.g., H. pratensis var. parviflora) are included in this account with only incomplete data. Habenaria angustifolia H.B.K. has been reported from the Guayana Region but is not included because it is known only from the type in Paris and has not been examined by contemporary orchidologists. The identity of this species could not be clarified, and its affinity with other species of Habenaria from the flora area is unknown. It has been suggested (Renz 1992) that this taxon might be close to H. pauciflora (as H. trifida), and the long spur indicated by the original description is also suggestive of this fact. Habenaria platydactyla Kraenzl., a member of the H. leprieuri complex, is likely to occur in the flora area since it is known from surrounding areas. If found, it would be easy to recognize among related species due to its white flowers and broad perianth segments, particularly the midlobe of the lip. Key to the Species of Habenaria 1. 1. 2(1). 2. 3(2).

3.

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

5.

Lip simple or (rarely) basally protuberant ............................................... 2 Lip conspicuously 3-lobed .......................................................................... 5 Spur ≥ 3.5 cm long; rachis completely hidden by the floral bracts ...................................................................................................... H. obtusa Spur 1.4–3 cm long, when > 2 cm the rachis not completely hidden by the floral bracts ............................................................................................ 3 Petals ca. 4 mm long, the posterior lobe linear-oblong, apex obtuse to acute, the anterior lobe triangular acute, ca. 2 mm long or at least 1/2 the length of the posterior lobe .................................................. H. sp. A Petals 3–11 mm long, the posterior lobe quadrate or oblong, the anterior lobe either absent or reduced to a teeth or else much shorter than the posterior lobe ......................................................................................... 4 Petals 3–3.2 × 1.7–2 mm, acute ................................................... H. quadrata Petals 3.7–11 × 1.5–6 mm, apically unequally 2- or 3-lobed .... H. floribunda Leaves basally crowded in a rosette, 2–4(5) in number, leaf blades elliptic, broadly elliptic to almost suborbicular, abruptly narrowing basally into their sheaths; distal to the basal rosette, the leaves abruptly transformed into bracts ± appressed to the stem or peduncle of the inflorescence, thus the plants resembling members of the Spiranthiinae; plants from forest environments ........................................................... 6 Leaves arranged distichously along the stem, if crowded on the base of the plant, the leaf blades narrowly elliptic to setaceous and gradually narrowing into their sheaths; distal to the basal rosette the leaves

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6(5).

6.

7(5). 7. 8(7). 8. 9(7).

9.

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

11.

12(10).

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

14.

gradually transform into bracts on the stem into the inflorescence; plants commonly from savanna environments, more rarely from forests ......... 7 Plants 7.2–10(–15) cm tall; leaves 1.1–2.2 cm long; petals simple; lip with lateral lobes much shorter than the midlobe; floral bracts much shorter than the ovary; peduncle with 2(3) internodes ....................... H. pygmaea Plants 15–50 cm tall; leaves 5–15 cm long; petals 2-lobed; lip with the lateral lobes about as long or longer than the midlobe; floral bracts about as long as the ovary or longer; peduncle with 3–5 internodes ..................................................................................................... H. distans Lobes of lip attenuated toward the base; flowers yellow or orange ......... 8 At least the central lobe of lip not attenuated toward the base; flowers greenish or white, rarely yellow ........................................................... 9 Dorsal sepal 15–17 mm long; lateral lobes of lip obovate, truncate, 2–4 times longer than wide ........................................................................... H. huberi Dorsal sepal 9–11 mm long; lateral lobes of lip oblong or narrowly spathulate, rounded, at least 6 times longer than wide .................... H. pratensis Leaf blades not sharply differentiated from their sheaths (the portion of the leaf that clasps the stem), usually narrower to as broad as or slightly broader than the sheaths and linear to setaceous, 2–4(–5) mm wide, usually subparallel to the stem or even clasping it, usually shorter than to as long as the internodes of the stem (rarely longer) .............................................................................................................. 10 Leaf blades well differentiated and longer than their sheaths (the portion of the leaf that clasps the stem), usually broader than the sheaths and lanceolate or narrowly elliptic to ovate-elliptic, (5–)8–20 mm long, often somewhat to sharply diverging from the stem and much longer than the internodes of the stem .......................................................... 15 Dorsal sepal 7–9 mm long ....................................................................... 11 Dorsal sepal 2.5–5.5 mm long ................................................................. 12 Leaves long, to 20 cm; dorsal sepal with 5(–7) conspicuous, dark green veins; the lip with a ca. 2 mm long undivided basal portion, lobes subequal ....................................................................................... H. sprucei Leaves rather short, usually not longer than 6 cm; dorsal sepal with 3–(5) veins; lip divided to its base, the lateral lobes somewhat narrower than the central lobe ............................................................................ H. setacea Spur longer than half the length of the ovary; the floral bract acuminate and ending in a bristle-like tip, reaching the tip or the lower half of the spur; fruits appressed to the rachis .................................................... 13 Spur half the length of the ovary or shorter; floral bract much shorter than the pedicellate ovary ................................................................... 14 Lateral lobes of lip longer than midlobe, the midlobe wider than the laterals ................................................................................ H. mesodactyla Lateral lobes of lip as long as or shorter than midlobe, the three segments about as wide ............................................................................ H. leprieuri Flowers bright yellow, distichously or cylindrically arranged on the rachis, the rachis short and densely flowered (3–5 flowers per side); all of the rachis partially overlapped by floral bracts ............... H. heptadactyla Flowers totally green, all facing the same side of the rachis; the rachis

Habenaria 375

15(9). 15. 16(15). 16. 17(16). 17.

18(17).

18.

19(16).

19.

20(19).

20.

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

22.

long and lax (5–10 flowers on a particular inflorescence), with conspicuous sections of naked rachis between flowers ....... H. ayangannensis Stems and foliar sheaths spotted; foliar margins scarious; flowers entirely white ........................................................................... H. monorrhiza Stems and foliar sheaths not spotted; foliar margins not scarious or when somewhat scarious the flowers usually not entirely white ................ 16 Petals trifid, with a tooth on each side, or with a single tooth on the upper side of the posterior lobe ..................................................................... 17 Petals bifid, rarely simple ....................................................................... 19 Petals with small teeth basally (one on each side of the posterior lobe, the posterior lobe ovate suborbicular; plants from cloud forests ..... H. nilssonii Petals trifid, or bifid with a small lobe or tooth pointing upward from the upper margin of the posterior lobe; the plants from lowland savanna environments ....................................................................................... 18 Petals with a well-developed anterior lobe, somewhat shorter than the posterior lobe, and a smaller lobe or tooth pointing upward above the base of the posterior lobe, or else the petal bilobed but the short, acute lobe on the “wrong side” and pointing upward; lip with a relatively long (ca. 1/4 total length) undivided portion at base .............................. H. sp. B Petals trifid, the upper lobe and the basal lobe ± equally developed, these narrower and shorter than the posterior lobe; lip divided from its base ....................................................................................................... H. trifida Plants aquatic, growing in shallow waters of permanently or seasonally flooded areas, like ditches, edges of lakes, ponds, swamps, or rivers, often growing on floating mats; rhizome ± elongated, with long roots; leaves with the midvein carinate on the underside ........................... 20 Plants of wet and dry areas but never entirely aquatic; rhizome abbreviated, roots short; leaves with the midvein not carinate on the underside .............................................................................................................. 21 Anterior lobe of the petals longer than the posterior lobe; lateral lobes of the lip longer than midlobe; spur 7–14 cm long; lip undivided in the lower 1/4; plants 50–150 cm tall ............................................ H. longicauda Anterior lobe of the petals shorter than posterior lobe; lateral lobes of the lip shorter than midlobe; spur 0.8–1.5 cm long; lip divided from its base; plants 30–60(–100) cm tall ................................................. H. repens Pedicel of ovary short, < 1 cm long, or the ovary sessile; flowers held ± appressed to the rachis ........................................................................ 22 Pedicel of ovary elongated, 2–3.5 cm long; flowers ± spreading from the rachis .................................................................................................... 23 Flowers widely spreading, in a ± right angle to the rachis, spur narrowly cylindrical, very slender throughout, the apex mostly hidden among the bracts; rostellum midlobe broadly triangular, slightly overtopping the anther loculi; lip below the lateral lobes with an undivided portion ..................................................................................................... H. ernestii Flowers slightly spreading; spur cylindrical, slightly thickened in the apical portion, frequently curved outwards; rostellum midlobe prominent and hooded, clearly overtopping the anther loculi; lip divided from its base .................................................................................... H. lehmanniana

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23(21). Spur (3–)5–12 cm long, as long or longer than the ovary with pedicel; plants with 1–3(–6) flowers; dorsal sepal 9.5–16 mm long ......... H. trifida 23. Spur 1.5–2.5 cm long longer than the ovary with pedicel; plants with 4– 15 flowers; dorsal sepal 4–7 mm long ................................................. 24 24(23). Leaves linear to narrowly lanceolate; flowers with white petals, lip, and spur ........................................................................................... H. caldensis 24. Leaves lanceolate, oblong-lanceolate, or broader; flowers with pale green petals, lip, and spur ............................................................................. 25 25(24). Lateral lobes of lip shorter than to as long as the central lobe ............. 26 25. Lateral lobes of lip conspicuously longer than the midlobe ................... 27 26(25). Leaves 4–25 cm long, narrowly linear-lanceolate; anterior lobe of petals reduced to a tooth or, if linear, < 1/3 the length of the posterior lobe; inflorescences densely multiflorous .......................................... H. parviflora 26. Leaves 2–3.5 cm long, narrowly elliptic; anterior lobe of lip well developed and slightly shorter than posterior lobe; inflorescences laxly multiflorous ................................................................................... H. roraimensis 27(25). Leaves (18–)20–25 mm wide, laxly arranged along the stem; spur 40– 100 mm long ......................................................................... H. quinqueseta 27. Leaves > 10 mm wide laxly or densely arranged along the stem or at its base; spur 6–32 mm long ..................................................................... 28 28(27). Leaves 2 or 3(–5), mainly in the lower portion of the stem, linear to linearlanceolate, (3–)6–11 mm wide, distally the leaves abruptly become bract-like, thus, the apical portion of the plant consists of a welldefined inflorescence with 1–5(–7) flowers ......................................... 29 28. Leaves several to many, distributed along the stem, narrowly lanceolate to narrowly ovate, (6–)8–15 mm wide, distally narrowed gradually into the many-flowered inflorescence ......................................................... 30 29(28). Ovary on top of a 10 mm long pedicel; spur 28–32 mm; dorsal sepal widely ovate to suborbicular, 5–6.2 mm long; flowers 1–3 ................... H. dusenii 29. Ovary sessile or almost; spur 10–25 mm long; dorsal sepal ovate, 5– 11; flowers 3–7 .................................................................. H. schomburgkii 30(28). Spur longer than the ovary; lip with an undivided portion on the lower 1/5 of its length; sepals acutely apiculate; lateral lobes of lip at most 1.5 times longer than and parallel to the midlobe .................... H. armata 30. Spur shorter than to as long to the ovary; lip divided to its very base; sepals not apiculate; lateral lobes of lip ca. twice as long as the midlobe and curving upward ........................................................ H. amambayensis Habenaria amambayensis Schltr., Repert. Spec. Nov. Regni Veg. 16: 353. 1920. Terrestrial; leaves many along a leafy stem, linear-lanceolate, acuminate; inflorescence densely many-flowered; lateral lobes of lip and anterior lobe of petals twice as long as the midlobe of lip and the posterior lobes of the petals; sepals green, petals and lip white. Expected to occur in the savannas of Bolívar and Amazonas. Guyana, French Guiana, Brazil.

Habenaria armata Rchb. f., Bonplandia (Hanover) 2: 10. 1854. Habenaria moritzii Ridl., Trans. Linn. Soc. London, Bot. 2: 284. 1887. Habenaria enthomantha auct. non (Llave & Lex) Lindl. 1835: sensu Foldats in Lasser, Fl. Venez. 15(1): 55. 1969, pro parte; Dunst. & Garay, Venez. Orchid. Ill. 2: 167. 1961. Terrestrial 10–40 cm tall; leaves 3–7, located in the basal 1/2 of the stem, 2–8.5 × 0.6

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–1.5 cm; inflorescences few- to many-flowered racemes; dorsal sepal 4–7 × 2–5 mm; lateral sepals 4.4–8 mm long, slightly narrower than dorsal one; posterior lobe of the petals 3.2–6 × 0.8–1.5 mm, anterior lobe narrower than the posterior one, 0.3 mm wide; central lobe of lip 3.4–7 × 0.5–1.3 mm, lateral lobes similar to the anterior lobe of the petals; spur 5–16 mm long; stigmatic prolongations conspicuous, not tubercular. Open habitats, (100–)800–1700 m; Bolívar (near El Paují, Gran Sabana), Amazonas (Coromoto, near Puerto Ayacucho). Anzoátegui, Miranda, Guárico; Colombia, Guyana. This concept is here broadly circumscribed to include a series of populations with almost identical flowers but strikingly different plants. The flowers of this complex are characterized by anterior lobes of the petals and lateral lobes of the lip filiform and much longer than the corresponding posterior lobes of the petal and lip midlobes. The plants are small to medium sized (to 40 cm tall) but leaves range from short, broad, and almost appressed to the stems (to 5.5 cm long) to linear-elliptic, acute and to 8.5 cm long. The plants with short, broad leaves have been treated in previous floristic treatments as Habenaria entomantha, a species endemic to central Mexico with white flowers, shorter anterior lobes and lateral lobes, and broader perianth segments. A detailed study of this complex of forms (also including H. moritzii Ridl., H. ernstii Schltr., and possibly other names) is required to ascertain whether this is a single, variable species, or several closely related taxa. At the heart of this problem is the identity of the type of H. armata. There are several collections in the Reichenbach herbarium referred by him to this species. These specimens represent populations of the two kinds of plants discussed above and a lectotypification is required to properly assign names should more than one taxon be recognized in the complex. Habenaria moritzii Ridl. was based on two syntypes, one from the Coastal Range, Moritz 630B (and conspecific with H. gollmeri Schltr., sensu Renz 1992, 492) and the other from Roraima (Im Thurn 367, K), here selected as the lectotype of the species. This lectotypification renders the name, H. moritzii, conspecific with (or at least a mem-

ber of the complex of) Habenaria armata Rchb. f. Habenaria ayangannensis Renz, Candollea 47(2): 490. 1992. Terrestrial, erect herb 20–30 cm tall; leaves ca. 2.5 cm long; flowers greenish, dorsal sepal ca. 2.5 mm long and wide. Expected to occur in the savannas of Bolívar. Guyana. Habenaria caldensis Kraenzl., Bot. Jahrb. Syst. 16: 128. 1894. Habenaria ernestii auct. non Schltr. 1914: sensu Foldats in Lasser, Fl. Venez. 15(1): 49. 1969, pro parte (as Habenaria rodeiensis Barb. Rodr.). Terrestrial 20–60(–100) cm tall; leaves distichous, 6–16 × 5–16 mm; flowers white; dorsal sepal 5–7.5 × 4–5.5 mm; lateral sepals 6–10 × 2–4.2 mm; posterior lobe of the petals 5–7 × 1.5–2.7 mm, anterior lobe linear-filiform, 6–8 mm long; central lobe of lip 5–9 × 0.7–1.5 mm, lateral lobes 6–11 × 0.3 mm; spur 3–4 cm long. Open, humid places, 1200– 2300 m; Bolívar (Auyán-tepui, Ptari-tepui, Sororopán-tepui). Guyana, Suriname, Peru, Brazil (Amazonas, Goiás, Minas Gerais). Habenaria distans Griseb., Cat. Pl. Cub. 270. 1866. Terrestrial 14–55 cm tall; leaves 2–6, the basal ones 5–15 × 1.7–5 cm; inflorescences few- to many-flowered racemes; flowers greenish, petals and lip paler; sepals 5–9 mm long; dorsal sepal 4–5 mm wide, the lateral ones 2–3 mm wide; posterior lobe of petals 5– 9 × 1–1.2 mm, anterior lobe nearly lacking to reduced to a tooth or > the posterior lobe; central lobe of lip 7–11 × 1–1.7 mm, lateral lobes 8–13 mm long; spur 12–17 mm long. Shrublands, forests, 1200–1900 m; Bolívar (headwaters of Río Erebato). Aragua, Distrito Federal, Guárico, Mérida, Miranda; U.S.A. (Florida), southern Mexico, Guatemala, Belize, Costa Rica, West Indies, Ecuador, Brazil. Habenaria dusenii Schltr., Repert. Spec. Nov. Regni Veg. 16: 251. 1919. Terrestrial 12–45 cm tall; leaves distichous, 3.5–7 × 0.3–0.6 cm; inflorescences laxly 1–6-flowered racemes; flowers greenish yellow, medium-sized; dorsal sepal 5–6.2 × 3.5–5.5 mm; lateral sepals 7–8 × 3–3.5 mm;

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posterior lobe of the petals 5–6 × 1.5–3 mm, anterior 10–12 × 0.5–0.6 mm; central lobe of lip 8–9 × 1–1.3 mm; lateral lobes 11–14 × 0.5–0.6 mm; spur 2.4–3.2 cm long. Shrublands, 50–600 m; Bolívar (Cerro Guacamaya), Amazonas (Puerto Ayacucho, Río Siapa, near Santa Bárbara del Orinoco). Suriname, French Guiana, Brazil. The illustration of this species in Foldats (Flora de Venezuela 15(1): 56. 1969) is a composite of a plant similar to Habenaria leprieuri and idealized flowers of H. dusenii based on the original description and illustration of this species. The collection cited in Foldats (1969, 55) seems to be related to, but not conspecific with, H. leprieuri. That plant features petals and sepals similar to H. leprieuri but with a much longer spur. More material of this taxon is required to ascertain its identity. Habenaria dusenii is included in this treatment on the basis of subsequent collections from other localities in Bolívar and Amazonas. Habenaria ernestii Schltr., Notizbl. Königl. Bot. Gart. Berlin. 6: 122. 1914. Habenaria caldensis auct. non Kraenzl.: sensu Dunst. & Garay, Venez. Orch. Ill. 5: 130. 1972, pro parte. Habenaria rodeiensis auct. non Barb. Rodr.: sensu Dunst. & Garay, Orchids Venez. Ill. Field Guide 345. 1979. Terrestrial 30–90 cm tall; leaves distichous, 6–12(–16) × 1.5–2.5 cm; inflorescences laxly to ± densely many-flowered racemes (usually with > 7 flowers in well-developed plants); flowers medium-sized; dorsal sepal 6.5–9 × 4.5–9 mm; lateral sepals 7–10 × 3–5 mm; petals with posterior lobe 4–6.5(–8) × 2.3–3.5 mm, anterior lobe 3–6.5 × 0.5–0.7 mm; lip basally simple and somewhat dilated, 2.5–3 × 2–2.5 mm, central lobe 6–9.7 × 1–2 mm, lateral lobes 6.5–9.5 × 0.5–0.7 mm; spur 23–50 mm long. Open forests, (100–) 800–2100 m; Bolívar (Auyán-tepui, Gran Sabana, Roraima-tepui, Sierra Parima), Amazonas (near Santa Bárbara del Orinoco, Sierra de la Neblina). Mérida(?), Táchira(?); Guyana, French Guiana. Habenaria ernestii is one of the most widespread species of the genus in the flora area and in the Guianas. To determine its actual distribution, the complex of species to which it belongs needs to be studied carefully. It is similar and closely related to H.

rodeiensis (with which it has been confused). Habenaria rodeiensis has, among other differences, longer lateral lip lobes. Habenaria floribunda Lindl., Gen. Sp. Orchid. Pl. 316. 1835. Habenaria petalodes auct. non Lindl. 1835: sensu Dunst. & Garay, Venez. Orchid. Ill. 1: 165. 1959. Terrestrial, exceptionally a subepiphyte, 40–180 cm tall; leaves distichous, 5–30 × 1.5–5.3 cm; inflorescences laxly to densely many-flowered racemes; flowers mediumsized; dorsal sepal 4.5–10 mm long and wide; lateral sepals 6–12 × 3.2–7 mm; petals simple or nearly so, 3.7–11 × 1.5–6 mm; lip simple or basally with a small tooth or auricle at both sides, 7–16 × 0.5–1.5 mm; spur 10–30 mm long. Humid gallery forests, 100– 1900 m; Bolívar (Canaima, La Escalera, Macizo del Chimantá [Apacará-tepui], Río Venamo). Aragua, Distrito Federal, Guárico, Miranda, Portuguesa, Sucre, Yaracuy, Zulia; U.S.A. (Florida), Mexico, Brazil, Paraguay. Habenaria heptadactyla Rchb. f., Linnaea 22: 812. 1849. —Habenaria leprieuri Rchb. f. var. heptadactyla (Rchb. f.) R.E. Schult., Bot. Mus. Leafl. 17(7): 193. 1956. Habenaria viridiaurea Lindl. ex Kraenzl., Bot. Jahrb. Syst. 16: 102. 1892, in syn. Habenaria leprieuri auct. non Rchb. f. 1846: sensu Foldats in Lasser, Fl. Venez. 15(1): 69. 1969, pro parte; Dunst. & Garay, Venez. Orchid. Ill. 5: 134. 1972. Terrestrial to 30 cm; leaves filiform, 2–3 cm long; inflorescences short, cylindrical, 4– 10-flowered, the flowers distichous, bright yellow or greenish; sepals 2.5–3.2 mm long; anterior lobes of the petals and lateral lobes of the lip reflexed, relatively broad; spur 1.5– 2 cm long. Savannas, 50–1500 m; Bolívar (Gran Sabana), Amazonas (near Puerto Ayacucho). Monagas, Anzoátegui; Panama, Colombia, Trinidad, Guyana, Suriname, Brazil, Bolivia. ◆Fig. 351. Previously, Habenaria heptadactyla has been included in the synonymy of H. leprieuri, but it is amply distinct (see key). Habenaria huberi Carnevali & Morillo, Ernstia 19: 6. 1983. Terrestrial; flowers large for the genus, yellow-orange; lateral lobes of lip very wide

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in comparison with other Venezuelan species. Growing on flooded savannas, 100–300 m; Amazonas (Río Culebra, Río Parú). Endemic. ◆Fig. 350. This species is closely related to Habenaria pratensis (Lindl.) Rchb. f. var. pratensis but the colors are different (petals and lip bright orange in H. huberi and pale yellow with green bases in H. pratensis) and the lip lobes are broader in H. huberi. Habenaria pratensis var. pratensis is a species of dry savanna environments while H. huberi occurs in flooded savannas. Habenaria lehmanniana Kraenzl., Bot. Jahrb. Syst. 16: 97. 1892. Habenaria caldensis auct. non Kraenzl. 1892: sensu Foldats in Lasser, Fl. Venez. 15(1): 49. 1969, pro parte; sensu Dunst. & Garay, Venez. Orchid. Ill.5: 130. 1972, pro parte. Terrestrial; leaves bluish green; anterior lobe of the petals as long as the posterior lobe; all three lobes of lip about equally long; sepals usually dark green to olive-green, petals and lip light green or whitish with some greenish. savannas and rocky places 1200– 2200 m; Bolívar (Auyán-tepui, La EscaleraCerro Venamo general zone, Ptari-tepui). Colombia, Guyana, French Guiana. According to Renz (1992), Habenaria lehmanniana can be confused with H. caldensis, which has smaller flowers and narrower leaves. In Habenaria caldensis the anterior lobes of the petals are longer than the posterior lobes and the lateral lobes of the lip are distinctly longer than the midlobe. More critical analysis of the material in the Habenaria ernestii–H. caldensis– H. lehmanniana complex is required to ascertain how many of these taxa actually occur in the flora area. The three species have been confused in the past, as the convoluted putative synonymy indicates. Here we are following Renz’s lead in recognizing three taxa in the area distinguished by the characters in the key. Habenaria leprieuri Rchb. f., Linnaea 19: 376. 1846. Habenaria culmiformis Schltr., Beih. Bot. Centralbl. 42(2): 70. 1925. Terrestrial 20–60 cm tall; leaves 1–7, distichous, 1–10 × 1–3 mm; inflorescences fewto many-flowered racemes; flowers small,

pale green, yellowish green, or yellowish; dorsal sepal 2.5–3.8 × 1.5–3 mm; lateral sepals 2.8–5 × 1.2–1.9 mm; posterior lobe of the petals 1.7–3.2 × 0.5–1.3 mm; anterior 0.5–4 × ca. 0.8 mm; central lobe of lip 2–5 × 0.6–1.1 mm, lateral lobes originating from the base of the lip, 2–4.4 × 0.4–0.9 mm; spur 5–15 mm long. Shrublands, savannas, 50–1100 m; Bolívar (Gran Sabana), Amazonas (La Esmeralda, near Puerto Ayacucho, Santa Bárbara del Orinoco). Anzoátegui; Panama, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil. As here defined, Habenaria leprieuri is a variable species. Material with narrower, filiform anterior lobe to the petals (0.1–0.2 mm) and lip lateral lobes (0.1–0.2 mm) has been annotated as H. culmiformis in the past. This species is here treated as a synonym of H. leprieuri. Habenaria longicauda Hook., Bot. Mag. t. 2957. 1829. Habenaria sartor auct. non Lindl. 1843: sensu Foldats in Lasser, Fl. Ven. 1: 93. 1969. Subaquatic 60–150 cm tall; leaves distichous, 8–27 × 1–2.5 cm; inflorescences racemes; flowers large; dorsal sepal 14–20 × 9–13 mm; lateral sepals 15–23 × 6–12 mm; petals with posterior lobe 15–20 × 1.8–4 mm, anterior lobe 22–30 × 0.5–1.5 mm; lip basally not divided, 4–7 × 2.5–3.5 mm, central lobe 14–22 × 1.2–3 mm, lateral lobes 20–32 × 0.9– 1.4 mm; spur 7–14 cm long. Mauritia palm swamps, edges of rivers and lagoons, 50– 1200 m; Delta Amacuro (Caño Acoimo near Río Orocoima), Bolívar (near Caicara del Orinoco, near Ciudad Piar, Serranía de los Pijiguaos), Amazonas (near Puerto Ayacucho, Sierra Parima). Guárico, Táchira, Zulia; Ecuador, Brazil. Habenaria macronectar (Vell.) Hoehne (syn. H. sartor Lindl.) from Brazil and H. pringlei B.L. Rob., from Mexico to Costa Rica, are closely related but in these taxa the spur is only about half as long. Habenaria mesodactyla Griseb., Fl. Brit. W. I. 664. 1864. Terrestrial 21–56 cm tall; leaves few, distichous, 3–8 × 1–4 mm; inflorescences fewflowered racemes; flowers relatively small; dorsal sepal 3–7.7 × 2–5 mm; lateral sepals 3.5–8.2 × 1.5–3.1 mm; posterior lobe of petals

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3–6 × 1–2 mm, anterior 3.5–8.5 × 0.3–0.6 mm; central lobe of lip 3.5–7.5 × 0.9–1.2 mm, lateral lobes 5–10 × 0.3–0.6 mm; spur 9–15 mm long. Humid savannas, shrublands, 1000–1300 m; Bolívar (El Dorado to Cerro Venamo, Gran Sabana). Miranda; Southern Mexico, Belize, Honduras, West Indies. Habenaria monorrhiza (Sw.) Rchb. f., Ber. Deutsch. Bot. Ges. 3: 274. 1885, non Cogn. 1909. —Orchis monorrhiza Sw., Prodr. 118. 1788. Habenaria maculosa Lindl., Gen. Sp. Orchid. Pl. 309. 1835. Habenaria speciosa Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 44. 1836. Terrestrial 20–120 cm tall; stems and sheaths basally spotted; leaves distichous, with membranous-scarious margins; blades 4–15 × 1.5–4 cm; inflorescences racemes; flowers small to medium-sized; sepals 5–14 mm long; dorsal sepal 4–7 mm wide; lateral sepals 3–6 mm wide; posterior lobe of the petals 5–13 × 1.5–3 mm, anterior lobe extremely variable, from reduced to a tooth to linear-filiform and much longer than the posterior one; central lobe of lip 6.5–15 mm long; lateral lobes linear-filiform, 7–16 mm long; spur 15–30 mm long. Savannas, humid forests, 1200–2200 m; Bolívar (Auyán-tepui, Cerro Venamo). Widespread in Neotropics. This species is widely ranging and extremely variable. A careful study of material referred to it may demonstrate that several taxa (closely related species or subspecific ranks) may warrant recognition within the broad concept of Habenaria monorrhiza taken here. Habenaria nilssonii Foldats, Bol. Soc. Venez. Ci. Nat. 100: 256. 1962. Terrestrial 28–80 cm tall; leaves distichous, 3–9 × 0.5–2 cm; flowers mediumsized; inflorescences few- to many-flowered racemes; dorsal sepal 4.3–6.5 × 3.3–6 mm; lateral sepals 4.8–7.5 × 2–4.5 mm; petals with the principal lobe 4–5 × 2–4 mm, anterior one to 2 mm long, posterior shorter than the anterior one; central lobe of lip 5–6.5 × 1– 1.2 mm, lateral lobes 3.5–4 × 0.5–0.6 mm; spur 10–35 mm long. On sandstone, 1200– 2200 m; Bolívar (Auyán-tepui, Luepa to Cerro Venamo). Endemic.

Habenaria obtusa Lindl., Gen. Sp. Orchid. Pl. 315. 1835. Terrestrial 30–115 cm tall; leaves distichous, 5–25 × 1.5–4 cm; inflorescences manyflowered racemes; bracts foliaceous, 2–6 × 0.6–2.2 cm; flowers relatively large; dorsal sepal 7–12 × 6–10 mm; lateral sepals 10–14 × 1.9–3 mm; petals simple, 6–10 × 2–4 mm; lip simple, 10.6–18 × 1.9–3 mm; spur 3.7–7 cm long. Savannas, sometimes in fire zones, forests, 100–2500 m; Bolívar (Canaima, La Escalera, Sabana de Arecuna, near Santa Elena de Uairén). Widespread elsewhere in Venezuela; Colombia, Suriname, Peru, Brazil, Paraguay. Habenaria parviflora Lindl., Gen. Sp. Orchid. Pl. 314. 1835. Habenaria demerarensis Rchb. f. ex M.R. Schomb., Reis. Br.-Guiana 3: 1123. 1848. Habenaria amalfitana F. Lehm. & Kraenzl., Bot. Jahrb. Syst. 16: 113. 1892. Habenaria arecunarum Schltr., Notizbl. Königl. Bot. Gart. Berlin 6: 121. 1914. Terrestrial 10–130 cm tall; leaves distichous, 4–25 × 0.2–1 cm; inflorescences racemes few- to many-flowered; flowers small; dorsal sepal 2.5–5 × 1.5–4 mm; lateral sepals 3–6 × 1.5–2.5 mm; petals with posterior lobe 2–4 × 1–1.7 mm, anterior lobe 0.6– 1.5 × 0.3–0.4 mm; central lobe of lip 2.8–5 × 0.8–1.3 mm, lateral lobes 2–5 × 0.2–0.5 mm; spur 4.5–12 mm long. Dry or humid savannas, 1100–2800 m; Bolívar (Auyán-tepui, Río Tírica, Roraima-tepui, Sororopán-tepui, Ueitepui). Apure, Distrito Federal, Miranda, Yaracuy; Colombia, Guyana, Brazil, Paraguay, Argentina, Uruguay. Habenaria parviflora is another widely ranging and variable species. The flowers, which are among the smallest in the genus, are borne in denser, more floriferous spikes toward the southern end of its range (southeastern Brazil, Argentina, and Uruguay) than they are in northern South America. Habenaria arecunarum and H. demerarensis are typical examples of forms within this broad circumscription of this species found in its northernmost range. Habenaria amalfitana, with its papillose end to the lateral lobes of the lip, might deserve recognition, but this form does not seem to occur in

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the flora area. Several taxa might be involved. Habenaria pratensis (Lindl.) Rchb. f., Linnaea 22: 813. 1949. —Bonatea pratensis Lindl., Gen. Sp. Orchid. Pl. 328. 1835. Suriname, French Guiana, Brazil; 2 varieties, one in Venezuela. H. pratensis var. parviflora Cogn. in Mart., Fl. Bras. 3(4): 86. 1893. Habenaria georgii Schltr., Beih.Bot. Centralbl. 42: 86. 1925. Erect terrestrial to 40 cm tall; flowers fragrant; sepals light green, petals and lip yellow with green bases. Bolívar (cited by J. Renz 1992). We have not seen authentic material of Habenaria pratensis var. parviflora from Venezuela, although the entity here treated as Habenaria sp. A might be conspecific or closely related. This entity is, however, readily distinguished by its petals with a tooth on the posterior margin of the posterior lobes. As conceived by Renz (1992), this variety seems distinct enough from Habenaria pratensis var. pratensis to warrant recognition at the specific level. Should it be recognized as a different species, the name H. georgii is available. Habenaria pygmaea C. Schweinf. & R.E. Schult., Amer. Orchid Soc. Bull. 23: 822. 1954. Habenaria guanchezii Carnevali & I. Ramírez, Ernstia 33: 14. 1985. Terrestrial or subterrestrial, growing on sand in shady places, 7–17 cm tall; leaves basal; flowers pale green; petals usually simple, sometimes 2-lobed. Scrublands, dwarf open and humid forests, 100–200 m; Amazonas (Río Guayapo). Amazonian Colombia and Ecuador. ◆Fig. 348. Habenaria quadrata Lindl., Gen. Sp. Orchid. Pl. 316. 1835. Terrestrial 30–90 cm tall; leaves distichous, 5–12 × 1.5–3 cm; inflorescences racemes, laxly many-flowered; dorsal sepal 4–4.5 × 4–4.3 mm; lateral sepals 5–6 × 3–4 mm; petals simple, 3–3.2 × 1.7–2 mm; lip

simple, 6–8 × 1–1.7 mm; spur 14–17 mm long. Forests, savannas, ca. 100 m; Bolívar (near Caicara), Amazonas (Raudal de Atures). Brazil. Habenaria quinqueseta (Michx.) Sw., Adnot. Bot. 46. 1829. —Orchis quinqueseta Michx., Fl. Bor.-Amer. 2: 155. 1803. Terrestrial 20–90(–105) cm tall; leaves distichous, 5–25 × 2–7 cm; flowers mediumsized; dorsal sepal 6–12 × 5–10 mm; lateral sepals 8–16 × 4–7 mm; petals with posterior lobe 6–15 × 2–4 mm, anterior lobe 13–30 × 0.6–1 mm; central lobe of lip 8–20 × 2–3 mm; lateral lobes 15–30 × 0.5–1 mm; spur 4–18 cm long. Flooded places, ca. 100 m; Bolívar (Río Caroní). Aragua, Carabobo; U.S.A., Mexico(?), Central America(?), West Indies, Guyana, Suriname. We have not seen authentic material of Habenaria quinqueseta from the flora area. The related H. macroceratitis Willd. has much longer anterior lobes on the petals, and, on the average, longer spurs. The two taxa are often treated as conspecific. The illustration in Foldats (1969, 88) seems to represent H. macroceratitis due to its extremely long anterior lobes to the petals. Habenaria repens Nutt., Gen. N. Amer. Pl. 2: 190. 1818. Subaquatic 30–100 cm tall; leaves distichous, (5–)8–26 × (0.4–)1–2.6(–3) cm; dorsal sepal 2.8–7.5 × 2.2–6.5 mm; lateral sepals 3.5–8 × 2–4 mm; petals with posterior lobe 2.5–7 × 0.8–2 mm, anterior lobe filiform or linear-filiform, 3–7.5 mm long; central lobe of lip 3.2–7 × 0.7–1.2 mm, lateral lobes filiform or linear-filiform, 3.5–11 mm long; spur 6–11 mm long. Flooded savannas, ca. 100 m; Delta Amacuro (Sacupana), Bolívar (Río Caroní, near Tumeremo). Apure, Distrito Federal, Mérida, Monagas, Táchira, Yaracuy; widespread in Neotropics. ◆Fig. 349. Over its enormous range in the tropics and subtropics of the New World, Habenaria repens is remarkably consistent in its overall morphology, despite being variable in vegetative size. It ranges from near sea level to elevations above 2500 m, always growing in standing water.

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Habenaria roraimensis Rolfe, Trans. Linn. Soc. London, Bot. 6: 65. 1901. Terrestrial 17–70 cm tall; leaves distichous, reduced and somewhat bract-like, 2–4 × 0.5–0.8 cm; flowers small; inflorescences few- or many-flowered racemes; dorsal sepal 3.5–6 × 3–5 mm; lateral sepals 4.2–7 × 2–3 mm; petals with posterior lobe 2.5–5.5 × 1.5– 2.5 mm, anterior lobe very short to 4.2 × 0.3 mm; central lobe of lip with recurved margins, 4.2–6.3 × 1–1.5 mm, lateral lobes 3.2–7 × ca. 0.3 mm; spur 6–13 mm long. Savannas, shrublands, (800–)1500–2800 m; Bolívar (Auyán-tepui, Kukenán-tepui, La Escalera, Roraima-tepui), Amazonas (Cerro Marahuaka, Sierra de la Neblina). Guyana, Brazil. Habenaria roraimensis is endemic to the high tepuis in the flora area and adjacent portions of Brazil and Guyana. Material included in this species in Foldats (1969, 91– 93) is referable to H. gollmeri Schltr. and other related taxa. The bluish tinge of the whole plant and the reduced leaves are diagnostic. Habenaria schomburgkii Lindl., London J. Bot. 2: 673. 1843. Terrestrial 23–125 cm tall; leaves distichous, (5–)8–20 × (0.5–)0.8–1.3 cm; inflorescences 3–7-flowered, usually racemes; flowers medium-sized; dorsal sepal 5–13 × 4–9 mm; lateral sepals 5–14 × 3–5 mm; petals with posterior lobe 4.5–10 × 1.5–3 mm, anterior lobe 9–20 mm long; central lobe of lip 7– 12 × 0.7–3 mm, lateral lobes filiform, 9–20 mm long; spur (2–)8–32 mm long. Savannas, shrublands, 1100–1500 m; Bolívar (La Escalera, near Santa Teresita de Kavanayén). Guyana, Suriname, Brazil. Habenaria setacea Lindl., Gen. Sp. Orchid. Pl. 312. 1835. Guyana, French Guiana, Brazil; 2 varieties, both in Venezuela, 1 in the flora area. H. setacea var. setacea Terrestrial; leaves grass-like; inflorescences narrow, 5–7-flowered. Savannas, 1200 m; Bolívar (Kukenán-tepui); Guyana, French Guiana, Brazil. Habenaria setacea var ecalcarata Renz was described from Táchira and is distinguished from the typical variety by its spurless lip.

Habenaria sprucei Cogn. in Mart., Fl. Bras. 3(4): 40. 1893. Habenaria leaoana Schltr., Beih. Bot. Centralbl. 42(2): 72. 1925. Terrestrial 21–32 cm tall; leaves 3–5, distichous, 3–8 × 1–3 mm; inflorescences laxly 2–7-flowered racemes; flowers mediumsized; sepals 7–12 mm long; dorsal sepal 5–6 mm wide; lateral sepals 2.4–4 mm wide; posterior lobe of the petals longer than to slightly shorter than the dorsal sepal, 1.8–2 mm wide; anterior equal to or shorter than the posterior, 0.2–0.5 mm wide; lateral lobes of lip placed somewhat above the base of the lip, 6.5–8 × 0.6–1.2 mm, central lobe as long as or longer than the lateral ones, 1–2 mm wide; spur 10–15 mm long. Riparian, ca. 1300 m; Bolívar (Río Kukenán basin). Guyana, Suriname, French Guiana, Brazil. Habenaria trifida H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 330. 1815 [1816]. Bonatea pauciflora Lindl., Gen. Sp. Orchid. Pl. 329. 1835. —Habenaria pauciflora (Lindl.) Rchb. f., Bonplandia (Hanover) 2: 10. 1854. Terrestrial 20–80 cm tall; leaves few, distichous, 4–20 × 0.5–1.8 cm; inflorescences laxly 1–6-flowered; flowers large; pedicellate ovary of well-developed flowers 4.5–12 cm long; sepals green, petals and lip white or creamy-white; dorsal sepal 7–15 × 4–9 mm; lateral sepals 7.5–16 × 3–6 mm; petals usually bifid, exceptionally trifid, with posterior lobe 5.5–13 × 1.5–4 mm, anterior lobe 4–10 × 0.5–1 mm; central lobe of lip 6–15 × 1–2 mm, lateral lobes 5–14 × 0.6–1 mm; spur 2.8–12 cm long, sometimes apically emarginate. Humid savannas, gallery forests, 1200–1300 m; Bolívar (near Santa Elena de Uairén), Amazonas (Cerro Autana, lower Río Ventuari). Aragua, Carabobo, Guárico, Miranda, Monagas, Zulia; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Paraguay. The authors have never seen the trifid petals often attributed to Habenaria trifida in past literature. Habenaria sp. A Terrestrial, vegetatively similar to Habenaria floribunda; floral bracts elliptic, acute, 8–11 mm long, the margins minutely papillose; dorsal sepal broadly elliptic, ob-

Habenaria 383

tuse, 4.2–4.4 mm long, ca. 3 mm wide; lateral sepals obliquely obovate, rounded apically, 2veined, 7–7.2 mm long, 3.6–3.8 mm long; petals with a linear-oblong posterior lobe, 4.2– 4.3 mm long; anterior lobe pointing downward, tooth-like or somewhat hook-like, ca. 1.4–1.4 mm long, the posterior margin of the posterior lobe at the base with another short, blunt tooth pointing upward (thus petals almost 3-lobed), the anterior margin of the posterior lobe thickened; lip 11.3–1.5 mm long, the midlobe ca. 1 mm wide, apex rounded and thickened, the lateral lobes tooth-like and perpendicular to the midlobe, 1.3–1.45 mm long, thus 3.8–3.9 mm across the spread lateral lobes. Evergreen lowland forests, 100–200 m; Amazonas (near Maroa). This species seems related to a complex of Brazilian species, including Habenaria josephensis Barb. Rodr. and H. gnoma Barb. Rodr., characterized by the dorsal sepal much smaller than the laterals, these obliquely obovate, rounded apically, the lip and petals almost simple with the lateral lobes reduced to teeth-like structures. In Habenaria sp. A, which is almost certainly an undescribed species, the posterior lobe of the petals is linear-oblong and proportionally short as compared to related taxa. In the flora area, the closest relative seems to be H. floribunda, but in that species the petals are simple or almost so, and the posterior lobe is subquadrate. Habenaria sp. B Terrestrial 30–35 cm tall; leaves 4–6, laxly arranged on the stem, 6–8 × 0.8–1.4 cm, narrowly lanceolate, acute, becoming bract-like distally; flowers 8–10 on the stem; sepals green, ca. 8–8.5 mm long, dorsal ca. 8 mm wide, laterals 9–10 × 3.5–4 mm wide, petals lighter green, posterior lobe 8.5–9.5 × 0.8–1.2 mm, anterior lobe ca. 5 × 0.5 mm, tooth or lobe on the posterior margin of the posterior lobe either conical and 2 mm long or linear and ca. 4 mm long, the bases of petals and sepals whitish; lip with midlobe 9–13 mm long, laterals 7–8 × 0.8 mm, the lip undivided for ca. 4 mm; pollinia light yellow; spur ca. 22 mm long. Seepages at edges of lajas and savannas, 50–100 m; Amazonas (Puerto Ayacucho). This distinctive taxon resembles vegetatively Habenaria alata Lindl., but the flow-

ers are strikingly different. The flowers suggest Habenaria pratensis var parviflora in having broad segments and an undivided basal portion of the lip. The floral segments in this entity are, however, entirely green and somewhat smaller, and the petals have the distinctive tooth or lobe on the posterior margin of the posterior lobe, making them 3-lobed.

Fig. 348. Habenaria pygmaea

Fig. 349. Habenaria repens

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Fig. 350. Habenaria huberi

Fig. 351. Habenaria heptadactyla

Helonoma 385

63. HELONOMA Garay, Bot. Mus. Leafl. 28: 327. 1982. —Beloglottis sect. Helonoma (Garay) Szlach., Fragm. Florist. Geobot. 41(2): 702. 1996. [Subtribe Spiranthinae]. Spiranthes Rich., De Orchid. Eur. 20, 28, 36. Aug.–Sept. 1817, nom. cons., pro parte. Cyclopogon C. Presl, Reliq. Haenk. 1: 93. 1827, pro parte. Beadlea Small, Fl. S.E. U.S. 319. 1903, pro parte. Beloglottis Schtr., Beih. Bot. Centralbl. 37(2): 364. 1920, pro parte. by Germán Carnevali, Ivón M. Ramírez-Morillo, and Carlos A. Vargas Terrestrial, humicolous, or ± aquatic herbs, small, erect, acaulescent. Roots thick, pubescent. Leaves thin, convolute in bud, broadly elliptic, ovate, or suborbicular, apex obtuse, rounded, or acute, basally attenuate into a ± well-developed pseudopetiole, frequently with darker green veins, forming a basal rosette. Inflorescence racemose, terminal, erect, longer than leaves, internodes sheathed, lax- or dense-flowered, few- to many-flowered. Flowers small, resupinate, green or yellowish, with a white or yellowish lip; floral bracts small but conspicuous, herbaceous; ovary fusiform, sessile, barely twisted. Sepals variously basally connate, subequal; petals coherent with dorsal sepal, smaller than sepals, sometimes forming a basal spur decurrent along the basal 1/2 of the spatulate column. Lip unlobed or with a hypochile and epichile, basally hastate. Column ± erect, with a frontal longitudinal groove, with a short, oblique foot on top of ovary, base of clinandrium adnate to dorsal sepal; anthers ovate-lanceolate, hooded, acuminate at apex; pollinia 4, linearclavate, viscidium small, round; rostellum short, triangular, obtuse, mostly ruptured or dissolved in autogamous flowers; stigmas 2, commonly free with sides barely touching one another. Endemic to the Venezuelan Guayana Shield; 3 species, all in the flora area. Some authors recently considered Helonoma as a part of Beloglottis. We chose a conservative approach considering them as distinct, but closely related genera. Floral features are similar between the two genera, but a very small habit, association with bogs, and rostellum reductions leading to autogamy are features typical only in Helonoma. Key to the Species of Helonoma 1. 1. 2(1). 2.

Inflorescence laxly 1- or 2-flowered; perianth segments apically entire ................................................................................................ H. americana Inflorescence densely many-flowered; all perianth segments apically bifid ................................................................................................................ 2 Lip ≥ 3 mm long, epichile well developed, as long as the hypochile, lip glands well developed .................................................................... H. bifida Lip ≤ 2.5 mm long, epichile rudimentary, shorter than the hypochile, lip glands inconspicuous ............................................................ H. chiropterae

Helonoma americana (C. Schweinf. & Garay) Garay, Bot. Mus. Leafl. 28: 328. 1982. —Manniella americana C. Schweinf. & Garay, Bot. Mus. Leafl. 20: 5, pl. 2. 1962. —Beloglottis americana

(C. Schweinf. and Garay) Szlach., Fragm. Florist. Geobot. 41(2): 702. 1996. Humicolous, small, erect, yellowish or greenish. Montane forests, tepui summits, usually on moist rocks along streams or in

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boggy places in sun or semishade, 500–2000 m; Bolívar (Amaruay-tepui, La Escalera to Cerro Venamo region), Amazonas (Cerro Aracamuni, Cerro Autana). Endemic. ◆Fig. 352.

Fig. 352. Helonoma americana

Helonoma bifida (Ridl. ex Thurn) Garay, Bot. Mus. Leafl. 28: 328. 1982. —Spiranthes bifida Ridl. ex Thurn, Timehri 5: 205. 1886. —Spiranthes bifida Ridl. ex Oliv., Trans. Linn. Soc. London, Bot. 2: 283. 1887, nom. superfl. —Cyclopogon bifidus (Ridl. ex Oliv.) Schltr., Beih. Bot. Centralbl. 37(2): 386. 1920. —Beadlea bifida (Ridl. ex Oliv.) Garay & Dunst. in Dunst. & Garay, Orchids Venez. Ill. Field Guide 23. 1979. —Beloglottis bifida (Ridl. ex Oliv.) Szlach., Fragm. Florist. Geobot. 41(2): 702, fig. 1. 1996. Terrestrial or muscicolous, small to medium-sized; racemes 15–50 cm tall; sepals green; petals and lip white. Shrublands or bogs on tepui summits, 1600–2500 m; Bolívar (Aprada-tepui, Kamarkawarai-tepui, Kukenán-tepui, Macizo del Chimantá, Ptari-tepui), Amazonas (Sierra de la Neblina). Endemic. Helonoma chiropterae (Szlach.) Carnevali & G.A. Romero in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1136. 2000. —Beloglottis chiropterae Szlach., Fragm. Florist. Geobot. 41(2): 703, fig. 2. 1996. Beadlea bifida (Ridl.) Garay & Dunst., non Spiranthes bifida Ridl. 1886: sensu Dunst. & Garay, Orchids Venez. Ill. Field Guide 23. 1979. Herb 20–30 cm tall; leaves very small, 21–29 × ca. 8 mm; spike 3 cm long, 20–25flowered; dorsal sepal 2.8–0.7 mm; lateral sepals ca. 3 × 0.8 mm; petals 2.2–0.8 mm; lip ca. 2.5 × 1.6 mm. Bogs on tepui summits, ca. 2500 m; Amazonas (Cerro Marahuaka). Endemic. ◆Fig. 353. Szlachetko (Fragm. Florist. Geobot. 41(2): 702. 1996) noticed that Garay and Dunsterville (G. C. K. Dunsterville & L. Garay, Orchids of Venezuela: An Illustrated Field Guide 1: 23. 1979) included a drawing of Helonoma bifida (Ridl.) Garay [Dunsterville 1157 AMES, line drawing, published as Beadlea bifida (Ridl.) Garay & Dunst.], which differs from the type of Spiranthes bifida Ridl. (Im Thurn 342, K).

Fig. 353. Helonoma chiropterae

Hexisea 387

64. HEXISEA Lindl., J. Bot. (Hooker) 1: 7. 1834. [Subtribe Laeliinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes or lithophytes, cespitose, erect, or arching to subpendulous when mature, mostly sun-loving. Roots many, thin; rhizomes short. Pseudobulbs heteroblastic (one internode) or homoblastic (more than one internode), spindle-shaped or narrow-oblong, terete or subquadrangular, frequently ridged, often reddish or purplish, 1–3-foliate apically, basally enveloped by several imbricate, scarious, eventually deciduous sheaths, apically producing more pseudobulbs and forming chains with 2–8 pseudobulbs, sometimes apical pseudobulbs producing roots basally. Leaves articulate, conduplicate, coriaceous, suberect or spreading, subopposite on top of stem due to abbreviation of apical pseudobulb internodes, sessile; blades oblong or oblong-elliptic, apex rounded, truncate, or emarginate. Inflorescences fewflowered, dense racemes, apical from and much shorter than upper pseudobulbs; peduncle short, enveloped by scarious sheaths. Flowers resupinate, several or all opening simultaneously, showy, bright red to orange; bracts conspicuous, scarious, mostly lanceolate; pedicellate ovary terete, often reddish or pale orange. Perianth segments campanulate to spreading; sepals lanceolate to oblong, acute to obtuse, subequal, free, or the lateral ones ± connate at the base forming a small mentum, petals similar to sepals but ± smaller or narrower. Lip fused in its basal 1/3 to the column, the fused lateral margins and the ventral face of the column forming a cup or deep sack; free portion of lip dilated into a simple, oblong-elliptic to subspatulate blade, strongly geniculate at the base; disk almost ecallose or bicallose. Column short and thick; anther operculate, incumbent, 2-locular; clinandrium 3-lobed, lateral lobes larger; pollinia 4, with yellow caudicles; rostellum transverse; stigma ventral. Capsules suborbicular to ellipsoid. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil; 3 species, 2 in Venezuela, both in the flora area. Recent phylogenetic analyses suggest that Hexisea would be better treated as congeneric with Scaphyglottis Poepp. & Endl. The differences in column-lip structure are adaptations to hummingbird pollination in Hexisea Key to the Species of Hexisea 1. 1.

Pseudobulbs terete in cross section; dorsal sepal oblong; lip obtuse or truncate .................................................................................... H. bidentata Pseudobulbs subquadrangular in cross section; dorsal sepal elliptic; lip acute ........................................................................................ H. imbricata

Hexisea bidentata Lindl., J. Bot. (Hooker) 1: 8. 1834. Epiphyte, rarely a lithophyte; pseudobulbs with many shallow ridges, often purplish; lip basally with 3 or 5 keels. Locally common in rain forests or open humid associations, 100–1500(–1900) m; widespread in Bolívar and Amazonas. Miranda, Táchira; other distribution as in genus.

Hexisea imbricata (Lindl.) Rchb. f. in Walp., Ann. Bot. Syst. 6: 470. 1862. —Diothonea imbricata Lindl., Sert. Orchid. pl. 40, fig. 1. 1841. —Hexisea bidentata var. imbricata (Lindl.) C. Schweinf., Bot. Mus. Leafl. 15: 106. 1951. Epiphyte; pseudobulbs with few, deep ridges, often purplish; lip longitudinally 2-

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callose at base. Lower montane forests, humid open places, 200–1300 m; Bolívar (Auyán-tepui, Cerro Tonoro, upper Río Caura basin near Santa Elena de Uairén, Serranía Marutaní), Amazonas (Cerro Autana, Cerro Duida, Cerro Marahuaka, Cerro Yapacana). Southern Mexico, Central America, Colombia. ◆Fig. 354.

Fig. 354. Hexisea imbricata

65. HOULLETIA Brongn., Ann. Sci. Nat. Bot. sér. 2, 15: 37. 1841. [Subtribe Stanhopeinae]. by Gustavo A. Romero-González Cespitose epiphytes or terrestrial herbs. Stem modified into ovoid to cylindric, shallowly sulcate pseudobulbs of 1 internode, when young entirely concealed by distichous, scarious sheaths, eventually naked. Leaf 1, lanceolate, acute, plicate, articulate, long-petiolate. Inflorescences lateral, arising from the base of the pseudobulbs, erect or pendent, racemose, few- to many-flowered. Flowers fleshy, waxy, resupinate, sometimes pendulous, showy; floral bracts concave, narrowly elliptic to obovate, shorter than the pedicellate ovary. Sepals and petals spreading; sepals concave, broadly elliptic to ovate, the lateral ones basally connate or not, oblique or not, adnate to the column foot; petals narrowly elliptic, sometimes slightly lobed or spatulate, generally smaller than the sepals. Lip fleshy, subsessile, divided into a hypochile, 3-lobed, sometimes concave, the lateral lobes very narrowly ovate to elliptic, sometimes falcate, the midlobe antrorse, subtriangular to truncate, and an epichile, adnate to the apical underside of the hypochile, sometimes articulate, simple or auriculate to 3-lobed, truncate, the lateral lobes narrowly triangular to elliptic, apiculate. Column erect, recurved, semiterete, sometimes produced into a very short foot; anther terminal, operculate, incumbent, 1-locular; pollinarium with 2 yellow pollinia, narrowly obtriangular to obovate, cleft, cartilaginous, and dorsoventrally flattened, the tegula long, linear, the viscidium narrowly triangular. Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 10 species, 3 in Venezuela, 2 of these in the flora area.

Houlletia 389

Key to the Species of Houlletia 1.

1.

Flowers facing downward; petals narrowly obovate; lip with an erect, fleshy, column-like basal callus, the lateral lobes narrowly triangular, falcate ................................................................................. H. odoratissima Flowers facing forward; petals elliptic; lip without an erect, column-like basal callus, the lateral lobes elliptic, apiculate ................. H. roraimensis

Fig. 355. Houlletia odoratissima

Fig. 356. Houlletia roraimensis

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O RCHIDACEAE

Houlletia odoratissima Linden ex Lindl. in Lindl. & Paxton, Paxt. Fl. Gard. 3: 172. 1853. Terrestrial herb; inflorescence erect, to 16-flowered; flowers fleshy, somewhat pendulous; sepals, petals, and lateral lobes of lip deep maroon, rest of lip and column white. Dense, shaded thickets on tepui summits, (1000–)1500–2000 m; Bolívar (Auyán-tepui, Cerro Jaua, Ilú-tepui, La Escalera), Amazonas (Cerro Duida, Cerro Sipapo). Colombia, Ecuador, Peru, Brazil (Roraima-tepui), Bolivia. ◆Fig. 355.

Houlletia roraimensis Rolfe, Trans. Linn. Soc. London, Bot. 6: 63. 1901. Terrestrial herb; inflorescence erect; sepals and petals greenish yellow with dull lavender spots; lip rich yellow or yellow spotted with purple. Forested slopes, Bonnetia forests on tepui summits; 1900–2300 m; Bolívar (Auyán-tepui, Macizo del Chimantá [Abacapá-tepui, Toronó-tepui], Ptari-tepui, Roraima-tepui), Amazonas (Cerro Sipapo). Guyana, Brazil (Roraima-tepui). ◆Fig. 356.

66. HUNTLEYA Bateman ex Lindl., Edwards’s Bot. Reg. 23: sub t. 1991. 1837. [Subtribe Zygopetalinae]. by Gustavo A. Romero-González Cespitose or rhizomatous, epiphytes, occasionally subterrestrial herbs. Stem abbreviate, not modified into pseudobulbs. Leaves numerous, distichous, in a flabelliform cluster, coriaceous, narrowly linear-ovate, narrowly oblong-ovate, or oblong-obovate, acute to acuminate, plicate, articulate. Inflorescences lateral, arising from the axil of the leaf sheaths, erect to slightly arching, 1-flowered, < to almost as long as the leaves. Flowers fleshy, star-shaped, resupinate, showy; floral bracts concave, appressed, ovate, < 1/2 as long as the pedicellate ovary. Sepals and petals similar, spreading, narrowly ovate to ovate, acuminate; lateral sepals slightly oblique, adnate laterally to the apex of the column foot. Lip clawed, articulate with the apex of the column foot, entire to ± 3-lobed, acuminate; disk with an elevated, semicircular or subquadrate callus, radially keeled, marginally fimbriate or laciniate. Column elongate, semiterete, produced into a short foot, apically winged; anther terminal; pollinia 4, yellow, cartilaginous, dorsoventrally flattened, clavate, attached in 2 pairs to a subtriangular or oblong tegula. Central America, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 6 species, 2 in Venezuela, 1 of these in the flora area. Huntleya meleagris Lindl., Edwards’s Bot. Reg. 23: sub t. 1991. 1837. Huntleya albido-fulva Lem., Ill. Hort. 15: t. 556. 1868. Fan-shaped epiphyte without pseudobulbs; sepals and petals fleshy, yellowish white, waxy; lip beige with some green in the

margins and a fimbriate crest near the base. Wet forests, 600–900 m; Bolívar (Altiplanicie de Nuria, Icabarú). Táchira; Nicaragua, Costa Rica, Panama, Colombia, Trinidad-Tobago, Guyana, Ecuador, Brazil (widespread). ◆Fig. 357.

Huntleya

391

5 cm

5 cm

1 cm Column and lip

5 mm Anther and Pollinia

10 cm Fig. 357. Huntleya meleagris

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O RCHIDACEAE

67. HYLAEORCHIS Carnevali & G.A. Romero in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1136. 2000. [Subtribe Bifrenariinae]. Bifrenaria Lindl., Gen. Sp. Orchid. Pl. 152. 1833, pro parte. Maxillaria Ruiz & Pav., Fl. Peruv. Prodr. 116, pl. 25. 1794, pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytic herbs, shade-loving. Rhizome shortly creeping, appressed to the host, completely enveloped by imbricate, scarious sheaths. Pseudobulbs pearshaped to narrowly ovoid, rarely somewhat ellipsoid, apically 1-leaved, entirely clothed by 3 or 4 imbricating, tubular sheaths, the sheaths scarious, red-brown when dry, somewhat pustulose, disintegrating in place, the longest much longer than the pseudobulb and enveloping the basal half of the petioles. Leaves with convolute vernation, petiolate; blades fleshy- coriaceous when fresh, coriaceous when dry, subplicate, oblong-elliptic to narrowly elliptic, acute to subacuminate, with 3 primary and 2 secondary veins at each side of the midvein, margins sometimes microscopically erose-ciliate, somewhat revolute, crenate at the apical 1/10–1/7 (sometimes only at one side of the midvein) due to the apparent termination of the lateral veins close to the margins; petiole variable in length, from much shorter than the blade to about half its length, terete, channeled distally. Inflorescences originating from pseudobulb base, abbreviate and almost completely hidden by the sheaths of the pseudobulbs, 1-flowered in small plants, in larger plants long-lasting and consisting of a continuously growing rachis bearing 1-flowered branches at internodes, in mature plants branching from the basal internodes. Flowers resupinate, of small size for the subtribe; pedicellate ovary subterete; floral bract shorter than pedicels; perianth segments free, subfleshy, maroon, maroon-purple, or wine red; tubular; lateral sepals inserted along the sides of the column foot, producing a prominent mentum; lip subentire to shallowly 3-lobed about middle, articulate with the column foot, apically truncate, margins microscopically ciliate; disk with a low longitudinal, oblong callus with dark maroon, shiny surface that appears wet. Column slightly clavate, basally produced into a pronounced foot; clinandrium subentire or finely ciliolate; anther apical, operculate, papillose; pollinia in two unequal pairs, viscidium thin, horseshoe-shaped. Capsule obovoid or ellipsoid, deeply ridged. Colombia, Amazonian Venezuela, Ecuador, Peru, Brazil; 1 species. Hylaeorchis differs from Bifrenaria in its successively 1-flowered, perennial inflorescence, its pear-shaped, nonangulate pseudobulbs, its horseshoe-shaped stipe and viscidium, and 2 unequal pairs of pollinia. The flowers are among the smallest in the subtribe Bifrenariinae and strongly resemble flowers in the genus Maxillaria Ruiz & Pav. This similarity has resulted in the repeated description or placement of this taxon in Maxillaria. However, the leaves clearly display convolute vernation, placing it closest to the Bifrenaria complex. The short column foot is unusual in the subtribe Bifrenariinae. Recent cladistic analyses of DNA sequence variation confirm the uniqueness of Hylaeorchis since they identify this taxon as the sister group of the anomalous genus Teuscheria Garay (N. Williams and M. Whitten, personal communication). Teuscheria, however, features subspherical pseudobulbs and spurred lip. Hylaeorchis petiolaris (Schltr.) Carnevali & G.A. Romero in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1136. 2000. —Maxillaria petio-

laris Schltr., Beih. Bot. Centralbl. 42(2): 133. 1925. —Maxillaria rudolfii Hoehne, Arq. Bot. Estado São Paulo 2(4): 72: 1947, nom. nov. superfl., non A. Rich.

Hylaeorchis 393

1882, nom. nud. —Bifrenaria rudolfii (Hoehne) Carnevali & G.A. Romero, Monogr. Syst. Bot. Missouri Bot. Gard. 45: 1257. 1993. Bifrenaria minuta Garay, Bot. Mus. Leafl. 18: 206. 1958, non Maxillaria minuta Cogn. 1904. —Maxillaria perparva Garay & Dunst. in Dunst. & Garay, Venez. Orch. Ill. 6: 37. 1976. Epiphyte, shade-loving, usually growing low on the other plant, often among bryophytes; pseudobulbs 7–20 mm long, basally 3–6 mm wide; leaves 5–40 cm long; inflorescences branching when old; flowers lasting 2–4 days, purple or maroon with the lip a much deeper hue and a fringe of yellow on front lip margin; pedicellate ovary 5–7 mm long, subterete, somewhat dilated apically; floral bract 3–5 mm long; sepals 5-veined; dorsal sepal 8–10 mm long, 4.5–5 mm wide; labellum 10–11 mm long, 6–7 mm wide, ovate-elliptic to elliptic-trullate or oblongpandurate. Lowland to montane forests, 50– 800(–1200) m, the larger forms occurring at lower altitudes; Bolívar (Auyán-tepui, Gran Sabana, Río Carrao), Amazonas (common and widespread). Distribution as in genus. ◆Fig. 358. Hylaeorchis petiolaris is very variable vegetatively, especially regarding size, relative dimensions, and leaf thickness. Plants from higher elevations, especially from the

Gran Sabana area in Bolívar state in Venezuela and adjacent Guyana, tend to be smaller with thicker leaves that can measure to 15 cm long and a petiole of ca. 1/3 that length. Plants from the Amazonian lowlands are much larger with leaves of to 40 cm long while the petioles can be to 1/2 the total length of the leaf. These two forms look very different, but in some places at intermediate elevations in Amazonian Venezuela (e.g., the

Fig. 358. Hylaeorchis petiolaris

394

O RCHIDACEAE

slopes of Cerro Marahuaka) a mixture of forms linking both extremes can be found occurring sympatrically. An attempt to correlate the variation in size and leaf thickness with any other character has been unsuccessful and the differences seem to be only a response to different environmental condi-

tions in a phenotypically plastic species. Flowers in both extreme forms are essentially identical in size, color, and overall morphology. If recognition of two taxa, however, would eventually be warranted, the name Bifrenaria minuta would be available for the smaller form.

68. IONOPSIS H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 348, t. 83. 1815 [1816]. [Subtribe Oncidiinae]. Cybelion Spreng., Syst. Veg. 3: 679. 1826. by Germán Carnevali and Ivón M. Ramírez-Morillo Small twig epiphytes, rarely subterrestrial, cespitose, erect to pendulous, often attached to hosts by only a few roots. Pseudobulbs absent or very reduced and hidden by leaf sheaths, if present, 1-foliate. Leaves and leaf-bearing sheaths conduplicate, equitant, fleshy to subterete or almost terete, articulated with their sheaths. Inflorescences subterminal or axillary, 1–3, racemose or paniculate, shorter, ± equal to much longer than leaves; peduncle thin, subterete, remotely sheathed. Flowers minute to relatively large and showy, resupinate, long-pedicellate, white, pink, or violet, often with darker veins; floral bracts triangular to elliptic-triangular, acute, concave, very small; pedicellate ovary terete. Sepals free or the lateral fused into a small synsepal and forming a small mentum under the lip, subequal, elliptic to triangular-elliptic, acute, 3-veined; petals nearly equaling sepals but somewhat broader, acute to rounded, 4- or 5-veined. Lip larger to much larger than perianth segments, flat or somewhat concave basally and convex apically, obovate or transversely reniform, truncate to deeply emarginate, basally clawed, continuous with column base; disk basally 2-callose, 2-keeled. Column relatively short, erect, thick, exauriculate, footless; anther terminal, operculate, incumbent, with reduced partitions; clinandrium shallow; pollinia 2, cartilaginous, subglobose to obovoid; rostellum ventral; stigma ventral. Capsules ovoid to ellipsoid. U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 or 3 species, 2 in Venezuela, both in the flora area. Species of Ionopsis tend to grow as twig epiphytes on Psidium, Citrus, and Crescentia trees in relatively, open humid places such as river edges. Key to the Species of Ionopsis 1. 1.

Leaves terete or subterete; inflorescences few-flowered racemes; lip unlobed, ca. 6 mm long ........................................................... I. satyrioides Leaves laterally flattened; inflorescences many-flowered panicles; lip 2lobed, ca. 15 mm long ...................................................... I. utricularioides

Ionopsis satyrioides (Sw.) Rchb. f. in Walp., Ann. Bot. Syst. 6: 683. 1863. —Epidendrum satyrioides Sw., Prodr. 123. 1788.

Epiphyte, minute to small, usually shadeloving; leaves subterete to terete; racemes shorter than or equal to leaves; flowers small with spreading perianth segments, yellowish

Isochilus 395

to usually whitish or violet. Lowland to lower montane forests, 100–1000 m; Delta Amacuro (Serranía de Imataca), Bolívar (Altiplanicie de Nuria, upper Río Caroní basin). Aragua, Distrito Federal, Miranda; Panama, West Indies, Colombia, Guyana, Suriname, Ecuador, Brazil. ◆Fig. 359. Ionopsis utricularioides (Sw.) Lindl., Coll. Bot. t. 39A. 1821. —Epidendrum utricularioides Sw., Prodr. 122. 1788. Ionopsis paniculata Lindl., Edwards’s Bot. Reg. 22: sub t. 1904. 1836. Epiphyte, rarely subterrestrial or a lithophyte, usually sun-loving; inflorescences paniculate; flowers different shades of brown, very small to medium-large and showy. Evergreen forests, 100–500(–1000) m; Delta Amacuro (Río Amacuro, Serranía de Imataca), Bolívar (La Escalera to Cerro Venamo region, basins of upper Río Caroní, Río Caura, and Río Parguaza), Amazonas (basin of Río Cataniapo, Río Ventuari).

Fig. 359. Ionopsis satyrioides

Anzoátegui, Aragua, Carabobo, Falcón, Lara, Miranda, Sucre; U.S.A. (Florida), widespread in Neotropics.

69. ISOCHILUS R. Br. in W.T. Aiton, Hortus Kew. ed. 2, 5: 209. 1813. [Subtribe Laeliinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Usually epiphytes, rarely lithophytes or subterrestrial herbs, erect or ascending to drooping, cespitose to short-creeping, usually sun-loving. Roots thicker than the rhizome; rhizome creeping, frequently branching and forming cespitose plants. Stems terete, straight, distichously foliose in the upper 3/4, basally enveloped by tubular sheaths. Leaves conduplicate, articulate, linear to linear-elliptic, apically 2lobed, suberect, shiny green. Inflorescences terminal, racemose, rarely 1-flowered, dense or lax, secund or distichous, often scorpioid. Floral bracts chartaceous, inconspicuous to longer than the terete or 3-angled pedicellate ovary. Flowers nonresupinate, erect to spreading, tubular or the apices of the perianth segments spreading, membranous, short-lived, bright rose to purple, rarely whitish. Sepals free or the laterals connate in the basal 1/2, dorsally ± keeled or winged, lanceolate, oblong, or obovate, obtuse to acuminate; laterals ± gibbous at base; petals basally ± somewhat attenuate to clawed, subspatulate to elliptic, ± oblique, usually shorter and wider than the sepals, unkeeled. Lip shortly clawed, adnate to base of column, erect, similar to the petals but narrower, flat or concave, often constricted at middle, obtuse to acute, basally S-shaped. Column erect, straight, semiterete, wingless, footless or with a short foot; clinandrium 3-dentate; anther terminal, operculate, incumbent, 2-locular, each locule with a longitudinal, imperfect partition; pollinia 4, waxy, ovoid-oblong, laterally compressed, with flat, conspicuous caudicles; rostellum transverse, erect; stigma subapical. Capsules erect, oblong, ± 3-edged. Southern Mexico (where most diverse), Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina; 8–10 species, 1 in Venezuela.

396

O RCHIDACEAE

Isochilus linearis (Jacq.) R. Br. in W.T. Aiton, Hortus Kew. ed. 2, 5: 209. 1813. —Epidendrum lineare Jacq., Enum. Syst. Pl. 29. 1760. Epiphyte or lithophyte 7–70 cm tall; flowers pink to purple, rarely whitish. Lower montane forests, 900–1100 m; Bolívar (Río Pacairao, near Santa Elena de Uairén). Widespread elsewhere in Venezuela; other distribution as in the genus. ◆Fig. 360.

Fig. 360. Isochilus linearis

70. JACQUINIELLA Schltr., Repert. Spec. Nov. Regni Veg. 7: 23. 1920. [Subtribe Laeliinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, lithophytes, or terrestrial herbs, minute to small, erect to subpendulous, cespitose. Rhizomes almost absent. Stems terete or laterally compressed, enveloped by leaf sheaths, leafy to base but sometimes leafless in the basal 1/3, straight or zigzag. Leaves conduplicate, subterete or laterally compressed, articulated with their sheaths; sheaths sometimes rugulose or striate. Inflorescences terminal, solitary or fasciculate, when pedunculate, peduncle flattened and bearing 1 or 2 terminal bract-like leaves. Flowers minute to small, opening successively, widely or more often with subparallel perianth segments, often autogamous or cleistogamous, resupinate; floral bracts relatively large, acute or obliquely subtruncate, apiculate; pedicels short; ovary often subglobose or subellipsoid, with a cuniculus (nectary concealed within the ovary). Perianth segments fleshy, usually greenish, often red- or brown-tinged. Sepals subequal, free or shortly connate at base, concave; petals subequal to sepals, sometimes shorter and narrower. Lip shortly adnate

Jacquiniella 397

to column base or totally free, sessile or shortly clawed, very fleshy, sometimes geniculate and contracted below middle, unlobed or apically 3-lobed, concave, usually elliptic or subrhombic. Column short, thick, subterete, somewhat dilated apically, footless; anther terminal or subdorsal, operculate, incumbent, 2–4-locular; clinandrium mostly small, sometimes inflated or elongate, margins entire, erose, or denticulate; pollinia 4, waxy, laterally compressed, affixed to a small viscidium; rostellum horizontal, slit-like, breaking longitudinally when pollinia are detached; stigma ventral. Capsules ovoid-globose to ellipsoid. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; ca. 8 or 9 species, 3 in Venezuela, all in the flora area. Key to the Species of Jacquiniella 1. 1. 2(1). 2.

Leaves laterally compressed ................................................... J. steyermarkii Leaves terete or nearly so .......................................................................... 2 Inflorescences sessile ....................................................................... J. globosa Inflorescences long-pedunculate ................................................. J. teretifolia

Jacquiniella globosa (Jacq.) Schltr., Repert. Spec. Nov. Regni Veg. 7: 124. 1920. —Epidendrum globosum Jacq., Enum. Syst. Pl. 29. 1760. Epiphyte or lithophyte, small to minute, erect to subpendulous, sun-loving; leaves subterete; flowers minute, ± nodding to sharply pendulous, not widely opening, green or green-purple or brown-tinged. Rain or

cloud forests or open shrublands, usually in exposed positions, (100–)500–1600(–1700) m; Bolívar (widespread), Amazonas (widespread). Aragua, Carabobo, Distrito Federal, Falcón, Miranda, Nueva Esparta, Sucre, rarer in Andean states (Lara, Táchira); Mexico, Central America, West Indies, South America. ◆Fig. 361.

2 mm

Fig. 361. Jacquiniella globosa

Fig. 362. Jacquiniella steyermarkii

398

O RCHIDACEAE

Jacquiniella steyermarkii Carnevali & Dressler, Lindleyana 7: 18. 1992. Epiphyte, minute to small, erect to subpendulous, sun-loving; flowers solitary, nodding, almost closed, green or greenpurple or brown-tinged. Montane forests, often riparian, open associations, 700–1400 m; Bolívar (basin of Río Caroní, Río Parguaza), Amazonas (Cerro Marahuaka). Guyana. ◆Fig. 362.

Fig. 363. Jacquiniella teretifolia

Jacquiniella teretifolia (Sw.) Britton & P. Wilson, Bot. Porto Rico 6: 340. 1926. —Epidendrum teretifolium Sw., Prodr. 122. 1788. Epiphyte or terrestrial, erect or ascendent; leaves ± terete; inflorescence 1-flowered; flowers greenish yellow. Rain or cloud forests, 600–2500 m; Bolívar (Río Caroní basin), Amazonas (Cerro Autana, Cerro Marahuaka, Cerro Sipapo, Cerro Yapacana, Cerro Yaví). Aragua, Distrito Federal, Lara, Monagas, Sucre, Portuguesa, Táchira, Zulia; southern Mexico, Central America, West Indies, Colombia. ◆Fig. 363.

71. KEGELIELLA Mansf., Repert. Spec. Nov. Regni Veg. 36: 60. 1934. —Kegelia Rchb. f., Bot. Zeitung (Berlin) 10: 670. 1852, non Sch. Bip. 1848. [Subtribe Stanhopeinae]. by Gustavo A. Romero-González Cespitose epiphytes. Stem modified into ovoid, slightly compressed, shallowly sulcate and/or angulate pseudobulbs of 1 internode, when young entirely concealed by distichous, scarious sheaths. Leaves 1–3, most often 2, narrowly elliptic to narrowly ovate, acute, plicate, articulate, short- to long-petiolate. Inflorescences 1–several, lateral, arising from the base of the pseudobulbs, pendent, racemose, few- to many-flowered; peduncle densely pubescent. Flowers resupinate, showy; floral bracts concave, narrowly ovate, shorter than the pubescent, pedicellate ovary. Sepals and petals membranous, spreading; sepals similar, narrowly to broadly ovate, acute to acuminate, externally with glandular trichomes; petals similar to the sepals but much narrower. Lip fleshy, clawed, deeply 3-lobed, the lateral lobes large, erect or spreading, ± hatchet-shaped, broadly triangular or rounded, oblique; disk, between the lateral lobes, with an erect, fleshy callus, laterally compressed, dorsally 1- or 2-sulcate, 2-lobed, sometimes tongue-shaped. Column slender, elongate, recurved, semiterete, apically winged, footless; anther terminal, operculate, incumbent, 1-locular; pollinarium with 2 yellow pollinia, narrowly obovate, cleft, cartilaginous, and dorsoventrally flattened, the tegula long, oblong-linear, the viscidium narrowly obovate. Central America, Jamaica, Trinidad-Tobago, Colombia, Venezuela, Suriname, French Guiana; 5 species, 2 in Venezuela, 1 of these in the flora area.

Koellensteinia 399

2 mm Anther and pollinia

1 cm

5 mm Face of column

5 mm Column and lip

5 cm

Fig. 364. Kegeliella orientalis

Kegeliella orientalis G. Gerlach in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1137. 2000. Epiphyte; raceme pendent; flowers pale

brown with maroon striping. Tall forests, ca. 1100 m; Bolívar (Guayaraca). Endemic. ◆Fig. 364.

72. KOELLENSTEINIA Rchb. f., Bonplandia (Hanover) 2: 17. 1854. [Subtribe Zygopetalinae]. by Gustavo A. Romero-González Cespitose, terrestrial, or sometimes epiphytes. Stem abbreviate, modified into small pseudobulbs, concealed by distichous, scarious sheaths. Leaves 1–3, membranous to coriaceous, narrowly ovate, rarely ovate, acute to acuminate, attenuate toward the base or petiolate, plicate, articulate. Inflorescences lateral, arising from the base of the pseudobulbs, erect to slightly arching, racemose, few- to many-flowered, shorter than to longer than the leaves. Flowers resupinate; floral bracts concave, appressed, oblong or narrowly oblong-ovate, obtuse to acute, < 1/2 as long as the pedicellate ovary. Sepals and petals spreading, sometimes shallowly concave,

400

O RCHIDACEAE

membranous; sepals narrowly oblong-ovate to narrowly elliptic-ovate, acuminate, the lateral ones sometimes slightly oblique, adnate laterally to the apex of the column foot; petals oblong to oblong-elliptic, obtuse to subacute, sometimes slightly oblique. Lip clawed, articulate with the apex of the column foot, fleshy, concave, basally cordate to cuneate, conspicuously 3-lobed, lateral lobes erect to spreading, semicircular, reniform, or obovate, sometimes falcate, the midlobe widely ovate to transversely narrowly elliptic, apically obtuse, apiculate, retuse, emarginate or 2lobed; disk with a 2- or 3-dentate, retrorse callus. Column short, arched, semiterete, produced into a short foot, apically auriculate or not; anther terminal; pollinia 4, yellow, cartilaginous, dorsoventrally flattened, subequal, obovate to clavate, attached in 2 pairs to a subtriangular tegula. Belize, Panama, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 10 species, 6 in Venezuela, all in the flora area. Key to the Species Koellensteinia 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4).

5.

Plants epiphytic, mature leaves < 10 mm wide .......................... K. graminea Plants terrestrial, mature leaves > 15 mm wide ...................................... 2 Inflorescence < 1/2 as long as the subtending leaf; lip lateral lobes dentate ........................................................................................... K. hyacinthoides Inflorescence at least as long as the subtending leaf, usually longer; lip lateral lobes entire ................................................................................. 3 Lip with a 3-lobed callus .................................................................... K. lilijae Lip with a 2-lobed callus ............................................................................ 4 Lip lateral lobes strongly falcate, overlapping with central lobe; lip base cuneate, callus placed near the ithmus ................................... K. roraimae Lip lateral lobes oblique, not overlapping with central lobe; lip base cordate or subcordate, callus placed near the lip base ............................. 5 Lateral lobes of the clinadrium extending beyond the central lobe; lip with color spots near the base and, when extended, usually > 10 mm between the lateral lobes, the central lobe acute, apiculate .............. K. carraöensis Lateral lobes of the clinadrium not extending beyond the midlobe; lip with purple (brown in dry flowers) streaks throughout and, when extended, usually < 10 mm between the lateral lobes, the central lobe rounded, sometimes emarginate at the apex ............................................. K. tricolor

Koellensteinia carraöensis Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 166. 1976. Terrestrial; inflorescences long, erect, > 1/2 as long to longer than the leaves; flowers white with maroon spots on the lip base; lip with a basal, 2-lobed, retrorse callus. Wet forests, ca. 500 m; Bolívar (Río Carrao). Endemic. ◆Fig. 365. Koellensteinia graminea (Lindl.) Rchb. f., Bonplandia (Hanover) 4: 323. 1856.

—Maxillaria graminea Lindl., Edwards’s Bot. Reg. 21: sub t. 1802. 1836. —Promenaea graminea (Lindl.) Lindl., Edwards’s Bot. Reg. 29: misc. 13. 1843. Small epiphyte; leaves long, linear, narrow (6–10 mm wide); flowers small (1–1.5 cm diameter), yellowish pink, with maroon transverse stripes, the lip with a basal, 2lobed, retrorse callus. Locally abundant and frequent in wet forests, 50–900 m; Delta Amacuro (San Víctor), Bolívar (Río Toro, Río Uaiparú, Urimán), Amazonas (Cerro Duida,

Koellensteinia 401

Cerro Sipapo, Río Cataniapo, Río Sipapo). Colombia, Guyana, Ecuador, Peru, Brazil (Amazonas), Bolivia. Koellensteinia hyacinthoides Schltr., Beih. Bot. Centralbl. 42(2): 126. 1925. Terrestrial; inflorescence erect, generally < 1/2 as long as the leaves; flowers light yellow throughout; lip lateral lobes finely dentate. Open sandy savannas, 100–200 m; Amazonas (Caño Ucata southeast of Síquita, Cerro Yapacana, near San Carlos de Río Negro, Yavita-Maroa road). Colombia, Brazil (Amazonas).

common species found in the Guayana region, such as K. eburnea (Barb. Rodr.) Schltr., a name based on a Barbosa Rodrigues’s specimen from Minas Gerais, Brazil, and K. kellneriana Rchb. f., a name based on a specimen collected by H. Wagener in Venezuela’s state of Trujillo. Whether Koellensteinia tricolor and K. kellneriana are conspecific is yet to be determined.

Koellensteinia lilijae Foldats, Bol. Soc. Venez. Ci. Nat. 22: 263. 1961. Terrestrial; inflorescences erect, generally longer than the leaves; flowers whitish yellow, the lip with a basal, 3-lobed, retrorse callus, lateral lobes with maroon stripes. Low vegetation on lajas, ca. 100 m; Amazonas (Río Atabapo basin). Endemic. Koellensteinia roraimae Schltr., Orchis 12: 29. 1918. Terrestrial; inflorescence erect, generally longer than the leaves; flowers whitish yellow. Habitat and elevation not indicated; Bolívar (Roraima-tepui). Endemic. Koellensteinia roraimae is known only from the type collection, which was destroyed in Berlin in 1943. Koellensteinia tricolor (Lindl.) Rchb.f. in Walp., Ann. Bot. Syst. 6: 552. 1863. —Zygopetalum tricolor Lindl., Edwards’s Bot. Reg. 32: sub t. 64. 1848. Cyrtopodium eburnea Barb. Rodr., Revist. Eng. 3: 74. 1881. —Koellensteinia eburnea (Barb. Rodr.) Schltr., Orchis 12: 28. 1918. Terrestrial; inflorescence longer than the leaves; flowers white to greenish white with some purple transverse stripes. Open wet forests, savannas, 500–1200 m; Bolívar (Auyán-tepui, Ilú-tepui, Río Carrao, Río Carún, around Santa Elena de Uairén), Amazonas (Caño Ucata southeast of Síquita). Colombia, Peru, Brazil (Mato Grosso, Minas Gerais). Koellensteinia tricolor was most likely one of the many live plants Robert H. Schomburgk sent to the Loddiges establishment from what is now Guyana. It pre-dates all the other names cited in the past for the

Fig. 365. Koellensteinia carraöensis

402

O RCHIDACEAE

73. LEOCHILUS Knowles & Westc., Fl. Cab. 2: 143. 1838. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Twig epiphytes, rarely lithophytes, small to medium-sized, cespitose. Pseudobulbs clustered on short, creeping rhizomes, ovoid to suborbicular, laterally compressed (ancipitous), 1- or 2-foliate apically, lower portions concealed by 2 or 3 leaf-bearing sheaths, uppermost larger, eventually deciduous. Inflorescences produced laterally from base of pseudobulb, very rarely apically, subtended by a leafbearing sheath, racemose, paniculate, or secondarily paniculate, erect to pendent, 1–many-flowered; floral bracts inconspicuous. Flowers small to medium-sized, inconspicuous, resupinate; pedicellate ovary often twisted, glabrous. Lateral sepals free or united to 2/3 of their length, adnate to lip base, sepals ± equal, narrowly ovate, dorsally keeled, shortly acuminate, margins entire; petals free, similar to sepals but usually broader, forward projecting, often concealing column. Lip broadly attached to base of column forming a nectar-secreting cavity, longer than other perianth segments, entire or slightly lobed, sometimes ± pandurate, apically slightly emarginate, obtuse to truncate, with a fleshy-smooth or rough-tuberculate (horned) basal callus. Column short, with 2 stigmatic arms parallel to the cavity side walls and perpendicular to the column, located midway to the anther, footless; anther terminal, operculate, incumbent, 1-locular; clinandrium truncate; pollinarium with 2 waxy, subspheroidal pollinia, a relatively long, narrow tegula, and an oval viscidium; rostellum transverse; stigma ventral. Capsules trigonous to spheroid, weakly keeled to winged. U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad-Tobago, Ecuador, Brazil; 9 species, 2 in Venezuela, 1 of these in the flora area. Leochilus labiatus (Sw.) Kuntze, Revis. Gen. Pl. 2: 656. 1891. —Epidendrum labiatum Sw., Prodr. 124. 1788. Twig epiphyte, small, sun-loving, wholly red or purple-tinged; inflorescences 1–20 (–40)-flowered; flowers small, sweet-scented, yellow to yellow-green with red to red-brown blotches near callus. Lower montane forests, often on cultivated trees, 700–900 m; Bolívar (Santa Elena de Uairén). Aragua, Distrito Federal, Miranda; Mexico, Central America, West Indies, Colombia, Trinidad-Tobago, Ecuador, Brazil. ◆Fig. 366.

Fig. 366. Leochilus labiatus

Lepanthes 403

74. LEPANTHES Sw., Nova Acta Regiae Soc. Sci. Upsal. ser. 2, 6: 85. 1799. [Subtribe Pleurothallidinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, lithophytes, or subterrestrial herbs, mostly cespitose, but often creeping, erect, or pendulous. Rhizomes short or elongate, ramicauls 1-foliate, very short to elongate, sometimes proliferous, densely or laxly covered by a series of tubular, ribbed, ± imbricate sheaths with oblique, dilated, marginate ostia (wide apical opening of the funnel-like ramicaul bract), the ostia margins macro- or microscopically ciliate or scabrous, occasionally glabrous. Leaves conduplicate, membranous or rarely ± fleshy, linear-elliptic to orbicular, apex usually 3-denticulate; blades mostly flat, sometimes concave or convex, often red-tinged. Racemes 1–several, originating from an annulus near the ramicaul apex, few- to many-flowered; rachis straight to zigzag, sometimes with a peculiar fish-bone pattern. Flowers several, opening singly or more commonly successively, usually small, not resupinate, often lasting only one day, commonly brightly colored, red, yellow, or pink; floral bracts small to conspicuous; pedicellate ovary terete or 3-angled. Sepals glabrous or ciliate, often dorsally keeled; lateral ones mostly fused into a synsepal, rarely almost free; dorsal sepal entirely free to variously connate with basal portion of synsepal; petals generally wider than long (transverse), very rarely the petals longer than wide and then linear or very narrowly triangular, often differentiated into distinct upper and lower lobes. Lip usually 3-lobed, very rarely entire; central lobe modified into a variously shaped and sculptured appendix; lateral lobes often peltate or semilunate and embracing the column; base of lip connate to the under surface of column, often near its base. Column footless, usually subcylindrical, very short to relatively elongated; anther operculate, apical, subapical, or ventral; pollinia 2, attached to a small viscidium; stigma apical, subapical, or ventral. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 600–700 species, ca. 60 in Venezuela, 10 of these in the flora area. There is material in several relevant herbaria of what appears to be additional unidentified entities of the genus from the flora area. The lack of well preserved or more complete material of these specimens rendered them impossible to identify with any degree of certainty. Key to the Species of Lepanthes 1. 1. 2(1). 2. 3(2).

3.

Peduncle of inflorescence longer than subtending leaf ............................ 2 Peduncle of inflorescence shorter than subtending leaf .......................... 4 Ramicauls proliferous; column ca. 2.5 mm long ................ L. marahuacensis Ramicauls not proliferous; column < 2 mm long ...................................... 3 Lateral sepals fused for 1/2 of their length; blade of lip not divided into 2 obliquely ovate, acute lobes; petals ca. 2 times broader than long ........................................................................................................ L. dussii Lateral sepals almost free, fused for 1/3 of their length; blade of lip divided into 2 obliquely ovate, acute lobes; petals 3 times as broad as long .............................................................................................. L. unitrinervis

404

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

7(5). 7.

8(7).

8.

9(8). 9.

O RCHIDACEAE

Dorsal sepal 1-veined, plants minute, ramicauls < 2.5 cm long .................................................................................................. L. duidensis Dorsal sepal 3-veined; plants usually larger, ramicauls ≥ 3.5 cm tall ..... 5 Margins on outer sides of petals rounded, flat, or apiculate, never concave or emarginate; upper lobe of petals < 1/2 the length of dorsal sepal .... 6 Margins on outer sides of petals concave or emarginate; upper lobe of petals > 1/2 the length of dorsal sepal ......................................................... 7 Leaves narrowly ovate or narrowly elliptic, acute; internodes of rachis much longer than pedicel length; above 1600 m ........................... L. exilis Leaves widely ovate to suborbicular, apex broad and abruptly acuminate; internodes of rachis shorter than pedicel length; below 400 m ............................................................................................ L. helicocephala Leaves very broadly ovate, obtuse to very shortly acuminate; upper lobe of petals acute; apices of both petal lobes ciliolate ..................... L. cercion Leaves narrowly elliptic, ovate, or broadly ovate, always long-acuminate; upper lobe of petals rounded, obtuse, or truncate, never acute; apices of petal lobes totally glabrous or very rarely glabrescent ....................... 8 Sinus between dorsal sepal and lateral sepals shallow, inconspicuous; dorsal sepal wide-triangular, widest at the very base; mature leaves with conspicuous reticulate veins .............................................. L. aithalos Sinus between dorsal sepal and lateral sepal deep, always conspicuous; dorsal sepal ovate-elliptic, widest just below midlength; mature leaves without conspicuous reticulate veins .................................................... 9 Sepals shortly acuminate; upper lobe of petals truncate to obliquely truncate; lateral sepals connate to 1/2 of their length ................... L. pariaensis Sepals obtuse to acute; upper lobe of petals rounded to obtuse; lateral sepals connate to ca. 2/3 of their length ....................................... L. pectinata

Lepanthes aithalos Carnevali & I. Ramírez in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1137. 2000. Epiphyte 5–6 cm tall, cespitose; inflorescences not exceeding the length of the leaf; flowers widely opening; dorsal sepal ca. 8 × 5.5 mm, sepals orange with a waxy texture; petals orange; lip red. Cloud forests, 1100– 1900 m; Bolívar (Jaua-Sarisariñama massif). Sucre. Lepanthes cercion Luer & Escobar, Orquideología 17: 158. 1987. Epiphyte 2–9 cm tall, cespitose, erect; sepals pale yellow-brown flushed with maroon; petals maroon and yellow with a yellow streak along outer margin; lip red-maroon grading to orange near apex of column; column maroon-pink; anther pinkish white with fine pink spotting. Cloud forests, ca. 1300 m; Bolívar (La Escalera to Cerro Venamo region). Aragua, Falcón, Mérida, Sucre; Colombia.

Lepanthes duidensis Ames & C. Schweinf., Bull. Torrey Bot. Club 58: 346. 1931. Epiphyte 2–3.5 cm tall, cespitose, erect; leaves frequently purple-tinged; flowers minute, rose-purple. Cloud forests, often growing on thin branches of shrubs on tepui summits, 1300–1900 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Marahuaka). Nueva Esparta. ◆Fig. 367. Lepanthes dussii Urb., Repert. Spec. Nov. Regni Veg. 15: 106. 1917. Epiphyte 3–5 cm tall in flower, cespitose, erect; rachis slightly zigzag; flowers with pale brown sepals largely suffused with red except at margins, the midvein dark red; petals yellow; lip orange; column violet-purple. Cloud forests, 1200–1900 m; Bolívar (La Escalera to Cerro Venamo region), Amazonas (Sierra de la Neblina). Guadeloupe, St. Lucia. We have seen unidentifiable specimens from the Macizo del Chimantá that are probably referrable to this species.

Lepanthes 405

Lepanthes exilis C. Schweinf., Fieldiana, Bot. 28: 178. 1951. Epiphyte 7–21 cm tall, cespitose, erect; rachis ± zigzag, filiform; flowers very variable in size, color, and form of petals; sepals yellow, yellow flushed with brownish orange, or orange flushed with maroon; petals orange or with red or green margins, sometimes almost entirely red, not ciliate; lip brown-orange to rose or red; column pink, anther white or rose-purple. Cloud or rain forests, 1600–2500 m; Bolívar (Auyán-tepui, Jaua-Sarisariñama massif, Macizo del Chimantá, Ptari-tepui). Mérida, Táchira, Trujillo. Lepanthes helicocephala Rchb. f., Xenia Orchid. 1: 150. 1856. Epiphyte 6–15 cm tall, cespitose; flowers showy; dorsal sepal 4.3–5 mm long, maroonorange; lateral sepals orange-yellow flushed

with pinkish along margin of their outer sides and a thin band of red on margin of their inner sides; lip red with prominent, fleshy, yellow apical tails; the midlobe is a small, pink, fleshy process with fine pink hairs; column red at base grading to orange at apex; anther pink. Lowland evergreen forests, 100–200 m; Bolívar (Río Toro), Amazonas (Río Autana, Río Casiquiare, Río Cataniapo, near San Carlos de Río Negro). Guyana, Suriname, Ecuador, Amazonian Brazil. ◆Fig. 368. Lepanthes marahuacensis Carnevali & I. Ramírez, Novon 3: 113, fig. 8. 1993. Epiphyte, lithophyte, or subterrestrial, erect, prostrate, or creeping; flowers dull red. Local in tepui shrublands, often growing low on shrubs, ca. 2700 m; Amazonas (Cerro Marahuaka). Endemic. ◆Fig. 369. Fig. 367. Lepanthes duidensis

Fig. 368. Lepanthes helicocephala

Fig. 369. Lepanthes marahuacensis

Fig. 370. Lepanthes unitrinervis

406

O RCHIDACEAE

Lepanthes pariaënsis Foldats, Acta Bot. Venez. 3: 339. 1968. Epiphyte 6–12 cm tall, cespitose; inflorescences 1–4 simultaneously; flowers small; dorsal sepal 2.5–2.8 mm long, sepals tan, marked with maroon or yellow with a reddish orange center; petals and lip red. Cloud forests, 700–1900 m; Bolívar (Auyán-tepui, Jaua-Sarisariñama massif, La Escalera to Cerro Venamo region, Sierra Pakaraima), Amazonas (Cerro Marahuaka, Sierra de la Neblina). Sucre; Amazonian Brazil. Lepanthes pectinata Luer, Phytologia 59: 448. 1986.

Epiphyte 7–15 cm tall, cespitose; leaves ± suffused with purple; raceme very congested; flowers small with sepals 2.5–4 mm long, red-brown; petals orange, red toward the base; lip red-orange. Cloud forests, 900–1700 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, La Escalera to Cerro Venamo region). Sucre. Lepanthes unitrinervis Carnevali & I. Ramírez, Novon 3: 115, fig. 7D–H. 1993. Lithophyte or erect epiphyte, cespitose; raceme laxly few-flowered; flowers reddish purple. Dwarf cloud forests on tepui summits, 2500–2700 m; Bolívar (Ilú-tepui, Roraima-tepui). Endemic. ◆Fig. 370.

75. LEPANTHOPSIS (Cogn.) Ames, Bot. Mus. Leafl. 1: 3. 1933. —Pleurothallis sect. Lepanthopsis Cogn. in Mart., Fl. Bras. 3(4): 381. 1896. [Subtribe Pleurothallidinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, rarely lithophytes or subterrestrial, minute to small, cespitose, erect, arching, or pendulous. Stems thin, terete, 1-leaved at apex, enclosed by a series of lepanthiform sheaths (tubular, ribbed, imbricating sheaths with oblique, dilated, margined ostia, the ribs and margins of the ostia are grossly or microscopically ciliate or scabrous); stems often apically proliferous and rooting at base and apex. Leaves conduplicate, articulate, subsessile or basally attenuated in a short pseudopetiole, elliptic, narrowly elliptic, ovate, or obovate, apically rounded, obtuse, or acuminate, flat or concave. Inflorescences 1–many, originating from near ramicaul apex, with an annulus, racemose, few- to many-flowered, shorter than to much longer than subtending leaf; peduncle short or elongate; rachis straight or zigzag. Flowers minute to small, widely opening, often opening all at once or in slow succession and then usually several to many flowers open simultaneously, resupinate, bronze-green, hyaline, yellowish to deep wine purple. Dorsal sepal frequently 3-veined; lateral sepals free to variously connate, usually 1-veined, similar to the dorsal sepal, often falcate or oblique, all sepals frequently acuminate; petals usually dissimilar to sepals, often shorter and narrower, 1-veined. Lip usually simple, rarely 3-lobed, often concave and somewhat concealing the column, generally much shorter than sepals; disk callose or ecallose, attached to column base. Column short and thick, apically dilated, footless; anther terminal, operculate, incumbent; pollinia 2, waxy, subpear-shaped, attached to a small viscidium; rostellum ± transverse; stigma apical, 2-lobed, continuous under a flap on the rostellum. Capsules ovoid, obovoid, or ellipsoid. U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 40 species, 7 or 8 in Venezuela, 4 of these in the flora area. Species of Lepanthopsis are usually plants from cloud forests at intermediate to high elevations, rarely from lowland rain forests. Lepanthopsis is closely related to Trichosalpinx Luer but the short column is usually broadly winged and has a 2lobed, apical stigmatic surface.

Lepanthopsis 407

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

All 3 sepals connate for > 4/5 of their length forming a tube or cup ............................................................................................ L. obliquipetala Sepals free or only the laterals connate .................................................... 2 Lateral sepals connate for 2/3 of their length, free apices acute or rounded, not acuminate ......................................................................... L. floripecten Lateral sepals connate for 1/3 of their length, free apices acuminate ...... 3 Dorsal sepal 3-veined; petals about 2.5 mm long, ca. 3 times longer than wide; flowers pink ..................................................................... L. pulchella Dorsal sepal 1-veined; petals 1–2 mm long, ca. 2 times longer than wide; flowers dark wine purple or bright purple ................................ L. vinacea

Lepanthopsis floripecten (Rchb. f.) Ames, Bot. Mus. Leafl. 33: 11. 1933. —Pleurothallis floripecten Rchb. f., Bonplandia (Hanover) 2: 25. 1854. Epiphyte; ramicauls 2–8 cm tall; inflorescences longer than subtending leaf, manyflowered; flowers opening simultaneously, 2sided on rachis, tinged lilac, dull lavender, dull reddish purple, dull magenta, buff, or bronze-brown, sometimes lip a darker hue. Cloud forests, often growing on thin branches,

Fig. 371. Lepanthopsis floripecten

900–1900 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Jaua-Sarisariñama massif, La Escalera to Cerro Venamo region, Macizo del Chimantá [Toronó-tepui], Serranía Marutaní), Amazonas (Cerro Aracamuni, Cerro Aratitiyope, Cerro Marahuaka, Cerro Parú). Distrito Federal, Sucre, Trujillo; Mexico, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil. ◆Fig. 371.

Fig. 372. Lepanthopsis obliquipetala

408

O RCHIDACEAE

Lepanthopsis obliquipetala (Ames & C. Schweinf.) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 39: 76. 1991. —Physosiphon obliquipetalus Ames & C. Schweinf., Sched. Orchid. 8: 12. 1925, non Pleurothallis obliquipetala Acuña & C. Schweinf. 1938. —Stelis obliquipetala (Ames & C. Schweinf.) L.O. Williams, Ceiba 5: 54. 1956. —Pleurothallis connata Luer, Selbyana 5: 388. 1981. —Trichosalpinx obliquipetala (Ames & C. Schweinf.) Luer, Phytologia 54: 396. 1983. Erect epiphyte; leaves 3.6–5.5 cm long; flowers inconspicuous with dorsal sepal 3.5– 4 mm long; sepal cup and lip cream-colored; petals dark wine red. Cloud forests on tepui summits, 1500–2000 m; Bolívar (Macizo del Chimantá), Amazonas (Cerro Aracamuni). Costa Rica, Colombia, Ecuador. ◆Fig. 372.

Lepanthopsis pulchella Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 3: 160. 1965. Epiphyte; stems 2–4 cm tall; inflorescence solitary, raceme much longer (almost 5 times) than subtending leaf; flowers pale pink; dorsal sepal 2–8 mm long. Cloud forests, 1200–1700 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region). Endemic. Lepanthopsis vinacea C. Schweinf., Bot. Mus. Leafl. 18: 109. 1958. Epiphyte; stems 2–6 cm tall; inflorescence many-flowered; flowers bright purple; dorsal sepal 2–3 mm long; lip darker purple; column creamy white. Cloud forests, ca. 2000 m; Bolívar (Auyán-tepui, Jaua-Sarisariñama massif, La Escalera to Cerro Venamo region, Ptari-tepui, Uei-tepui). Ecuador.

76. LEUCOHYLE Klotzsch, Ind. Sem. Hort. Bot. Berol. app. 1. 1854. [Subtribe Oncidiinae]. Trichopilia Lindl., Intr. Nat. Syst. Bot. ed. 2, 446. 1836, pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, cespitose, erect. Rhizome thin, very short. Pseudobulbs oblong to narrow-ellipsoid, somewhat compressed or not, apically 1-foliate, clothed by 1 or 2 tubular sheaths that are eventually deciduous. Leaves linear to linear-elliptic, acute, fleshy and in some cases almost subterete, articulate. Inflorescences lateral, basal, emerging from axils of pseudobulb sheaths, pendulous or rarely arching, a (2)3–5(–8)-flowered raceme; peduncle short, with 1 or 2 sheaths; rachis ± dense, straight or ± zigzag; floral bracts conspicuous, chartaceous, shorter than pedicellate ovary. Pedicellate ovary terete. Flowers medium-sized, resupinate, with widely spreading segments, membranous, opening simultaneously, lasting several days, white, cream, yellowish, or somewhat hyaline, sometimes with red, purple, or pink spots. Petals and sepals narrow-lanceolate, acute. Lip slightly fleshier than the other perianth segments, free or fused to basal 1/4 of column, unlobed to slightly lobed, ovate to suborbicular, concave, margins smooth to laciniate; disk with a platelike or V-shaped callus, or 2-carinate; blade smooth or longitudinally 2-keeled. Column conspicuously shorter than lip, straight or geniculate and sometimes winged apically, footless; clinandrium ciliolate or denticulate, its margin lobed or simple; anther operculate, incumbent, 1-locular; pollinia 2, tegula elongate, linear-obovate, viscidium small, elliptic; stigma ventral, longer than wide; rostellum transversely perpendicular to column. Southern Mexico, West Indies, Central America, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, Ecuador, Peru, Brazil; 3 or 4 species, 2 in Venezuela, 1 of these in the flora area.

Ligeophila

409

Leucohyle mutica (Lindl.) Schltr., Orchideen 469. 1914. —Macradenia mutica Lindl., Edwards’s Bot. Reg. 25: misc. 20. 1839. —Trichopilia mutica (Lindl.) Rchb. f. & Wullschl. ex Rchb. f., Ann. Bot. (London) 6: 679. 1863. Herb 10–20 cm tall; flowers white, with pink or reddish tinge, especially on lower 1/2 of lip. Lowland forests, usually low on trees or close to rivers, ca. 50–500 m; Delta Amacuro (Río Cuyubini), Bolívar (middle Río Caura). Trinidad-Tobago, Guyana, Suriname. ◆Fig. 373.

Fig. 373. Leucohyle mutica

77. LIGEOPHILA Garay, Bradea 2: 194. 1977. [Subtribe Goodyerinae]. Erythrodes Blume, Bijdr. 410, t. 72. 1825, pro parte. by Germán Carnevali, Ivón M. Ramírez-Morillo, and Carlos A. Vargas Terrestrial to humicolous herbs, small to medium-sized, shade-loving. Rhizome cauliform, basally creeping, apically erect and foliose in the upper 1/3–1/2. Leaves ± fleshy, convolute, not articulate, usually elliptic or ovate-elliptic, acute to acuminate, with 1 or 2 conspicuous, longitudinal veins at each side of midvein, basally attenuated into a channeled pseudopetiole; sheaths invaginating the stem. Raceme erect, short-pedunculate, densely few- to many-flowered, shorter to slightly longer than the upper leaves. Flowers small to medium-sized, resupinate; floral bracts concave, shorter to ± equal to the ovary, ovate to elliptic, acute to acuminate, glabrous to pubescent; ovary arched, shortly pedunculate, cylindric to spindleshaped, usually pubescent. Sepals free, ± similar, elliptic to ovate; dorsal sepal somewhat concave; lateral sepals slightly oblique, apically spreading; petals connivent with dorsal sepal, obovate to obtrullate, oblique. Lip with a pair of fleshy plates or ridges at base leading to the entrance to the spur, adnate to column base, divided into a basal, ventricose hypochile, which is basally elongated into a well-developed spur, and an apical, transversely elliptic or rhombic to anchor-shaped epichile, often with reflexed lateral lobes; hypochile and epichile often separated by a well-developed claw. Column short; rostellum articulate, triggered, initially pointed outward, erect after release of the pollinia, 2-lobed; anther initially inclined, suberect after release of pollinia, concave; pollinia 4, sectile; caudicles well

410

O RCHIDACEAE

developed with a rather large viscidium adnate to the depressed rostellum; stigma confluent, terminal, horizontal. Capsule spindle-shaped. Central America, Colombia, Venezuela, Guyana, Suriname. French Guiana, Colombia, Ecuador, Peru, Brazil (most diverse in the Amazon basin); ca. 10 species, 4 in Venezuela, 3 of these in the flora area. Key to the Species of Ligeophila 1. 1. 2(1). 2.

Petals apically 2-lobed; inflorescence longer to ± equal to the upper leaves, many-flowered .............................................................. L. juruensis Petals unlobed apically; inflorescence shorter than the upper leaves, fewflowered .................................................................................................. 2 Petals oblong-ligulate, slightly oblique; epichile subsessile on the hypochile ................................................................................ L. amazonica Petals obliquely obtrulliform; hypochile separated from the epichile by a conspicuous claw ............................................................... L. stigmatoptera

Ligeophila amazonica Garay, Bradea 2: 194, fig. 2A. 1977. Humicolous or terrestrial; flowers green and white. Forests on tepui slopes, 1000– 1200 m; Amazonas (Cerro Huachamacari). Endemic. Ligeophila juruensis (Hoehne) Garay, Bradea 2: 195. 1977. —Physurus juruensis Hoehne, Comiss. Linhas Telegr. Estratég. Mato Grosso Amazonas, anexo 5, Bot. 1: 30. 1910. Physurus santensis Kraenzl., Kongl. Svenska Vetenskapsakad. Handl. 46(10): 33. 1911. —Erythrodes santensis (Kraenzl.) C. Schweinf., Bot. Mus. Leafl. 9: 128. 1941. Terrestrial or humicolous, to 33 cm tall; stems sometimes branching near base, 3- or 4-foliate in the upper 1/3; flowers with spreading sepals; lip with an apically 2-lobed epichile, apical lobes reflexed. Montane forests, 700–1200 m; Bolívar (Kukenán-tepui, La Escalera to Cerro Venamo region, Uaipán-tepui), Amazonas (Río Cataniapo). Colombia, Suriname, Peru, Brazil. Ligeophila stigmatoptera (Rchb. f.) Garay, Bradea 2: 195. 1977. —Physurus stigmatopterus Rchb. f., Xenia Orchid. 2: 185. 1873. —Erythrodes stigmatoptera (Rchb. f.) Pabst, Orquídea (Niteroi) 18: 215. 1957. Humicolous or terrestrial, 10–40 cm tall,

Fig. 374. Ligeophila stigmatoptera

Liparis 411

basally decumbent and then erect; inflorescence short, prominently bracteate; sepals and petals buff-salmon; lip white and green. Lower montane forests, 500–1200 m; Bolívar

(near Cerro Venamo, Río Paragua basin), Amazonas (Cerro Yapacana, near Puerto Ayacucho). Colombia, Guyana, Suriname, Peru, Brazil. ◆Fig. 374.

78. LIPARIS Rich., De Orchid. Eur. 21, 30, 38. Aug.–Sept. 1817, nom. cons. [Tribe Malaxideae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial, lithophytic, or epiphytic herbs, small to medium-sized. Rhizomes short to elongate, creeping or scandent. Stems absent, inconspicuous to well developed, frequently pseudobulbous-thickened; pseudobulbs spherical to narrowly spindle-shaped. Leaves terminal, conduplicate to plicate, bases sheathing, sometimes pseudopetiolate, usually deciduous, distichous to subopposite. Inflorescence terminal, usually long-pedunculate, racemose, few- to many-flowered, peduncle and rachis winged. Flowers minute to relatively showy, resupinate, purplish, reddish, yellowish, or greenish; perianth segments widely spreading, sometimes reflexed, membranous to fleshy. Sepals free, ± equal or the lateral ones somewhat oblique, frequently with revolute margins; petals narrower than sepals, usually filiform. Lip free or shortly adnate to column base, sessile to shortly clawed, usually unlobed, rarely 3-lobed, laciniate or dentate, flat or convex, much wider than other perianth segments, usually with 2 small basal callosities. Column elongate, usually somewhat arched, slender to relatively thick, usually with 2 small apical wings, semiterete, footless; anther terminal, operculate, incumbent; pollinia 4, in 2 pairs, waxy, usually ovoid; stigma entire, ventral. Capsule ellipsoid to ovoid. Cosmopolitan (most diverse in Tropical Asia and Oceania), in the Americas found in U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina; ca. 250 species, 5 in Venezuela, 3 of these in the flora area. Liparis is closely related to Malaxis Sol. ex Sw., from which it differs chiefly by its long column and incumbent anther. Key to the Species of Liparis 1.

1.

2(1).

2.

Plants small, 4–5 cm tall, with a long taproot; pseudobulbs apically aphyllous but totally enveloped by 2 or 3 sheathing leaves; inflorescences ca. 5 cm long, with verticils of 2–4 flowers; lip elliptic-lanceolate, acute, the margins erose .......................................... L. verticillata Plants larger, (6–)9–20(–35) cm tall; roots fasciculate; pseudobulbs with an apical leaf and variously naked or totally sheathed by leaf sheaths; inflorescences longer (< 7 cm long), racemose; lip either obovate or ovate-elliptic, apically truncate to obtuse, the margins entire ............ 2 Leaves 1 per pseudobulb, subconduplicate or of subfleshy-herbaceous texture with only the midvein apparent; lip wider near base, ecallose or almost at base; plants sun-loving ........................................ L. jamaicensis Leaves 2–4 per pseudobulb, with submembranous texture and conspicuously plicate (several parallel veins conspicuous); lip wider near apex, with 2 small, conical calli at base; plants usually shade-loving ..................................................................................................... L. nervosa

412

O RCHIDACEAE

Fig. 375. Liparis jamaicensis

Liparis jamaicensis Lindl. ex Griseb., Cat. Pl. Cub. 261. 1866. Sturmia elliptica Rchb. f., Linnaea 22: 833. 1849. —Liparis elliptica (Rchb. f.) Rchb. f. Ann. Bot. Syst. 6: 218. 1861, non Wight 1851. Liparis vexillifera auct. non (Llave & Lex.) Cogn.: sensu Dunst. & Garay, Venez. Orchid. Ill. 3: 163. 1965. Foldats in Lasser, Fl. Venez. 15(1): 455. 1970. Terrestrial, rarely lithophytic herb; pseudobulbs mostly subspheric; peduncle and rachis 2-winged; flowers minute to small, greenish or yellowish, sometimes purple-tinged with perianth segments spreading; dorsal sepal 6–13 mm long. Savannas or savanna-like tepui formations, sometimes in places subject to annual burning, 1000–1300 m; Bolívar (Cerro Kukenán, Gran Sabana). Aragua, Distrito Federal, Mérida, Miranda; Costa Rica, Panama, West Indies, Colombia. ◆Fig. 375.

Fig. 376. Liparis nervosa subsp. nervosa

Liparis nervosa (Thunb.) Lindl., Gen. Sp. Orchid. Pl. 26. 1830. —Ophrys nervosa Thunb., Fl. Jap. 27. 1784. Sturmia kappleri Rchb. f., Linnaea 22: 833. 1849. —Leptorchis kappleri (Rchb. f.) Kuntze, Revis Gen. Pl. 671. 1891.

Lockhartia 413

—Liparis kappleri (Rchb. f.) Rchb. f., Ann. Bot. Syst. 6: 218. 1864. —Liparis nervosa f. kappleri (Rchb. f.) Christenson & Carnevali, Lindleyana 11: 17. 1996. Terrestrial; pseudobulbs conical to spindle-shaped; leaves deciduous; flowers purplish, reddish, brownish, or greenish; dorsal sepal 5–8 mm long. Semideciduous forests to rain forests, generally in places with a well-defined dry season, 50–300 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela and New World tropics and subtropics; 2 subspecies, both in the flora area. This species is variable and may encompass several taxa. Key to the Subspecies of L. nervosa 1. Leaves usually > 8 cm long in mature plants, often 10–30 cm long, usually > 4 cm wide, often to 10 cm wide, usually elliptic, of a firmer texture; plants usually growing on soil, often in boggy places or as subepiphytes in extremely humid places .......................... subsp. nervosa 1. Leaves (3–)5(–8) cm long, (1.6–)2.9(–3.9) cm wide, usually narrowly elliptic, membranaceous, pale yellow-green upon drying; inflorescences 5–12 cm long; plants growing on granitic outcrops ................ ............................................... subsp. A L. nervosa subsp. nervosa. —Ophrys nervosa Thunb., Fl. Jap. 27. 1784. Liparis elata Lindl., Bot. Reg. 14: t. 1175. 1828. Terrestrial or subepiphytic herb 10–30 cm tall. Growing on soil, often in boggy places, or as subepiphytes in extremely humid places,

near sea level to 300 m; Delta (Caño Simoina), Bolívar (basin of Río Caura), Amazonas (basin of Río Casiquiare). Rest of distribution as in species. ◆Fig. 376. L. nervosa subsp. A Lithophytic herb 8–15 cm tall. Often on vertical rock walls or cliffs, 50–200 m; Bolívar (basins of Río Parguaza and Río Orinoco in the area of the lajas), Amazonas (Caño Coromoto, basins of Río Casiquiare and Río Cataniapo). Endemic (but likely to occur in neighboring Colombia). This entity is known from a few collections and grows in seasonally dry areas in environments that are much drier than those where the typical subspecies grows. Although very similar to the autonym, the plants of this subspecies are smaller, the leaves are narrower and dry a distinctive pale yellow-green or pale yellow-brown color. The inflorescences are also proportionately shorter. The flowers are somewhat smaller but otherwise identical to those of subspecies nervosa. More material could indicate that this entity should be recognized at the species level. Liparis verticillata G.A. Romero & Garay, Harvard Pap. Bot. 4: 484, fig. 7. 1999. Terrestrial herb 5–7 cm tall; corms emerging from a thick taproot to 2 cm long; leaves 9–11 mm long and wide; inflorescences to 5 cm long, angled in section, bearing a few irregular verticils of 3 or 4 flowers each toward the apex of the peduncle; sepals ca. 6 mm long, 1 mm wide, translucent white, the midvein red; lip with a prominent red midvein. Growing among mosses or bryophytes, ca. 2500 m; Bolívar (Cerro Kukenán). Endemic.

79. LOCKHARTIA Hook., Bot. Mag. 54: t. 2715. 1827. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, small to medium-sized, erect to pendulous, cespitose. Stems elongate, subterete to flattened, wholly covered by short, imbricating, laterally compressed sheaths. Leaves laterally compressed, narrow-oblong to narrow- or broadtriangular, acute to rounded, sometimes dorsally keeled, basally imbricate and articulate with their sheaths. Inflorescences originating from the leaf axils, usually subapically, frequently from the middle or lower internodes of the stem, 1–many simultaneously, 1–many-flowered, when many-flowered short, successively to simultaneously few-flowered racemes, rarely short, simultaneously flowered lax panicles; bracts of rachis and peduncle usually ± foliaceous, flat to concave, relatively con-

414

O RCHIDACEAE

spicuous. Flowers resupinate or not, inconspicuous to relatively showy, usually yellow with purple or maroon tinges or spots, or totally white; floral bracts usually ± foliaceous, flat to concave, relatively conspicuous; ovary terete. Perianth segments membranous or subfleshy, widely spreading; petals and sepals similar, sometimes the petals somewhat wider, usually elliptic or oblong, rarely ovate or obovate, acute, rounded, or truncate. Lip usually flat to convex, unlobed to variously lobed, when lobed usually with a pair of erect, antrorse to retrorse, short, slender basal lobes, frequently ± pandurate; apex obtuse to deeply emarginate; disk ecallose or variously callose, the upper surface of lip glabrous to variously pubescent. Column relatively short, subparallel with the lip forming an obtuse to acute angle with it, laterally produced in a pair of well-developed wings; anther apical to ventral, operculate, incumbent; pollinarium consisting of a small viscidium, a short or elongate hyaline tegula and 2 obpear-shaped pollinia; rostellum transverse; stigma ventral or subapical. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 35 species, 7 in Venezuela, 3 of these in the flora area. Key to the Species of Lockhartia 1. 1.

2(1).

2.

Inflorescence a short, lax panicle; flowers white; lip deeply emarginate ......................................................................................................... L. acuta Inflorescence a 1-flowered or successively few-flowered raceme; flowers yellow; lip not deeply emarginate, generally obtuse, rounded, truncate, or shallowly emarginate ........................................................................ 2 Lip not enveloping the column in fresh condition, narrow-ovoid or elliptic in general outline when open, with 2 well-developed calli, conspicuously 3-lobed in the basal 1/3; column wings erose or dentate .................................................................................................. L. imbricata Lip enveloping the column in fresh condition, suborbicular or transverseelliptic when open, essentially ecallose, unlobed; column wings entire ................................................................................................... L. latilabris

Lockhartia acuta (Lindl.) Rchb. f., Bot. Zeitung (Berlin) 10: 767. 1852. —Fernandezia acuta Lindl., Edwards’s Bot. Reg. 21: t. 1806. 1836. Lockhartia pallida Rchb. f., Bonplandia (Hanover) 2: 14. 1854. Lockhartia lasseri Schnee, Revista Fac. Agron. (Maracay) 1: 120. 1952. Epiphyte 15–50 cm long, erect or pendulous; flowers minute. Humid forests, in shady places, near sea level to 50 m; Delta Amacuro (locality not given). Aragua, Barinas, Lara, Miranda, Zulia; Panama, Colombia, Trinidad-Tobago. Lockhartia acuta was cited from Delta Amacuro by E. Foldats (Flora de Venezuela 15(5): 99. 1970) with the notation “Woolley”

Fig. 377. Lockhartia imbricata

Lueddemannia

415

in parenthesis. It is unknown if “Woolley” collected the plant, cultivated the plant, published a paper, or told Foldats the information. Lockhartia imbricata (Lam.) Hoehne, Arq. Bot. Estado São Paulo 2: 139. 1952. —Epidendrum imbricatum Lam., Encycl. 1: 189. 1783. Lockhartia weigelti Rchb. & Rchb. f., Bot. Zeitung (Berlin) 10: 767. 1852. Epiphyte 8–30 cm tall, erect to pendulous; flowers yellow, tinged or spotted with purple or maroon. Shady places in humid forests, 50–500 m; Delta Amacuro (Cuyubini, basin of Río Acure), Bolívar (Río Parguaza), Amazonas (basin of Río Cataniapo, Río Ventuari). Anzoátegui, Apure, Barinas, Monagas, Yaracuy; Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil. ◆Fig. 377. Lockhartia latilabris C. Schweinf., Fieldiana, Bot. 28: 200. 1951. Epiphyte 14–40 cm tall, pendulous; flowers yellow with a few maroon or orange spots in the lip. Cloud forests on tepui summits or slopes, 600–1700 m; Bolívar (Amaruaytepui, La Escalera to Cerro Venamo region, Sororopán-tepui). Guyana. ◆Fig. 378.

Fig. 378. Lockhartia latilabris

80. LUEDDEMANNIA Linden & Rchb. f., Bonplandia (Hanover) 2: 281. 1854. [Subtribe Stanhopeinae]. by Gustavo A. Romero-González Cespitose, robust epiphytes. Stem modified into ovoid, shallowly sulcate pseudobulbs of 1 internode, when young entirely concealed by distichous, scarious sheaths. Leaves 2, narrowly to broadly ovate, acute, plicate, articulate, subpetiolate. Inflorescences 1–several, lateral, arising from the base of the pseudobulbs, pendent, racemose, many-flowered. Flowers waxy, resupinate, showy; floral bracts concave, obovate, shorter than the velutinous pedicellate ovary. Sepals similar, fleshy, concave, elliptic to broadly ovate; petals smaller than the sepals, narrowly obovate to ovate-spatulate. Lip fleshy, clawed, concave, apically 3-lobed, the claw semiterete, with a high, dark, erect, conspicuous, pilose tooth; lateral lobes erect, subtriangular; midlobe triangular; disk with a prominent axial callus. Column slender, elongate, recurved, semiterete, apically winged, footless; anther terminal, operculate, incumbent, 1-locular; pollinarium with 2 yellow pollinia, obovate, cleft, cartilaginous, and dorsoventrally flattened, the tegula short, linear, the viscidium subtriangular. Colombia, Venezuela, Ecuador, Peru; 2 species, 1 in Venezuela.

416

O RCHIDACEAE

Lueddemannia pescatorei (Lindl.) Linden & Rchb. f., Bonplandia (Hanover) 2: 281. 1854. —Cycnoches pescatorei Lindl. in Lindl. & Paxton, Paxt. Fl. Gard. 1: 123. 1853 [1850]. Inflorescence to 20-flowered or more; flowers reddish yellow, fleshy, the lip claw with a dark maroon, pilose tooth. Montane forests, ca. 1500 m; Bolívar (Cerro Jaua). Central Coastal Cordillera, Táchira; Colombia, Ecuador, Peru. ◆Fig. 379.

Fig. 379. Lueddemannia pescatorei

81. LYCASTE Lindl., Edwards’s Bot. Reg. 29: misc. 14. 1843. [Subtribe Lycastinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, lithophytes, or subterrestrial herbs, medium- to rather large-sized, erect, cespitose. Rhizome very short. Pseudobulbs heteroblastic (1 internode), (1)2or 3(4)-foliate apically; broadly ellipsoid, ovoid, or suborbicular, slightly compressed, somewhat angled and ribbed, especially when dry, enveloped by sheaths of which the innermost 1–3 may have foliar blades. Leaves large, plicate, elliptic, attenuate basally into a variously developed petiole, usually deciduous and lasting only one growing season, sometimes longer-lasting, when deciduous leaving an abscission and armed scar on the apex on pseudobulb. Inflorescences lateral, originating from pseudobulb base, 1-flowered, 1–many flowering simultaneously in the same plant, shorter to much longer than pseudobulb, but usually shorter than leaves, in some species originating from mature pseudobulbs when plants are defoliated or foliated, in other species appearing with a new shoot; peduncle terete, erect, covered by several bracts. Flowers showy, resupinate, fleshy, nodding to spreading, usually long-lasting; floral bracts usually large, fleshy, hooded, and partially or totally concealing the terete pedicellate ovary. Sepals free, subequal or the lateral ones decurrent on column foot and sometimes oblique, green, yellow, white, pink, or purple, widely spreading to subparallel to column and lip, usually elliptic, apex acute to rounded; petals usually smaller than sepals, color same as or very different than sepals, generally parallel to column and forming a hood over it, apex sometimes reflexed or spreading. Lip erect, hinged to column foot, nearly unlobed to

Lycaste 417

sharply 3-lobed, basal lobes enfolding the column, sometimes fimbriate, usually of the same color as petals but with darker zones or stripes; disk ecallose. Column erect, elongate, subterete, straight to ± arched, sometimes ventrally pubescent, basally produced into a conspicuous foot; anther apical, operculate, incumbent; pollinia 4, with relatively small, elongate tegula and viscidium; stigma ventral. Capsules oblong to spindle-shaped. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia (most diverse in Central America and the Andes); ca. 30 species, 4 in Venezuela, 2 of these in the flora area. Key to the Species of Lycaste 1.

1.

Central lobe of lip with fimbriate margins, especially in the basal 1/2; perianth segments acute; floral bract ± equal to the pedicellate ovary ................................................................................................. L. fulvescens Central lobe of lip with glabrous margins; perianth segments obtuse to rounded; floral bract longer than the pedicellate ovary ...... L. macrophylla

Lycaste fulvescens Hook., Bot. Mag. 71: t. 4193. 1845. Epiphyte, lithophyte, or subterrestrial; flowers somewhat nodding; sepals and petals 6–8 cm long, cream-colored, flushed with light tan in the apical 1/2; lip pale orange to dark cream, callus and central lobe darker. Cloud forests, ca. 1900 m; Bolívar (JauaSarisariñama massif). Táchira; Colombia, Ecuador. Lycaste macrophylla (Poepp. & Endl.) Lindl., Edwards’s Bot. Reg. 29: misc. 14. 1843. —Maxillaria macrophylla Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 37, t. 64. 1835 [1836]. Subterrestrial, lithophyte, or epiphyte; flowers spreading; sepals 4–7 cm long, light green flushed with coppery brown; petals creamy white flushed with pink toward apex; lip creamy white with some yellow on throat

Fig. 380. Lycaste macrophylla

and reddish purple on veins and callus. Rain forests, 500–600 m; Bolívar (Auyán-tepui). Aragua, Distrito Federal, Miranda; Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 380.

418

O RCHIDACEAE

82. LYROGLOSSA Schltr., Beih. Bot. Centralbl. 37(2): 448. 1920. [Subtribe Spiranthinae]. Spiranthes Rich., De Orchid. Eur. 20, 28, 36. Aug.–Sept. 1817, nom. cons., pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Small terrestrial herbs, usually leafless during the flowering season (hysteranthous). Roots fleshy, tuberous, spindle-shaped, fasciculate, pubescent. Leaves convolute, rosulate, basal, few, small or sheath-like. Inflorescence erect, slender, remotely several-bracteate, ending in a rather loose, several-flowered, ± spirally twisted spike. Flowers small, resupinate, not widely opening; floral bracts rigid, concave, erect or suberect, acuminate, pubescent; ovary subsessile, arched, spindle-shaped to cylindric, ± twisted. Sepals similar, divergent from a subparallel base, pubescent outside; dorsal sepal ovate-elliptic, deeply concave; lateral sepals spreading with ± reflexed apex, basally decurrent on a short column foot without forming a true mentum; petals variable, the internal margin firmly adherent with dorsal sepal and with oblique, nondecurrent base. Lip membranous, pandurate, subsessile with a very short claw, the margins linear, callose thickened; the margins of the blades above base adherent with column on both sides. Column short, dilated above, sulcate and pubescent in front, basally produced into a short foot of same length; anther ovate-hooded, acute; pollinia 4, soft, clavate with a small, narrowly oblong viscidium; rostellum rigid, acuminate; stigmas 2, terminal, approximate, touching each other at the middle, ± cleft by a median furrow in front of column. Central America, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil; 3 species, 1 in Venezuela. Lyroglossa grisebachii (Cogn.) Schltr., Anexos Mem. Inst. Butantan, Secc. Bot. 1(2): 27. 1921. —Spiranthes grisebachii Cogn. in Mart., Fl. Bras. 3(4): 207. 1895. Terrestrial, sun-loving; leaves few; inflorescence erect, few-flowered in the apical 1/3; flowers small, white; lateral sepals apically reflexed-spreading; lip white with purple veins. Open savannas, savanna-like associations, elevation not known; Bolívar (locality not given). Miranda; Trinidad-Tobago, Brazil. ◆Fig. 381. Bolívar was cited without locality by Rudolph Schlechter (Repert Spec. Nov. Regni Veg. Beih. 6. 1919.

Fig. 381. Lyroglossa grisebachii

Macradenia

419

83. MACRADENIA R. Br., Bot. Reg. 8: t. 612. 1822. [Subtribe Oncidiinae]. Rhynchadenia A. Rich. in Sagra, Hist. Phys. Cuba, Bot. Pl. Vasc. 2: 248, fig. 85. 1853. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, small to medium-small, erect, cespitose, shade-loving. Pseudobulbs subcylindric to narrowly ovoid, 1-foliate, enveloped by 1–3 scarious, not leaf-bearing sheaths. Leaves conduplicate early and later flattened or not, erect to horizontal, narrowly elliptic, narrowly oblong-elliptic, or elliptic, acute to subacuminate, pseudopetiolate, subfleshy to coriaceous. Inflorescences basal, racemose, rarely pseudopaniculate (a new raceme is borne at the base of the old rachis after the flowers of the first rachis have wilted), pendulous, laxly to ± densely few- to many-flowered; peduncle relatively short, straight, few-sheathed, the rachis longer. Flowers small to medium-sized, resupinate, floral bracts small, concave; pedicellate ovary thin, subcylindric. Perianth segments usually widely spreading, often apically reflexed, ± fleshy, green, greenish white, bronzy maroon, or red, usually with paler, often yellow or bronze edges; sepals free, subequal or the dorsal broader, usually narrow-elliptic, acute or acuminate; lateral sepals sometimes oblique; petals similar to lateral sepals, but often broader. Lip sessile, adnate to column base, 2- or 3-lobed, blade flat or ± concave, usually white, yellowish, or maroon, with purple or maroon longitudinal stripes, central lobe longer and narrower than lateral ones, frequently linear, narrow-oblong to narrow-elliptic, acute to acuminate, flat or decurved. Column terete, ventrally grooved, exauriculate, footless; anther subdorsal, erect, elongate; clinandrium large, subdorsal to terminal, marginally dentate to laciniate; pollinia 2, cartilaginous, with a long, membranous, narrow-obovate tegula; viscidium small; rostellum prominent; stigma ventral. Capsul ellipsoid, attenuate on both extremes. U.S.A. (Florida), southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 5–7 species, 3 in Venezuela, all in the flora area. Key to the Species of Macradenia 1.

1.

2(1).

2.

Dorsal sepal 16–23 mm long; central lobe of lip 7–14 mm long, longer than basal blade, lip base cuneate; flowers dark maroon-red or redbrown, with a yellow border ................................................ M. brassavolae Dorsal sepal 8–14 mm long; central lobe of lip 3–5.5 mm long, shorter than or subequal to basal blade; lip base rounded to truncate; flowers greenish, yellowish, purple, or bronzy .................................................. 2 Lateral sepals 3.3–5.5 mm wide; central lobe of lip shorter than the width of the flattened basal portion; blade with 3 low longitudinal keels; perianth segments faintly maroon toward apex .................... M. lutescens Lateral sepals 2.3 mm wide; central lobe of lip much longer than the width of the flattened basal portion; blade with a high longitudinal keel; perianth segments bronzy and brown ........................... M. rubescens

420

O RCHIDACEAE

Macradenia brassavolae Rchb. f., Bot. Zeitung (Berlin) 10: 734. 1852. Serrastylis modesta Rolfe, Bull. Misc. Inform. Kew 1894: 158. 1894. —Macradenia modesta (Rolfe) Rolfe, Orchid Rev. 4: 357. 1896. Epiphyte; perianth segments dark maroon-red or brown with a yellow border; lip white with maroon-red longitudinal stripes. Lowland forests, 100–200 m; Amazonas (Río Ventuari basin). Carabobo; Mexico, Central America, Colombia, Ecuador. ◆Fig. 382. Macradenia lutescens R. Br., Bot. Reg. 8: t. 612. 1822. Twig epiphyte, shade-loving; perianth segments greenish or yellowish, often sparsely purple-spotted; lip white with purple streaks. Lowland forests, 50–600 m; Bolívar (Río Paragua basin), Amazonas (Río Cataniapo basin). Aragua, Miranda, Táchira, Zulia; U.S.A. (Florida), West Indies, Colombia, Ecuador, Peru, Suriname. Macradenia rubescens Barb. Rodr., Gen. Spec. Orchid. 1: 139. 1877. Twig epiphyte; perianth segments bronzy suffused with pale green and brown; lip lilacwhite with lavender streaks. Lowland forests, ca. 50 m; Delta Amacuro (Caño Araguao). Amazonian Brazil, Bolivia.

Fig. 382. Macradenia brassavolae

84. MACROCLINIUM Barb. Rodr., Gen. Spec. Orchid. 2: 236. 1882. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Twig epiphytes, fan-shaped, minute to small, cespitose, erect to pendulous, mostly sun-loving. Roots thin and long. Stems short, often pseudobulbous, very rarely monopodial. Pseudobulbs usually laterally compressed, ovoid, ellipsoid, or oblong, apically 1-foliate, totally or partially enveloped by leaf-bearing sheaths. Leaves and sheaths equitant, laterally compressed, articulate, or leaf sheaths absent; blade flat to triquetrous, fleshy to fleshy-coriaceous, usually obliquely triangular or elliptic to narrowly ovate, acute, often covered with reddish or brownish pustules or warts. Inflorescences lateral, originating from the base of the pseudobulbs or from the axils of the leaf sheaths, usually long-pedunculate and much longer than subtending leaf, erect to very rarely pendulous, raceme with a short rachis usually forming a capitate raceme. Flowers usually opening simultaneously, small but large for plant size, resupinate or not; floral bracts small, usually narrow; pedicellate ovary filiform. Perianth segments membranous to thinly fleshy, translucent, pink or purple, often with purple or pink spots, especially on petals, lip frequently white. Sepals free or the laterals somewhat connate basally; petals subequal or somewhat

Macroclinium 421

narrower than the sepals. Lip linear to (rarely) ovate, basally clawed, unlobed or lobed, frequently pandurate, often with a pair of retrorse lobes above the claw, apical lobe usually sagittate or cordate, apically truncate, rounded, acute, or more frequently long-acuminate. Column footless, elongate, linear to filiform, terete, apically expanded and frequently geniculate; anther dorsal, operculate, incumbent, often long-rostrate apically, basally subcordate; pollinia 2, hard, yellow, obovoid to subspheroid; tegula elongate, narrow, apically expanded, viscidium very small; clinandrium shallow; rostellum rostrate; stigma small, narrow. Capsules ellipsoid. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia; ca. 25 species, 3 in Venezuela, 2 of these in the flora area. Macroclinium is closely related to Notylia Lindl. and Macradenia R. Br., differing from them by its laterally compressed leaves. Flowers are pollinated by euglossine bees. There is a specimen of a third species of the genus represented by a sterile specimen from Gavilán, Amazonas state (Romero 1290, VEN), which is accompanied by a color photograph. This taxon is closely related to Macroclinium mirabile but the surface of the leaves is scabrous, and the lip is much longer than the other perianth segments. It probably represents the Peruvian species Macroclinium chasei Dodson & Bennett but good flowering material is required to confirm this determination. Key to the Species of Macroclinium 1. 1.

Pseudobulbs absent; leaves 5–8 mm long, surfaces smooth; column ca. 1/2 the length of the lip; lip apically obtuse .................................. M. mirabile Pseudobulbs present but very small; leaves 10–30 mm long, surfaces scabrous; column much longer than the lip; lip apically acute-acuminate ................................................................................ M. wullschlaegelianum

Macroclinium mirabile (C. Schweinf.) Dodson, Icon. Pl. Trop. 10: pl. 938. 1984. —Notylia mirabilis C. Schweinf., Bot. Mus. Leafl. 12: 202, pl. 22, figs. 8–14. 1946. Notylia norae Garay, Bot. Mus. Leafl. 18: 214, fig. 42. 1958. —Macroclinium norae (Garay) Dodson, Icon. Pl. Trop. 10, t. 938. 1984. Twig epiphyte 1.5–2.5 cm tall, including the inflorescence; dorsal sepal 2–3.5 mm long, pinkish, transluscent, petals with 2 or 3 darker pink spots. Lowland forests, often growing on shrubs close to waterfalls, 50– 600 m; Bolívar (La Escalera to Cerro Venamo region, Río Chicanán, Río Parguaza, Río Uaiparú), Amazonas (Río Cataniapo, Río Casiquiare). Peru, Amazonian Brazil. ◆Fig. 383.

Fig. 383. Macroclinium mirabile

422

O RCHIDACEAE

Macroclinium wullschlaegelianum (H. Focke) Dodson, Icon. Pl. Trop. 10, t. 939. 1984. —Notylia wullschlaegeliana H. Focke, Bot. Zeitung (Berlin) 11: 343. 1853. Twig epiphyte 3.5–5 cm tall, including the inflorescence; flowers large for the plant with dorsal sepal 4.8–7 mm long, translucent purple, petals with several dark purple spots. Lowland forests; 50–700 m; Bolívar (Altiplanicie de Nuria, La Escalera to Cerro Venamo region, Río Caura basin). Monagas, Sucre; Costa Rica, Suriname, Ecuador, Peru, Brazil. ◆Fig. 384.

Fig. 384. Macroclinium wullschlaegelianum

85. MALAXIS Sol. ex Sw., Prodr. 119. 1788. [Tribe Malaxideae]. Microstylis Nutt., Gen. N. Amer. Pl. 2: 196. 1818. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial or rarely epiphytes, mostly shade-loving, very variable as to habit and ecological preferences. Rhizomes well developed and creeping, or very short, prostrate or ascending to erect. Stems pseudobulbose-thickened or not, when thickened, narrowly pear-shaped to subcylindric, erect, covered by sheaths, with 1–3 apical to subapical leaves or distichously foliose. Leaves plicate, nonarticulate, membranous to subfleshy, contracted below into a long-sheathing petiole; the petioles enveloping the basal part of the peduncle. Inflorescences terminal, racemose; peduncle terete or ± compressed, usually much longer than the rachis, often the rachis very short and the flowers in an umbel-like raceme; floral bracts inconspicuous. Flowers small or minute, usually greenish or yellowish, usually nonresupinate. Perianth segments subfleshy, spreading; sepals free or the laterals connate below the basal 1/2, subequal; petals usually narrower than the sepals, often filiform, sometimes twisted. Lip sessile, nonarticulate, fleshier and broader than the rest of the perianth segments, unlobed or 2- or 3-lobed; base concave, rounded, cordate, auriculate, or sagittate; margin entire to lacerate. Column very short, erect, terete; clinandrium conspicuous, membranous, oblique; anther terminal, erect, 2-locular; pollinia 4, waxy, in 2 pairs, with small caudicles; rostellum erect; stigma ventral to subterminal. Capsules ovoid. Cosmopolitan (most diverse in tropical Asia and Oceania), in the Americas found in U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, northern Argentina; ca. 200 species, ca. 15 in Venezuela, 1 of these in the flora area. A collection from Roraima-tepui (Im Thrun 289, K), supposedly referable to Malaxis umbelliflora, has been reported in the literature (Foldats in Lasser, Fl. Venez. 15(1): 445. 1969). It has never been collected again, and the known distribution of M. umbelliflora makes it very unlikely to occur in the flora area. This report is probably based on a misidentification. Malaxis maguirei C. Schweinf., Bull. Torrey Bot. Club 75: 217. 1948. Malaxis excavata auct. non (Lindl.) O. Ktze., 1891: sensu Foldats in Lasser, Fl.

Venez. 15(1): 430. 1969, pro parte. Terrestrial 10–50 cm tall, with 2 subopposite broadly ovate leaves and an umbellike inflorescence; flowers greenish, yellow-

Masdevallia 423

ish, or white. Montane to upper montane forests, 1100–2300 m; Bolívar (La Escalera to Cerro Venamo region, Macizo del Chimantá [Apacará-tepui, Toronó-tepui], Roraimatepui), Amazonas (Sierra de la Neblina). Suriname. ◆Fig. 385. The Venezuelan material of Malaxis maguirei may represent an undescribed species.

Fig. 385. Malaxis maguirei

86. MASDEVALLIA Ruiz & Pav., Fl. Peruv. Prodr. 122, t. 27. 1794. [Subtribe Pleurothallidinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Subterrestrial, lithophytes, or epiphytes, shade- to sun-loving, minute to medium-large, erect to pendulous, mostly cespitose but rarely creeping. Ramicauls 1foliate, short to almost absent, very rarely somewhat elongate, mostly terete, rarely angled, enveloped by 1–several sheaths, sometimes pustulose or blackish. Leaves coriaceous to fleshy, conduplicate, flattened to subterete, usually narrowly obovateelliptic but often elliptic or linear-elliptic, sessile, long-petiolate; petiole channeled. Inflorescences emerging with an annulus below the abscission layer, short- to longpedunculate, 1-flowered or successively 1–few-flowered, rarely racemose with several flowers opening simultaneously; peduncle terete or triquetrous, sometimes basally triquetrous and apically terete, rarely somewhat flattened, remotely 1–4sheathed. Flowers small to large and showy, mostly resupinate, long- to short-pedicellate. Ovary smooth, verrucose or rugulose. Sepals variously connate, forming a basal cup or tube, rarely almost free, usually the lateral forming a conspicuous synsepal, apices variously developed into tails to almost without tails, color

424

O RCHIDACEAE

variable; petals fleshy to membranous, free, much smaller than sepals, usually oblong or elliptic, apically lobed, dentate or rarely entire, with a callus margin often developed into a tooth. Lip small, ± strap-shaped, fleshy, hinged to an incurved extension from the apex of the column foot, often constricted about or above the middle and then with an apical lobe and basal blade. Column terete, straight or recurved, winged or not; anther ventral or subterminal, operculate, incumbent, 1-locular; clinandrium entire or variously dentate; pollinia 2, ovoid, waxy, attached to a small viscidium; rostellum transverse; stigma ventral. Capsule smooth or with sharp points (erinaceous). Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 350 species, 40 in Venezuela, 6 of these in the flora area. Key to the Species of Masdevallia 1. 1. 2(1). 2. 3(2). 3. 4(1).

4.

5(4). 5.

Peduncle of the inflorescence distinctly triquetrous in the apical 1/3 ...... 2 Peduncle of the inflorescence apically terete (rarely somewhat triquetrous basally) ......................................................................................... 4 Lateral sepals free almost to base; ovary verruculose or rugulose .................................................................................................. M. picturata Lateral sepals basally united forming a distinct synsepal; ovary smooth ................................................................................................................ 3 Petioles blackish; tails distinctly longer than synsepal, very narrow, ca. 1.5 cm long .......................................................................... M. guayanensis Petioles green or greenish; tails shorter than or rarely ± equal to synsepal, triangular, 1 cm long ........................................................... M. norae Leaves distinctly petiolate, the petiole black; tails of the sepals ca. 2 times as long as synsepal or longer; dorsal sepal 38–40 mm long ................................................................................................ M. manarana Leaves basally gradually attenuated, never distinctly petiolate, the petiole green; tails of the sepals ± equal to the synsepal length; dorsal sepal 25 mm long ............................................................................................. 5 Synsepal tails abruptly attenuated; lip distinctly constricted about the middle; inflorescences successsively 1–3-flowered .................... M. sprucei Synsepal tails gradually attenuated; lip not distinctly constricted about the middle; inflorescences 1-flowered ............................ M. wendlandiana

Masdevallia guayanensis Lindl., London J. Bot. 2: 673. 1843. Epiphyte, small, cespitose; inflorescences 1-flowered; flowers with a yellow sepal cup. Montane forests; Bolívar (Roraima-tepui). Guyana, Brazil. Masdevallia manarana Carnevali & I. Ramírez, Ernstia 44: 15. 1987. Epiphyte, cespitose, erect; ramicauls blackish; inflorescences 1-flowered, ± equal to the leaves; flowers medium-sized for the

genus, white, with long sepal tails. Montane forests, 1800–1900 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 389. Masdevallia norae Luer, Lindleyana 3: 44. 1988. Masdevallia guttullata auct. non Rchb. f. 1877: sensu Dunst. & Garay, Venez. Orchid. Ill. 2: 195. 1961; Foldats in Lasser, Fl. Venez. 15(3):34. 1970. Epiphyte, small to medium-sized, cespitose; flowering successively (rarely 2 at a

Masdevallia 425

time); sepal cup dull olive green, diffusely dotted with purple, tails dull yellow-orange; petals and lip yellowish with some red-spotting. Lowland to lower montane forests, (100–)300–1000 m; Bolívar (Río Caroní basin), Amazonas (basins of Río Casiquiare and Río Negro). Endemic. ◆Fig. 387. Masdevallia picturata Rchb. f., Otia Bot. Hamburg. 1: 16. 1878. Epiphyte, small, cespitose; inflorescences ± equal to the leaves, 1(–3)-flowered; ovary verruculose; sepals almost free; sepal cup pale green or pale lavender, purple-spotted, tails green or reddish; petals white; lip white or yellow with lavender areas. Montane forests, 1300–2000 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region, Macizo del Chimantá), Amazonas (Cerro Sipapo, Sierra Fig. 386. Masdevallia wendlandiana

Fig. 387. Masdevallia norae

de la Neblina). Aragua, Distrito Federal; Costa Rica, Colombia, Guyana, Bolivia. Masdevallia sprucei Rchb. f., Otia Bot. Hamburg. 1: 17. 1878. Epiphyte, small, cespitose, shade-loving; inflorescences basally triquetrous, apically terete, ± equal to or surpassing the leaves, successively 1–3-flowered; sepal cup yellow, internally with 2 large patches of dark maroon and red-purple tails; petals and lip white or cream. Lowland to lower montane forests, 100–900 m; Bolívar (Perai-tepui), Amazonas (Mawishiña on upper Río Cunucunuma, Río Casiquiare basin, San Carlos de Río Negro). Brazil. ◆Fig. 388. Masdevallia wendlandiana Rchb. f., Gard. Chron. ser. 3, 1: 174. 1887. Epiphyte, small, cespitose, shade-loving; inflorescences ± equal to leaves, 1-flowered; sepal cup white with purple splashes or streaks within; tails yellow or white. Lowland to lower montane forests, 100–l000 m; Bolívar (basins of Río Caroní and Río Caura), Amazonas (basins of Río Casiquiare and Río Sipapo). Colombia, Ecuador, Peru, Brazil. ◆Fig. 386.

Fig. 388. Masdevallia sprucei

Fig. 389. Masdevallia manarana

426

O RCHIDACEAE

87. MAXILLARIA Ruiz & Pav., Fl. Peruv. Prodr. 116, pl. 25. 1794. [Subtribe Maxillarinae]. Ornithidium Salisb., Trans. Hort. Soc. London 1: 293. 1812. Camaridium Lindl., Bot. Reg. 10: t. 848. 1824. Heterotaxis Lindl., Bot. Reg. 12: t. 1028. 1826. Dicrypta Lindl., Gen. Sp. Orchid. Pl. 44. 1830. Marsupiaria Hoehne, Arq. Bot. Estado São Paulo 2: 68. 1947. Pseudomaxillaria Hoehne, Arq. Bot. Estado São Paulo n.s. formato maior 2: 71. 1947. by Germán Carnevali and Ivón M. Ramírez-Morillo Generally epiphytes, but often lithophytes or terrestrial herbs, inconspicuous to very large, cespitose to creeping or climbing, erect to pendulous. Rhizome short to very long, appressed to substrate, ascendent, arching or pendulous, usually enclosed in sheaths, the sheaths sometimes leaf bearing. Plants with pseudobulbs and then sympodial or without pseudobulbs and then often monopodial. Pseudobulbs various, usually spheric, ovoid, or oblong, rarely ± 4-edged, enveloped by few to several sheaths of which the innermost may have foliar blades. Leaves conduplicate, articulate with their sheaths or with the apical internodes of the pseudobulb, 1–3(4) at the apex of each pseudobulb, sessile or attenuated into a channeled or terete pseudopetiole, usually oblong or oblong-elliptic, but often linear, subterete, broadly elliptic, or obovoid, apex symmetrical to irregularly oblique (one of the halves of the blade longer than the other). Inflorescences 1-flowered, 1–many, erect, ascendent or pendulous, originating from the base of pseudobulbs or from the axils of the rhizome or stem sheaths, usually clothed by scarious sheaths, shorter to longer than pseudobulbs. Flowers inconspicuous to large and showy, usually resupinate, fugacious to rather long-lived, variously colored, but often yellow, brown, white, or purple; floral bracts inconspicuous to large and completely covering the pedicellate ovary; pedicellate ovary cylindrical to 3-edged. Perianth segments free, submembranous to rather fleshy, subparallel to column and the flowers not widely spreading. Sepals usually longer and broader than petals, the lateral usually oblique and decurrent along column foot; petals frequently subparallel to column. Lip fleshier than the other perianth segments, auriculate or not to the apex of the column foot, simple to 3-lobed, central lobe usually larger than the lateral ones, disk usually with one or more longitudinal, often waxy calli, other calli frequently found on the central lobe. Column usually long, cylindrical to hemicylindrical, straight or arched; anther apical, operculate, incumbent; pollinia 4, in 2 superposed, unequal pairs, viscidium usually well developed, often widely lunate; stipes often lunate or apparently missing, sometimes elongate; rostellum transverse, stigma ventral, transverse, very narrow. Capsules mostly ellipsoid or subspheric, usually with smooth surface, sometimes 3-edged. U.S.A. (Florida), Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, northern Argentina; ca. 400 species, ca. 100 in Venezuela, 59 of these plus 1 hybrid in the flora area. The circumscription of Maxillaria is problematic and most recent authors include Dicrypta, Heterotaxis, Marsupiaria, Pseudomaxillaria, Camaridium, and Ornithidium within its boundaries, but some of these groups might merit generic rank. A thorough phylogenetic analysis of the genus (and of subtribe Maxillariinae)

Maxillaria 427

is currently underway, and until such a study is completed, we will refrain from recognizing at the generic level any of the distinctive groups here included within Maxillaria. Similarly, delimitation of species within the genus is also often difficult within some of the complexes that comprise the genus. Since more comprehensive collection efforts in the Neotropics, particularly in the Andean countries and southern Central America, are still revealing many undescribed taxa, this makes it difficult to estimate the total number of species in Maxillaria at this time. Many species of Maxillaria seem to be pollinated by wax-gathering bees or wasps; euglossine-bee pollination has been reported for M. rufescens and its allies (although we have never seen any of these bees attracted to a member of the M. rufescens complex), some other species or populations seem to be always autogamous. Many species of the Ornithidium-complex with brightly colored, cup-like flowers fit the hummingbird-pollination syndrome. Maxillaria ponerantha Rchb. f., known from the Venezuelan coastal range, Suriname, and French Guiana, is likely to be found in the flora area. Key to the Species of Maxillaria 1. 1. 2(1). 2.

3(2).

3.

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

6.

7(4).

Leaves terete, subterete, or laterally compressed .................................... 2 Leaves conduplicate ................................................................................... 4 Leaves laterally compressed; plants monopodial, without pseudobulbs; lip dark maroon .............................................................................. M. equitans Leaves terete or subterete; plants sympodial, with pseudobulbs (even though very small and/or totally enclosed by sheaths); lip white or creamy yellow ........................................................................................ 3 Leaves and pseudobulbs densely crowded on the rhizome, the distances between the pseudobulbs shorter to a little longer than pseudobulb length; lip base shaply attenuated into a narrow claw, the apex rounded ........................................................................................ M. uncata Leaves and pseudobulbs sharply distant on the rhizomes, the distances between the pseudobulbs much longer than pseudobulb length; lip base gradually attenuated into a broad claw; the lip apex 2-lobed ................ ................................................................................................. M. squamata Plants without pseudobulbs ....................................................................... 5 Plants with well-developed pseudobulbs, although these sometimes hidden by sheaths ....................................................................................... 7 Flowers yellowish or white, produced singly in the axils of the leaves; perianth segments > 2 cm; lip articulate with column foot ....... M. meridensis Flowers clear to bright yellow, produced in fascicles; perianth segments < 1.2 cm; lip rigidly attached to the column foot .................................. 6 Leaves obtuse; ovary conspicuously 3-edged; petals obovate, conspicuously attenuated basally; lip very fleshy, unspreadable, conspicuously 3-lobed ......................................................................................... M. alticola Leaves acute to acuminate; ovary terete or inconspicuously 3-edged; petals elliptic, inconspicuously attenuated basally; lip submembranous, easy to spread, inconspicuously 3-lobed ........................................ M. aurea Lip rigidly attached to the column foot; lateral sepals connate in the basal

428

O RCHIDACEAE

1 3

7.

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

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

11.

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

13.

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

16(8).

16. 17(16).

/ ; flowers produced in short, dense fascicles with the peduncles and pedicellate ovaries completely hidden by densely packed, imbricate sheaths ....................................................................................... M. conferta Lip articulate with column foot; lateral sepals free; flowers produced singly or in fascicles but rarely if ever with the peduncles and pedicellate ovaries completely hidden by sheaths .................................................. 8 Pseudobulbs apically 2- or 3-leaved .......................................................... 9 Pseudobulbs apically 1-leaved ................................................................. 16 Rhizome abbreviate, bearing crowded pseudobulbs, usually attached to substrate, forming cespitose plants .................................................... 10 Rhizome elongate, bearing pseudobulbs at intervals, creeping or attached to substrate only at base, forming erect, ascendent, or pendulous plants .............................................................................................................. 11 Sheaths enveloping the pseudobulbs not leaf bearing; lip orange or yellow-brown, about 2 times longer than wide ..................... M. grobyoides Sheaths enveloping the pseudobulbs leaf bearing; lip dark purple or maroon, 3 or more times longer than wide .............................. M. proboscidea Flowers originating from the base of the pseudobulbs; lip conspicuously clawed with a triangular blade and a conspicuous V-shaped, waxy callus .................................................................................. M. notylioglossa Flowers originating from the leaf axils or young growths; lip not clawed, blade of the lip not triangular and if with a waxy callus, this not V-shaped ............................................................................................... 12 Lip not lobed, may be somewhat constricted or dilated around the middle but without forming distinct lobes ...................................................... 13 Lip lobed ................................................................................................... 15 Inflorescences concealed by 2 or more, acute, somewhat inflated bracts; pedicellate ovary totally hidden by floral bracts; all perianth segments narrowly elliptic, stiffly coriaceous; petals 5 times longer than wide .............................................................................................. M. scorpioidea Inflorescences naked or only partially concealed by bracts; pedicellate ovary naked or nearly so; perianth segments oblong-elliptic to ovateelliptic; fleshy; petals 2–3 times longer than wide ............................. 14 Leaves < 10 mm wide; lip basically white, with or without red spots .............................................................................................. M. stenophylla Leaves > 10 mm wide (very rarely only to 7 mm wide); lip deep maroon or deep yellow with a maroon callus .................................... M. guareimensis Rhizome strong or weak but not rigid, totally enveloped by sheaths; apical lobe of the lip shorter than basal lobe; sepals obtuse ............ M. camaridii Rhizome rigid, not totally enveloped by sheaths, leaving portions naked; apical lobe of the lip longer than basal lobe, deeply emarginate; sepals acute .............................................................................................. M. rigida Rhizome elongate, bearing pseudobulbs ± approximate to remote, creeping or attached to substrate only at base, forming erect, ascendent, or pendulous plants .................................................................................. 17 Rhizome abbreviate, with aggregate pseudobulbs or these somewhat creeping but always closely approximate ........................................... 29 Sheaths enveloping pseudobulbs without foliar blades or when rarely

Maxillaria

429

present, these soon deciduous ............................................................. 18 17. Sheaths enveloping pseudobulbs with conspicuous leaf blades ............. 19 18(17). Inflorescences originating from the axils of sheaths from new growths; flowers white; lip subsimple; pedicellate ovary and capsules subcylindric; plants from lowlands ............................................................... M. alba 18. Inflorescences originating from the base of young pseudobulbs base; flowers yellow; lip deeply 3-lobed; pedicellate ovary and capsule 3-edged; plants growing above 1200 m ................................................... M. alpestris 19(17). Perianth segments < 0.8 cm long ............................................................ 20 19. Perianth segments > 1.2 cm long ............................................................ 21 20(19). Pseudobulbs 1–2 cm long; petals linear-elliptic to linear-obovate; lip subsimple, constricted about the middle ............................ M. mapiriensis 20. Pseudobulbs 1.4–8 cm long; petals oblong or ovate; lip deeply 3-lobed ..................................................................................................... M. ramosa 21(19). Leaves > 15 times as long as wide ........................................................... 22 21. Leaves < 10 times as long as wide ........................................................... 25 22(21). Rhizome relatively short; central lobe of the lip acute; dorsal sepal linearelliptic ............................................................................................ M. tenuis 22. Rhizome very elongate bearing pseudobulbs at long internodes; central lobe of lip obtuse or rounded; dorsal sepal elliptic to oblong-elliptic .............................................................................................................. 23 23(22). Central lobe of lip elliptic; pseudobulbs 4–5 cm long ............... M. schlechteri 23. Central lobe of lip ovate or triangular-ovate; pseudobulbs to 3.5 cm long .............................................................................................................. 24 24(23). Peduncle > 10 cm long; dorsal sepal > 40 mm .................... M. ×dunstervillei 24. Peduncle 1.5–3 cm long; dorsal 17–22 mm long ...................... M. meridensis 25(21). Dorsal sepal > 30 mm long; plants from high tepuis .............................. 26 25. Dorsal sepal < 25 mm long (very rarely to 27 mm long); plants from elevations usually below 1500 m ................................................................. 27 26(25). Petals and sepals concolorous or with the apical half pale maroon; leaves linear-oblong or linear-elliptic, acute; midlobe of lip smooth or slightly scabrous ............................................................................ M. ×dunstervillei 26. Petals and sepals with the apical half dark maroon; leaves oblong-lanceolate to oblong-oblanceolate, obtuse to broadly acute; midlobe of lip verruculose ................................................................................. M. quelchii 27(25). Central lobe (or apical) of lip deeply emarginate; petals obtuse; plants from Amazonas state .................................................................... M. rigida 27. Central lobe (or apical) of lip obtuse, acute, or subrounded, not deeply emarginate; petals acute; plants from Bolívar state .......................... 28 28(27). Flowers membranous; lip 3-lobed at the basal 1/2; petals acute or obtuse ................................................................................................. M. imbricata 28. Flowers subfleshy; lip only constricted below middle with a pair of lateral auricles (rarely these missing); petals acuminate ..................... M. lasallei 29(16). Sheaths enveloping pseudobulbs without leaf blades ............................ 30 29. Sheaths enveloping pseudobulb conspicuously leaf bearing .................. 42 30(29). Dorsal sepal > 4 cm .................................................................................. 31 30. Dorsal sepal < 3.7 cm long (usually < 3.4 cm long) ................................. 32 31(30). Pseudobulbs pear-shaped to ellipsoid, only slightly flattened; leaves long-

430

O RCHIDACEAE

31.

32(30). 32. 33(32).

33.

34(33).

34. 35(33). 35. 36(35). 36. 37(36). 37.

38(32). 38. 39(38). 39. 40(39).

40.

41(39). 41.

petiolate; apical lobe < 1/2 the length of the lip; margins of clinandrium not fimbriate ............................................................................ M. luteoalba Pseudobulbs suborbicular to broadly ellipsoid, strongly flattened; leaves short-petiolate; apical lobe 1/2 or more the length of lip; margins of clinandrium fimbriate ............................................................... M. setigera Inflorescences shorter than to subequaling the pseudobulb length, when longer < 2 times the pseudobulb length .............................................. 33 Inflorescences > 2.5–3 times longer than pseudobulbs (generally much longer) .................................................................................................. 38 Flowers yellow, orange, yellowish orange, or greenish; lip 3-lobed below its middle, yellow or orange with purple, maroon, or brown blotches; leaves sessile ........................................................................................ 34 Flowers yellow-brown, deep purple, or maroon; lip 3-lobed around or above its middle, maroon or deep purple; leaves very rarely sessile .............................................................................................................. 35 Central lobe of lip tapering toward the apex; sepals 9–15 mm long; leaves mostly 5–10 cm long, narrowly elliptic to elliptic; pseudobulbs usually ellipsoid to ± pear-shaped ....................................................... M. acutifolia Central lobe of lip oblong, often broader toward apex; sepals 20–25 long; leaves usually > 15 cm long, oblong; pseudobulbs oblong ...... M. rufescens Sepals widely spreading with petals subparallel to column; central lobe of lip about as wide as long .................................................... M. desvauxiana Sepals and petals subparallel to column; central lobe of lip conspicuously longer than wide .................................................................................. 36 Floral bracts narrowly elliptic; lip 3 times longer than wide .... M. foldatsiana Floral bracts widely elliptic; lip 2 times longer than wide ..................... 37 Plants terrestrial to lithophytic in open places; sheaths of the pseudobulbs not disintegrating; petiole thick, variable in length ...... M. auyantepuiensis Plants epiphytic in cloud or rainforests; sheaths enveloping pseudobulbs defibrating and persisting; petioles thin, always longer than pseudobulb .......................................................................................... M. longipetiolata Perianth white; dorsal sepal < 0.45 cm wide .................................. M. kegelii Perianth yellow, orange, brown, purple, or combinations of these colors; dorsal sepal at least 0.6 cm wide (usually wider) ............................... 39 Dorsal sepal < 2.3 cm long ....................................................................... 40 Dorsal sepal > 2.6 cm (usually to 3 cm or more long) ............................. 41 Leaves conspicuously petiolate; blades 3–5 times longer than wide; flowers brown-purple or pink; sepals acute; lip only shallowly lobed ............................................................................................ M. cryptobulbon Leaves sessile or inconspicuously petiolate; blades 6–10 times longer than wide; flowers yellow, creamy yellow, or orange, usually purple-, brown-, or red-tinged toward apex; sepals obtuse to rounded; lip clearly 3-lobed ................................................................................................... M. porrecta Leaves (in the flora area) 20–25 mm wide; central lobe of lip broader toward apex; dorsal sepal 6–9 mm wide .................................. M. loretoensis Leaves at least 28 mm wide (usually > 30 mm); central lobe of lip broader toward base or at central portion; dorsal sepal 11–18 mm wide ..................................................................................................... M. parkeri

Maxillaria 431

42(29). Dorsal sepal 2.9–4.5 times longer than lip .............................................. 43 42. Dorsal sepal < 2.4 times longer than lip ................................................. 48 43(42). Lip simple, slightly pandurate, 3-lobed, or basal lobes separated from central lobe by smooth, rounded indentations; lip white with a yellow callus ......................................................................................... M. albiflora 43. Lip sharply 3-lobed, lateral lobes separated by conspicuous indentations from central lobe; lip purple, yellow, or white with a deep orange central lobe and apices of lateral lobes ..................................................... 44 44(43). Column foot (measured from point of insertion of dorsal sepal to point of insertion of lip) about 3/4 of column length; central lobe of lip < 1/5 of total lip length .....................................................................M. marmoliana 44. Column foot > 1/4 column length; central lobe of lip > 1/3 of total lip length .............................................................................................................. 45 45(44). Sepals and petals deep reddish purple or maroon at apical 1/2; lip purple or maroon; central lobe of lip truncate, about as long as wide; plants from tepui summits usually above 1600 m ........................................ M. quelchii 45. Sepals and petals white; lip white with deep orange at central lobe and apices of lateral lobes; central lobe of lip rounded or obtuse, longer than wide; plants from lowlands to 1500 m (very rarely higher) ............... 46 46(45). Pseudobulbs oblique with a ring at their apical zone; the external sheaths of the pseudobulbs bearing small leaf blades while the innermost are similar to the apical leaf; margin of clinandrium ciliate; flowering 1–3 contemporaneously .......................................................... M. splendens 46. Pseudobulbs not oblique and without a ring below point of leaf abscission; blades of sheaths all subequal; margin of clinandrium glabrous; flowers many simultaneously ........................................................................... 47 47(46). Floral bracts longer than pedicellate ovary; inflorescences shorter, subequaling to 1.4 times longer than pseudobulb; calli 3–5 times longer than wide ............................................................................... M. chlorantha 47. Floral bracts shorter to subequaling pedicellate ovary; inflorescences always longer than pseudobulb, 1.5–2 times longer than the pseudobulb; calli 2 times longer than wide .............................................. M. ochroleuca 48(42). Floral bract < 1/4 the length of pedicellate ovary; inflorescences with peduncles shorter to subequaling the pseudobulbs, rarely somewhat longer .................................................................................................... 49 48. Floral bracts > 1/2 the length of pedicellate ovary; inflorescences longer or shorter than the pseudobulb ............................................................... 53 49(48). Leaves fleshy-coriaceous, often furrowed on drying, 5–22 mm wide; pseudobulbs almost totally hidden by leaf sheaths, to 4.2 cm long (usually smaller) ......................................................................................... 50 49. Leaves coriaceous, 29–60 mm wide; pseudobulbs not totally hidden by leaf sheaths, 5–15 cm long .......................................................................... 52 50(49). Lip 20–40 mm long, 8–12 mm wide; dorsal sepal 24–37 mm long ..................................................................................... M. violaceopunctata 50. Lip 10–16 × 4–7 mm; dorsal sepal 12–22 mm long ................................. 51 51(50). Leaves 3.5–7 times longer than wide, the lower surface usually purple- or maroon-tinged; petals 4–5 times longer than wide; lip shallowly 3-lobed ........................................................................................ M. discolor

432

O RCHIDACEAE

51. 52(49).

52.

53(48). 53. 54(53).

54.

55(54). 55. 56(55).

56.

57(56).

57.

58(56).

58.

59(53). 59.

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

Leaves at least 10 times longer than wide, concolorous; petals 3–3.5 times longer than wide; lip conspicuously 3-lobed ............................... M. villosa Leaves 5–10 mm wide, the pseudopetioles usually twisted; petals 10– 10.5 mm long, 3.2–3.7 times longer than wide; central lobe of lip broadly ovate or broadly triangular ........................................ M. santanae Leaves (6–)12–23 mm wide, the pseudopetioles straight or only inconspicuously twisted; petals 12–17 mm long, 4–5.5 times longer than wide; central lobe of lip triangular ......................................... M. superflua Lip simple or nearly so, when there is some lobing, no sharp sinus separates lateral lobes from central, or lip is slightly pandurate ............. 54 Lip sharply 3-lobed .................................................................................. 59 Leaves 4–14 mm wide, fleshy-coriaceous; lip apically rounded, basally gradually attenuated; column foot longer than the column ................... ............................................................................................. M. bolivarensis Leaves (16–)23–80 mm wide, coriaceous; lip apically obtuse or truncate, basally rounded or obtuse, if attenuated then shortly and abruptly so; column foot shorter than the column, rarely nearly equaling it ........ 55 Dorsal sepal 2.5–2.9 times longer than wide, acute or obtuse ..... M. eburnea Dorsal sepal 3.4–5.2 times longer than wide, acuminate to rarely acute .............................................................................................................. 56 Pseudobulbs surrounded by 1(2) leaf-bearing sheaths, these conspicuously smaller than the pseudobulb leaf; column foot about 1/2 the length of column .............................................................................................. 57 Pseudobulbs surrounded by (2)3–5 leaf-bearing sheaths, these sheaths ± similar to the pseudobulb leaf; column foot about < 1/4 the length of column .................................................................................................. 58 Flowers with sepals externally dull maroon, internally dull orange-maroon, lip dull yellow tinged with maroon toward base, underside of lip with dull dark maroon toward apex; peduncle of inflorescence at least 5 times longer than pseudobulbs; lip broader toward apex or middle ................................................................................................. M. colemanii Flowers with sepals white or pale yellow with a yellow lip; peduncle of inflorescence as long as to 3 times longer than pseudobulbs; labellum broader toward base ................................................................. M. albiflora Sepals widely spreading, apices obtuse or acute; bracts of the inflorescence not inflated; floral bract shorter than the pedicellate ovary .............................................................................................. M. proboscidea Sepals either subparallel to the petals or reflexed, apices acuminate; lip subacuminate or acute; bracts of the inflorescence usually ± inflated; floral bract longer or nearly equaling the pedicellate ovary ...... M. nasuta Plants small to minute; leaves < 35 × 11 mm; pseudobulbs to 9 mm long; leaf apex minutely serrulate; lip 7–11 mm long ................................. 60 Plants larger; leaves at least 80 mm long (usually > 100 mm long), 9.5– 80 mm wide; pseudobulbs 15–95 mm long; leaf apex entire; lip 9.5– 20 mm long ........................................................................................... 62 Leaves 10–11 mm wide; lip 10–11 mm long ......................... M. brachybulbon Leaves 4–8 mm wide; lip 6–8 mm long ................................................... 61 Leaves linear oblong, acute; sepals 7–10 times longer than wide; column

Maxillaria 433

61. 62(59). 62. 63(62). 63. 64(62). 64.

65(64). 65. 66(65). 66. 67(66).

67.

68(65). 68. 69(68).

69.

foot well developed; lobes of labellum acute ....................... M. xylobiiflora Leaves linear-spathulate, obtuse; sepals 4–5 times longer than wide; column foot inconspicuous; lobes of lip obtuse to rounded ...... M. pterocarpa Sepals obtuse or rounded ......................................................................... 63 Sepals acute or acuminate ....................................................................... 64 Flowers brownish or reddish; leaves 2–6 times longer than wide, petiole of pseudobulb leaf about 1/2 the blade length ........................ M. cryptobulbon Flowers whitish or pale yellow; leaves 10 times longer than wide; petiole of pseudobulb ± equaling leaf length ....................................... M. connellii Sepals acute; petals and sepals bright yellow; lip entirely deep purple; mature leaves 23–90 cm long ...................................................... M. nasuta Sepals acuminate; petals and sepals yellowish, greenish, purple or white; lip variously colored, never deep purple (if so, only central lobe); mature leaves to 30 cm long ..................................................................... 65 Dorsal sepal 2–3 times longer than lip .................................................... 66 Dorsal sepal to l.5 times longer than lip ................................................. 68 Leaves 10 times longer than wide .................................................... M. tenuis Leaves 2–6 times longer than wide ......................................................... 67 Leaf on top of the pseudobulb sessile or inconspicuously petioled, when rarely present petiole < 1/5 the length of the leaf blade; pseudobulbs apically symmetrical, broadly elliptic to suborbicular in outline, clothed with 1 leaf-bearing sheath at each side; leaves white-spotted, central lobe of lip obtuse, borne in the apical 3/4 of total lip length; plants usually from elevations above 600 m .............. M. reichenheimiana Leaf on top of the pseudobulb conspicuously petioled, petiole at least 1/4 the total length of the leaf blade; pseudobulbs apically oblique; elliptical to narrowly oblong-elliptical, clothed with (1)2 leaf-bearing sheaths on each side; leaves plain green; central lobe of lip acute, borne in the apical 2/3 of total lip length; plants from elevations below 300 m ................................................................................................ M. pauciflora Lip 4 times as long as wide, its apical lobe occupying its apical 1/4; petiole of pseudobulb leaf at least 1/3 of blade length ......................... M. nuriensis Lip 2–3 times as long as wide, its apical lobe occupying its apical 1/3 or 1/2; petiole of pseudobulb leaf < 1/4 of blade length .............................. 69 Leaves elliptic, 3–4 times as long as wide; sheaths not verrucosepustulose; pseudobulbs ovoid or circular; central lobe of lip triangular, representing about 1/2 of lip total length; plants from cloud forests, 1200–2500 m; perianth segments red-purple ................ M. guadalupensis Leaves linear-oblong, > 10 times as long as wide; sheaths verruculosepustulose; pseudobulbs oblongoid; central lobe of lip ovate, representing 1/3 of lip length, flowers white; plants from lowland rainforests ....................................................................................................... M. tenuis

Maxillaria acutifolia Lindl., Edwards’s Bot. Reg. 25: misc. 92. 1839. Maxillaria rufescens auct. non Lindl. 1835 [1836]: sensu Foldats in Lasser, Fl. Venez. 15(4): 521. 1970, pro parte; Dunst. & Garay, Venez. Orchid. Ill. 2:

218. 1961. Epiphyte, usually growing in shady places; flowers sweet-fragrant; dorsal sepal and petals hooded over the column; petals and sepals greenish or yellowish to translucent white, often with pink to maroon tinges;

434

O RCHIDACEAE

lip cream to orange with pale or dark redmaroon spots. Uncommon in montane forests, 1000–1400 m; Bolívar (La Escalera to Cerro Venamo region). Aragua, Carabobo, Distrito Federal, Portuguesa; Colombia, Ecuador, Peru, Guyana, Amazonian Brazil. Maxillaria acutifolia is usually included within a broadly conceived M. rufescens. It is a smaller-flowered member of the M. rufescens complex, and is common in Guyana, Suriname, French Guiana, and the Venezuelan coastal range, but its distribution outside these areas is uncertain due to confusion with members of the M. rufescens complex. Maxillaria alba (Hook.) Lindl., Gen. Sp. Orchid. Pl. 143. 1832. —Dendrobium album Hook., Exot. Fl. 2: t. 142. 1825. —Camaridium album (Hook.) Hoehne, Arq. Bot. Estado Sâo Paulo 2: 72. 1947. Maxillaria lactea Schltr., Repert. Spec. Nov. Regni Veg. Beih. 29: 233. 1923. Epiphyte; erect to subpendent, usually sun-loving; leaves 15–50 cm long, linear to linear-ligulate; flowers not widely spreading, white with some yellow at lip; dorsal sepal 17–27 mm long. Locally common in lower montane forests, 400–600 m; Bolívar (Río Carrao), Amazonas (Río Ararí). Anzoátegui, Aragua, Miranda, Barinas; Guatemala, Honduras, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil (Amazon basin, Bahia). Maxillaria albiflora Ames & C. Schweinf., Sched. Orchid. 8: 56. 1925. Maxillaria melina auct. non Lindl.: sensu Foldats in Lasser, Fl. Venez. 15(4); 481. 1970, pro parte; Dunst. & Garay, Venez. Orchid. Ill. 1: 228. 1959; 2: 204. 1961. Maxillaria anatamorum auct. non Rchb. f.: sensu Foldats in Lasser, Fl. Venez. 15(4): 410. 1970, pro parte. Epiphyte, small to medium-sized, cespitose; pseudobulbs small with leaf-bearing sheaths; leaves grayish on drying; inflorescences longer than pseudobulb; flowers medium-sized to rather large and showy, with perianth segments subparallel to column, white with a yellow callus and some purple or maroon in the inner basal part of the lip. Montane and tepui forests, 800–2000 m; Bolívar (Kukenán-tepui, La Escalera to

Cerro Venamo region, Macizo del Chimantá, Ptari-tepui). Distrito Federal, Miranda, Nueva Esparta, Sucre, Táchira; Trinidad-Tobago, Guyana. Maxillaria connelli and M. nuriensis are closely related but clearly distinct taxa. Maxillaria albiflora, as presently circumscribed, includes much internal variation, and might include several different species. A thorough revision of the smaller-flowered of the M. grandiflora complex (e.g., the species related to M. melina) is required. Maxillaria alpestris Lindl. in Benth., Pl. Hartw. 154. 1839 [1845]. Epiphyte or lithophyte, sun-loving, creeping and occasionally forming large, dense masses; flowers opening well, medium-sized to small, with reddish brown perianth segments, a yellow to salmon red lip. Cloud forests at tepui summits or slopes, usually in relatively open places, 1300–2300 m; Bolívar (Auyán-tepui, Jaua-Sarisariñama massif, La Escalera to Cerro Venamo region, Macizo del Chimantá, Ptari-tepui), southern Amazonas (Cerro Aracamuni, Cerro Avispa, Sierra de la Neblina). Colombia, Guyana, Ecuador, Peru, Bolivia. ◆Fig. 391. The systematics of Maxillaria alpestris and its allies is unclear and for this reason no synonyms are cited. The distribution of this species and related taxa will only be determined with certainty when the systematics of the M. alpestris complex is resolved. Maxillaria alticola C. Schweinf., Bot. Mus. Leafl. 11: 261. 1945. Ornithidium serrulatum Lindl. in Benth., Pl. Hartw. 153. 1839 [1845], non Maxillaria serrulata Ames & Correll 1943. Epiphyte or terrestrial, monopodial or pseudomonopodial, usually growing in open places; stems cane-like, 20–100 cm long; flowers yellow; dorsal sepal 6–9.5 mm long. Locally common in tepui shrublands, 1900– 2600 m; Bolívar (Kamarkawarai-tepui, Macizo del Chimantá), Amazonas (Cerro Marahuaka, Sierra de la Neblina). Mérida, Táchira; Colombia, Ecuador, Peru. Maxillaria aurea (Poepp. & Endl.) L.O. Williams, Caldasia 1(3): 14. 1941. —Ornithidium aureum Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 57, t. 96. 1836.

Maxillaria 435

Usually terrestrial but often an epiphyte, 30–100(–150) cm tall, erect, sun-loving; sepals 8–11 mm long. Montane to upper montane forests, where locally common on open, steep slopes, (900–)1200–2100(–2400) m; Bolívar (Auyán-tepui, Ilú-tepui, JauaSarisariñama massif, La Escalera to Cerro Venamo region, Macizo del Chimantá, Roraima-tepui), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Marahuaka, Cerro Sipapo, Cerro Yaví, Sierra de la Neblina). Distrito Federal, Táchira; Colombia, Ecuador, Peru, Amazonian Brazil, Bolivia. ◆Fig. 394. Maxillaria auyantepuiensis Foldats, Bol. Soc. Venez. Ci. Nat. 22: 266. 1961. Sand-loving herb or lithophyte, sun-loving; flowers vertical, with parallel perianth segments, yellowish heavily tinged with pale to dark maroon; lip of a darker hue; dorsal sepal 20–21 mm long. Open sandy places or low open sclerophyllous shrublands on sandy substrate or sandstone outcrops; 500–1400 m; Bolívar (Gran Sabana, Jaua-Sarisariñama massif). Guyana, Suriname, Ecuador, Peru, Brazil (Roraima). Most reports of Maxillaria auyantepuiensis for other countries outside of the Guyana, Suriname, and French Guiana area are referable to the closely related M. longipetiolata. Maxillaria bolivarensis C. Schweinf., Bot. Mus. Leafl. 20: 22. 1962. Epiphyte; pseudobulbs hidden by 2 leafbearing sheaths, sheaths and inflorescence bracts scarious, dark purple-punctulate; flowers large for the plant; perianth segments and lip white, yellowish, or greenish, frequently with darker-hued tips; callus yellow; dorsal sepal 25–38 mm long. Lowland to montane forests, 50–1400 m; Delta Amacuro (Río Amacuro, Serranía de Imataca), Bolívar (Altiplanicie de Nuria, Cerro Venamo, Gran Sabana, Jaua-Sarisariñama massif, Serranía de Imataca), Amazonas (Sierra Parima). Ecuador, Peru, Brazil. ◆Fig. 396. Maxillaria brachybulbon Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19: 55. 1923. Epiphyte, minute to small; pseudobulbs with 3 or 4 leaf-bearing sheaths; inflorescence longer than pseudobulb; flowers large

for the plants; perianth segments subparallel, yellow or greenish; lip yellow, longitudinally purple-striped or purple at the base; dorsal sepal 15–18 mm long. Rain forests, 100–1400 m; Bolívar (La Escalera to Cerro Venamo region), Amazonas (Cerro Marahuaka, Río Atacavi, Serranía Batata). Sucre; Costa Rica, Guyana, Suriname, French Guiana, Ecuador, Brazil. ◆Fig. 399. Maxillaria camaridii Rchb. f., Allg. Gartenzeitung 19: 547. 1863. Camaridium amazonicum Schltr., Beih. Bot. Centralbl. 42(2): 135. 1925. Camaridium ochroleucum Lindl., Edwards’s Bot. Reg. 10: t. 844. 1824. —Cymbidium ochroleucum (Lindl.) Lindl., Gen. Sp. Orchid Pl. 168. 1833. Epiphyte, erect to pendulous, sun-loving; perianth segments white with some yellow on lip; dorsal sepal 18–35 mm long. Locally common in moist forests, 50–400 m; Delta Amacuro (Río Grande, Güiniquina), Bolívar (throughout), Amazonas (throughout). Common elsewhere in Venezuela; Guatemala to Panama, West Indies, Colombia, Suriname, Ecuador, Peru, Brazil. Maxillaria cozieriana H.G. Jones, described from neighboring Guyana, is very closely related to M. camaridii, but apparently has smaller flowers of a greenish color. The type of the species, however, was not available for comparison so the actual status of M. cozieriana is impossible to ascertain at this time. Maxillaria chlorantha Lindl., Edwards’s Bot. Reg. 23: sub t. 1986. 1837. Epiphyte; pseudobulbs flattened; perianth segments subparallel, white, tinged with pale yellow at apex; lip apically bright orange; dorsal sepal 40–50 mm long. Cloud forests, 1400–1500 m; Amazonas (Cerro Aracamuni). Táchira; Colombia, Guyana, Brazil. Maxillaria colemanii Carnevali & W. Fritz, Selbyana 21: 145. 2000. Terrestrial or lithophyte (perhaps also sometimes an epiphyte), cespitose; sepals externally dull maroon, internally dull orangemaroon; dorsal sepal 23–25 mm long, tinged with dull maroon toward base; petals exter-

436

O RCHIDACEAE

nally pale cream tinged with dull maroon toward the apices; internally dull cream yellow; lip dull yellow with dull maroon tinges toward base; callus and central lobe pale bright yellow, underside of lip clear yellow with dull dark maroon at base of lateral lobes; column and anther dull pale yellow. Usually growing under shrubs in open shrublands, ca. 1500 m; Bolívar (La Escalera to Cerro Venamo region). Endemic. ◆Fig. 390. Maxillaria colemanii is closely related to M. porrecta, but is easily distinguished by the leaf-bearing sheaths enveloping the pseudobulb. It probably also occurs in neighboring Guyana. Maxillaria conferta (Griseb.) C. Schweinf. ex León, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 8: 395. 1946. —Ornithidium confertum Griseb., Fl. Brit. W. I. 626. 1864. Ornithidium chloroleucum Barb. Rodr., Gen. Spec. Orchid. 2: 208. 1882. —Pseudomaxillaria chloroleuca (Barb. Rodr.) Hoehne, Arq. Bot. Estado São Paulo n.s. formato maior 2: 72. 1947. Ornithidium vestitum Rchb. f. in Walp., Ann. Bot. Syst. 6: 491. 1863, sine syn. et non (Sw.) Rchb. f. nec auct. post. Maxillaria surinamensis H. Focke ex Rchb. f. in Walp., Ann. Bot. Syst. 6: 492. 1863. Maxillaria similans Ames & C. Schweinf., Sched. Orchid. 10: 99. 1930. Maxillaria purpurea auct. non (Spreng.) Ames & Correll 1943: sensu Foldats in Lasser, Fl. Venez. 15(4): 511. 1970. Epiphyte or rarely a lithophyte; rhizome ascendent to pendulous, totally enclosed in tubular, scarious, imbricate sheaths; leaves 5–27 cm long; flowers campanulate, white with yellow in the lip; dorsal sepal 3.5–5.5 mm long. Rain forests, 100–600(–1000) m; Delta Amacuro (Río Amacuro), Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region, Río Canaracuni, Río Paragua), Amazonas (Cerro Huachamacari, Río Ararí, near San Carlos de Río Negro, Sierra Parima). Aragua, Distrito Federal, Miranda, Monagas, Portuguesa, Táchira, Yaracuy; TrinidadTobago, Guyana. Maxillaria connellii Rolfe, Trans. Linn. Soc. London, Bot. 6: 63. 1901.

Epiphyte, shade-loving; pseudobulbs small for plant size; flowers white, with perianth segments subparallel to column, sepals 14– 20 mm long; column foot longer than column. Cloud forests, 1600–2000 m; Bolívar (Macizo del Chimantá, Ptari-tepui, Roraima-tepui). Distrito Federal, Monagas; Guyana. ◆Fig. 403. Maxillaria cryptobulbon Carnevali & J.T. Atwood, Novon 1: 159. 1991. Maxillaria brunnea auct. non Linden & Rchb. f. 1854: sensu Dunst. & Garay, Venez. Orchid. Ill. 1: 221. 1962. Maxillaria trinitatis auct. non Ames 1923: Dunst. & Garay, Venez. Orchid. Ill. 2: 223. 1962. Epiphyte or rarely a lithophyte, 20–45 cm tall; leaves (20–)25–35(–42) cm long; flowers with subparallel segments, sepals dark dull red-brown or bright maroon-red; petals pink or yellow-orange within, tinged with dull red-brown without; lip yellow cream with bright yellow callus, the underside of the central lobe very dark red-purple; dorsal sepal 26–28 mm long. Lower montane forests, 400–700 m; Bolívar (Altiplanice de Nuria, La Escalera to Cerro Venamo region), Amazonas (Sierra de la Neblina). Miranda; Costa Rica, Ecuador, Peru. The presence of leaf-bearing sheaths enveloping the pseudobulbs is a variable character in Maxillaria cryptobulbon. Some individuals have both pseudobulbs that are enveloped by the sheaths and others that are not. Maxillaria desvauxiana Rchb. f., Bonplandia (Hanover) 3: 67. 1855. Maxillaria coriacea Barb. Rodr., Gen. Spec. Orchid. 2: 199. 1882. Maxillaria petiolaris A. Rich. in Saunders, Refug. Bot. 2: sub t. 134. 1882, nom. nud. Maxillaria verrucifera C. Schweinf., Bot. Mus. Leafl. 11: 292, fig. 19. 1945. Epiphyte or lithophyte; leaves 15–30 cm long; petioles 3–15 cm long; flowers erect, showy; perianth segments yellow-brown to red-brown with a red-brown lip; sepals spreading, petals subparallel to column; dorsal sepal 20–32 mm long. Lowland to montane forests or exposed cliffs, 50–1400 m; Bolívar (Río Carrao), Amazonas (basins of Río Casiquiare, Río Sipapo, and Río Ven-

Maxillaria 437

tuari). Miranda; Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Amazonian states). ◆Fig. 405. Maxillaria discolor (Lodd. ex Lindl.) Rchb. f. in Walp., Ann. Bot. Syst. 6: 529. 1863. —Dicrypta discolor Lodd. ex Lindl., Edwards’s Bot. Reg. 25: 91. 1839. Dicrypta bicolor Paxton ex Planch., Hort. Donat. 73. 1858. Epiphyte, shade-loving; leaves 10–35 cm long, frequently purplish on the lower surface; flowers fleshy, fetid; perianth segments yellow; lip yellow-orange or orange with purple spots; callus waxy; dorsal sepal 12–22 mm long. Lowland to (more rarely) cloud forests, 50–600(–1000) m; Delta Amacuro (Río Acure), Bolívar (basin of Río Caroní and Río Cuyuní). Apure, Miranda, Sucre, Táchira; Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 393. Maxillaria ×dunstervillei Carnevali & I. Ramírez, Novon 3: 106. 1993. Epiphyte or humicolous herb, shade-loving; flowers showy, variable in color, perianth segments dull or bright yellow to orange-yellow with paler bases, with some maroon tinges; lip pale cream yellow, with a dark maroon patch on the underside of the isthmus; dorsal sepal 31–48 mm long. Local on bogs or tepui shrublands, 2000–2400 m; Amazonas (Sierra de la Neblina). Brazil (Serra da Neblina). The available evidence strongly suggests that Maxillaria ×dunstervillei is a natural hybrid between M. meridensis Lindl. and M. quelchii Rolfe. Although it has been only recorded so far from the summit of Sierra de la Neblina, Maxillaria ×dunstervillei is likely to occur in other tepuis where both putative parents are sympatric and appropriate environmental conditions are available. The plants of this nothotaxon sometimes will key out to any of the putative parents. Maxillaria eburnea Lindl., Sert. Orch. t. 40, fig. 2. 1840. Maxillaria grandiflora auct. non (H.B.K.) Lindl. 1832: sensu Foldats in Lasser, Fl. Venez. 15(4): 454. 1970, pro parte. Epiphyte or muscicolous herb; leaves 11– 30 cm long; flowers showy with sepals 3.5–5 cm long, with spreading sepals and petals

parallel to column, all perianth segments white except the yellow lip apex or some purple at its margins. Cloud forests, elevation uncertain but probably high; Amazonas (near Cerro Marahuaka). Endemic. This apparently showy species has not been recollected. Maxillaria equitans (Schltr.) Garay, Bot. Mus. Leafl. 18: 208. 1958. —Camaridium equitans Schltr., Repert. Spec. Nov. Regni Veg. Beih. 7: 176. 1920. —Marsupiaria equitans Arq. Bot. Estado São Paulo 2: 71. 1947. Camaridium vandiforme Schltr., Beih. Bot. Centralbl. 42(2): 174. 1925. —Maxillaria vandiformis (Schltr.) C. Schweinf., Bot. Mus. Leafl. 11: 291. 1945. Maxillaria mattogrossensis Brade, Arq. Serv. Florest. 1: 46. 1939. —Marsupiaria mattogrossensis (Brade) Hoehne, Arq. Bot. Estado São Paulo 2: 71. 1947. Epiphyte, erect, arching, or pendulous; leaves distichous along the stem; inflorescences originating from leaf axils; flowers with widely spreading sepals; petals subparallel to the column; perianth segments white or yellowish; lip dark maroon with a yellow or whitish apex; dorsal sepal 14–17 mm long. Moist lowland forests, 100–300 m; Amazonas (Paso de Ganado, Río Mavaca). Apure, Táchira; Colombia (Amazonian states), Ecuador, Peru, Brazil, Bolivia. Maxillaria vandiformis and M. mattogrossensis might eventually prove to be distinct species. The laterally compressed leaves on a pseudomonopodial stem are a unique feature of these species. Maxillaria foldatsiana Carnevali & I. Ramírez, Ann. Missouri Bot. Gard. 76: 376. 1989. Apparently terrestrial; leaves 23–25 cm long; flowers erect, medium-small with perianth segments and lip subparallel to the column; dorsal sepal 2–2.1 cm long. Montane forests, ca. 1200 m; Bolívar (Cerro Guaiquinima, Gran Sabana, Uaipán-tepui). Guyana. ◆Fig. 395. Maxillaria grobyoides Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 5: 186. 1972.

438

O RCHIDACEAE

Epiphyte; pseudobulbs ovoid or orbicular; leaves 15–20 cm long; inflorescences longer than pseudobulbs; flowers with spreading segments; perianth segments yellow, yelloworange, or greenish yellow; lip yellow with purple-brown or orange-brown at callus and central lobe. Lower montane forests, 400– 900 m; Bolívar (upper Río Caroní basin), Amazonas (Cerro Aracamuni). Amazonian portions of Colombia, Guyana, Ecuador, Peru, Brazil. ◆Fig. 406. Maxillaria guadalupensis Cogn. in Urb., Symb. Antill. 6: 604. 1910. Maxillaria attenuata Ames & C. Schweinf., Sched. Orchid. 10: 89. 1930. Epiphyte to 10 cm tall; flowers large for the plant with subparallel perianth segments; perianth segments yellow or maroonred with reddish tips and a red or pink lip; dorsal sepal 1.8–3 cm long. Cloud forests 1300–2500 m; Bolívar (La Escalera to Cerro Venamo region, Ptari-tepui, Sierra Parima), Amazonas (Cerro Marahuaka). Guadalupe, Colombia, Ecuador, Peru. ◆Fig. 409. As presently circumscribed, Maxillaria guadalupensis has a lot of internal variation and might include several distinct species. Maxillaria guareimensis Rchb. f., Bonplandia (Hanover) 2: 16. 1854. Maxillaria spilotantha auct. non Rchb. f. 1854: sensu Foldats in Lasser, Fl. Venez. 15(4): 458. 1970. Epiphyte, medium-sized to rather large, cespitose, erect to pendulous; plant attached to substrate only at base, with elongate stems bearing pseudobulbs at internodes; inflorescences originating from leaf sheaths; flowers small, purple or yellow; lip purple or yellowish lip with a wet black “callus.” Moist montane forests, 700–1200 m; Bolívar (Cerro Guaiquinima, Roraima-tepui), Amazonas (Cerro Yutajé). Aragua, Falcón, Miranda, Yaracuy, Zulia; Colombia, Ecuador, Peru. ◆Fig. 404. The distribution of Maxillaria guareimensis outside of the flora area will remain unclear until a revision of the M. guareimensis-spilotantha-saxatilis complex is completed. Maxillaria imbricata Barb. Rodr., Gen. Spec. Orchid. 1: 120. 1877. —Camari-

dium imbricatum (Barb. Rodr.) Hoehne, Arq. Bot. Estado São Paulo 4: 72. 1947. Maxillaria iguapensis Hoehne & Schltr., Arq. Bot. Estado São Paulo 1: 271. 1927. —Camaridium iguapensis (Hoehne & Schltr.) Hoehne, Arq. Bot. Estado São Paulo 4: 72. 1947. Epiphyte, erect to pendulous; flowers widely spreading; white or greenish yellow with pink or purplish on lateral lobes of the lip; dorsal sepal 12–20 mm long. Moist montane forests, 1000–1200 m; Bolívar (upper basins of Río Caroní and Río Cuyuní). Aragua, Distrito Federal, Falcón, Mérida, Yaracuy; Suriname, Ecuador, Brazil, Bolivia. Material that has been included in Maxillaria imbricata from southern Central America is probably best referred to another species, such as M. bracteata (Schltr.) Ames & Correll or M. gomeziana J.T. Atwood. Maxillaria kegelii Rchb. f., Linnaea 41: 127. 1877. Maxillaria amazonica Schltr., Beih. Bot. Centralbl 42(2): 130. 1925. Maxillaria lactiflora Pabst, Orquídea (Niteroi) 29: 114. 1967. —Maxillaria lactea Schltr., Beih. Bot. Centralbl. 42(2): 131. 1925, non Maxillaria lactea Schltr. 1923, syn. nov. Maxillaria taracuana Schltr., Repert. Spec. Nov. Regni Veg. Beih. 42: 134. 1925. Epiphyte, medium-sized, cespitose, erect; pseudobulbs 1- leaved, without leaf-bearing sheaths; inflorescences shorter than leaves; flowers medium-sized with widely spreading sepals and petals subparallel to column, white with some yellow on labellum. Evergreen lowland forests, 100–200 m; Amazonas (Río Mawarinuma). Brazil (Amazonas). Guyana, Suriname, French Guiana. ◆Fig. 392. This species belongs in the very difficult Maxillaria brunnea Linden & Rchb. f. complex. Within this complex, the Venezuelan material seems to match very well the description of M. taracuana Schltr., here considered a synonym of M. kegelii. Maxillaria lasallei Foldats, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 3: 2. 1961. Epiphyte or lithophyte, creeping to pendulous; pseudobulbs at intervals or clumped;

Maxillaria 439

inflorescences longer than pseudobulb; flowers yellowish; lateral sepals spreading; dorsal sepal and petals forming a hood over the column and lip; dorsal sepal 2–2.7 cm. long. Cloud forests, 700–1500 m; Bolívar (Auyántepui, La Escalera to Cerro Venamo region). Guyana. Maxillaria longipetiolata Ames & C. Schweinf., Sched. Orchid. 8: 61. 1925. Maxillaria riopalenquensis Dodson, Selbyana 7: 355. 1984. Maxillaria auyantepuiensis subsp. epiphytica Carnevali & I. Ramírez, Ann. Missouri Bot. Gard. 76: 375. 1989. Maxillaria auyantepuiensis auct. non Foldats sensu: Foldats in Lasser, Fl. Venez. 15(4): 419. 1970, pro parte; Dunst. & Garay, Venez. Orchid. Ill. 3: 181. 1965, pro parte; Bennet & Christenson, Icon. Orch. Peruv. Pl. 309, 1995. Epiphyte, usually shade-loving; flowers horizontal, with subparallel perianth segments; dorsal sepal 16–17 mm long. Lowland to montane forests, (100–)700–1400 m; Bolívar (Cerro Guaiquinima, La Escalera to Cerro Venamo region), Amazonas (Maroa). Barinas, Miranda; Costa Rica, Panama, Suriname, Ecuador, Peru. The plants of Maxillaria longipetiolata from the eastern side of the Andes seem to have smaller flowers with a somewhat different-shaped lip. A revision of the M. desvauxiana complex is urgently required to decide how many species there are around M. auyantepuiensis. If the eastern Andean, Amazonian, and Guyanan populations prove to be distinct, a new combination at the species level based on Maxillaria auyantepuiensis subsp. epiphytica Carnevali & I. Ramírez would have to be proposed. In the meanwhile, a conservative approach is here taken. Maxillaria loretoensis C. Schweinf., Amer. Orchid Soc. Bull. 13: 92. 1944. Epiphyte, erect, cespitose; inflorescences erect, longer than pseudobulb; flowers with widely spreading sepals and petals subparallel to the column; yellow or creamy yellow, often with longitudinal purple streaks internally; petals sometimes apically white, lip white with longitudinal purple streaks;

dorsal sepal 2.6–3.7 cm long. Rain forests, 50–400 m; Bolívar (Macizo del Chimantá), Amazonas (basin of Río Casiquiare and Río Negro). Amazon basin in Peru and Brazil. ◆Fig. 397. Maxillaria loretoensis is closely related to M. parkeri but the former has narrower leaves and sepals, and the lip with the central lobe broader at the apical half. Maxillaria luteoalba Lindl., Orchid. Linden. 20. 1846. Epiphyte, medium-sized to rather large, cespitose; pseudobulbs 1-foliate, without leaf-bearing sheaths, inflorescences longer than pseudobulb; flowers large and showy with spreading sepals; petals archedsubparallel to column; perianth segments white, lip yellowish with a white margin and lateral lobes purple-striped. Rain forests, 100–800 m; Bolívar (Cerro Guaiquinima), Amazonas (lower Río Ventuari basin). Distrito Federal, Mérida, Miranda; Costa Rica, Colombia, Ecuador. ◆Fig. 402. Maxillaria mapiriensis (Kraenzl.) L.O. Williams, Caldasia 3: 16. 1842. —Ornithidium mapiriense Kraenzl., Repert. Spec. Nov. Regni Veg. 25: 23. 1928. Maxillaria condensata C. Schweinf., Fieldiana, Bot. 28: 194, fig. 38. 1951. Epiphyte or rarely a lithophyte, mediumsized to rather large, creeping, bearing 1-foliate pseudobulbs at intervals; inflorescences originating directly from the rhizome internodes or from the leaf sheath axils; flowers minute to small with spreading sepals and petals parallel to column; perianth segments whitish, yellowish, or greenish, sometimes the petals brown or maroon; lip yellow, apically with maroon or violet blotches. Montane and cloud forests, often in exposed places, sometimes the plants from higher elevations with abbreviated internodes (compare M. condensata), 700–2600 m; Bolívar (widespread above 700 m), Amazonas (Cerro Aracamuni, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú, Sierra de la Neblina). Aragua; Colombia, Suriname, Ecuador, Brazil, Bolivia. Maxillaria mapiriensis is extremely variable vegetatively. The flowers are remarkably consistent in size, color, and overall morphology.

440

O RCHIDACEAE

Maxillaria marmoliana Dodson, Icon. Pl. Trop. 1: 156. 1980. Maxillaria arachnitis auct, non Rchb. f.: sensu Foldats in Lasser, Fl. Venez. 15(4): 414. 1970. Epiphyte, cespitose; inflorescences longer than pseudobulbs; leaves 15–30 cm long, linear-lanceolate; flowers showy with yellowgreen sepals, basally purple and brown; petals white; lip yellow or white with brown or brown-purple central lobe. Montane forests, 500–1500 m; Bolívar (Río Carrao), Amazonas (Cerro Aracamuni). Miranda; Ecuador. Maxillaria marmoliana is one of the few taxa of the M. lepidota complex in the lowlands east of the Andes. Maxillaria meridensis Lindl., Orchid. Linden. 19. 1846. Epiphyte, lithophyte, or subterrestrial, medium-sized to large, creeping, erect, or scandent; rhizomes elongate, sometimes branching, bearing pseudobulbs only at mature parts of the sympodium; flowers emerging from rhizome leaf axils, small to medium-sized; perianth segments yellowish, brownish yellow to orange, subparallel. Cloud forests, frequently in open places, 1300–2500 m; common species in highlands of the whole flora area. Aragua, Mérida, Yaracuy; West Indies, Colombia, Guyana, Ecuador, Peru, Brazil (Amazonas, Roraima), Bolivia. Maxillaria nasuta Rchb. f., Beitr. Orchid.K. C. Amer. 104. 1866 [1867]. Maxillaria brevipedunculata Ames & C. Schweinf., Sched. Orchid. 10: 91. 1930. Epiphyte, lithophyte, or terrestrial, medium-sized to rather large, cespitose; pseudobulbs 1-foliate, with 2–4 leaf-bearing sheaths, inflorescences relatively short, but longer than pseudobulb; flowers mediumsized to rather large with widely spreading sepals and petals subparallel to the column; perianth segments yellow or greenish yellow, frequently with purple or greenish apex; lip shiny red, purple, or maroon, frequently with a greenish or yellowish apex. Lowland to montane forests, frequently in open places, (100–)400–1200(–1700) m; Bolívar (Río Caroní and Río Cuyuní basins), Amazonas (Cerro Marahuaka, Río Sipapo and Río Casiquiare basins, Sierra Parima). Aragua, Distrito Federal, Miranda, Sucre,

Yaracuy; Mexico, Central America, Colombia, Guyana, Ecuador, Peru, Brazil (Amazonas, Roraima), Bolivia. Maxillaria notylioglossa Rchb. f., Bonplandia (Hanover) 2: 16. 1854. Epiphyte or rarely a lithophyte, small to medium-sized, rhizomatous, creeping; pseudobulbs 2-foliate, inflorescences shorter than leaves, flowers small, with widely spreading sepals and petals subparallel to column; perianth segments greenish or yellowish; lip with a white waxy callus. Lower to montane forests, 400–2000 m; Bolívar (Río Caura basin), Amazonas (Cerro Aratitiyope, Cerro Autana, Cerro Marahuaka, Cerro Sipapo, Río Ocamo basin). Aragua, Distrito Federal, Falcón, Lara, Miranda, Portuguesa, Sucre, Táchira, Yaracuy; Colombia, Ecuador, Brazil (Amazonas, Roraima), Bolivia. Maxillaria nuriensis Carnevali & I. Ramírez, Novon 3: 109. 1993. Epiphyte, medium-sized, cespitose; pseudobulbs 1-foliate, with leaf-bearing sheaths; flowers relatively small with subparallel perianth segments; perianth segments creamy. Low-altitude cloud forests, 600–700 m; Bolívar (Altiplanicie de Nuria). Endemic. ◆Fig. 411. Maxillaria ochroleuca Lodd. ex Lindl., Gen. Sp. Orchid. Pl. 143. 1832. Colombia, Ecuador; 2 subspecies, 1 in Venezuela. M.

ochroleuca var. longipes Sander, Sander’s Orch. Guide 121. 1901. Terrestrial, lithophyte, or epiphyte, medium-sized, cespitose, erect; pseudobulbs flattened, 1-foliate, with 1–3 leaf-bearing sheaths; inflorescences much shorter than leaves; flowers medium-sized, with subdivergent sepals and petals subparallel (but crossing over) the lip; perianth segments white grading to yellow at apices; lip creamy white with an orange central lobe. Lowland to montane forests, 400–1500 m; Bolívar (JauaSarisariñama massif, upper Río Caura basin), Amazonas (Cerro Aracamuni, Cerro Autana, Sierra Parima). Brazil. Maxillaria parkeri Hook., Bot. Mag. 54: t. 2729. 1827.

Maxillaria 441

Epiphyte, medium-sized to rather large, cespitose, erect; pseudobulbs without leafbearing sheaths; inflorescences longer than pseudobulbs; flowers medium-sized to rather large with widely spreading sepals and petals subparallel to column; sepals buff to clear yellow, darker apically, often flushed with purple externally; petals yellow or white, sometimes with basal purple lines; lip white or clear yellow with purple streaks on lateral lobes and often white-bordered, central lobe. High and dense to low and open rain forests, 50–900 m; Bolívar (Río Caroní, Río Paragua), Amazonas (Río Casiquiare, Río Cataniapo, Río Sipapo). Guyana, Ecuador, Peru, Brazil (Amazonas, Pará, Roraima). ◆ Fig. 398. See coments under Maxillaria loretoensis. Maxillaria pauciflora Barb. Rodr., Gen. Spec. Orchid. 1: 116. 1877. Epiphyte 20–24 cm tall; leaf blades 14–15 × 2.5–2.8 cm; sepals widely spreading, petals subparallel to lip and column, petals and sepals pale yellow; lip yellow with deep redpurple longitudinal veins and central lobe; dorsal sepal 23–26 mm long, lip 12–14 mm long. Uncommon in lowland forests, 200–300 m; Amazonas (Misión Río Mavaca). Amazonian Brazil, possibly Amazonian Colombia. This rare species belongs in a complex of taxa around Maxillaria splendens, represented in the flora area by other species such as M. ochroleuca and M. tenuis. These taxa feature pseudobulbs that display an oblique petiole abscission line, (1)2 or 3 leaf-bearing sheaths, flowers with narrow perianth segments, and a narrow but thickly fleshy, blackish apical lobe of the lip. Most of these species occur at relatively low elevations and flower prolifically. Among other taxa in the complex that occur in the Venezuelan Guayana, M. pauciflora is easy to distinguish because of its smallish flowers produced singly or in pairs, and yellow perianth segments. Maxillaria porrecta Lindl., Edwards’s Bot. Reg. 24: misc. 192. 1838. Maxillaria brunnea Linden & Rchb. f., Bonplandia (Hanover) 2: 281. 1854. Maxillaria trinitatis Ames, Sched. Orchid. 2: 34. 1923. —Maxillaria trinitatis Ames ex Broadway, Orchid Rev. 34: 202. 1926, “trinitensis,” nom. superfl.

Maxillaria ringens auct. non Rchb. f. 1863: sensu Dunst. & Garay, Venez. Orchid. Ill. 2: 216. 1961. Maxillaria amazonica auct. non Schltr. 1925: sensu Dunst. & Garay, Venez. Orchid. Ill. 2: 198. 1961; Foldats in Lasser, Fl. Venez. 15(4): 409. 1970. Epiphyte, lithophyte, or rarely humicolous herb, generally sun-loving, cespitose; leaves 25–45 cm long, linear to elliptic; inflorescences usually longer than pseudobulb length; flowers with subparallel perianth segments, the lateral sepals frequently with the apices curved inward; perianth segments cream-yellow or orange-yellow, tinged with purple at apex; lip cream or orange with a deep purple central-lobe; dorsal sepal 2.5– 3.2 mm long. Locally common in lower montane to cloud forests, frequently in ± open places, 500–1500(–1800) m; Bolívar (widespread), Amazonas (Cerro Aracamuni, Sierra Parima). Carabobo, Miranda, Nueva Esparta, Sucre, Táchira; apparently widespread in tropical America, from south of Nicaragua to Ecuador and Peru. ◆Fig. 401. The name Maxillaria brunnea has been applied to this widely ranging, common species, but it is certainly conspecific with the earlier described M. porrecta. The type of M. brunnea is in very poor shape, but material collected very close to the type locality matches this widespread species closely. Maxillaria porrecta seems to be widespread and somewhat variable, but many of the synonyms usually attributed to M. brunnea (M. ringens Rchb. f., M. pubilabia Schltr., M. rousseauae Schltr., etc.) are different, though closely related species, and some of them are synonyms of other species (e.g., M. amazonica Schltr., M. taracuana Schltr., etc.). Until the systematics of this complex of species is better understood, we prefer not to include most of the names traditionally considered synonyms of M. brunnea in the synonymy of M. porrecta. Maxillaria porrecta is also distinguished usually by a glaucous bloom covering the lower surface of the leaves, which is entirely lacking in M. colemanii. Among the members of the M. porrecta complex, only M. cryptobulbon Carnevali & J.T. Atwood displays the same glaucous bloom. Maxillaria proboscidea Rchb. Bonplandia (Hanover) 2: 16. 1854.

f.,

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Maxillaria nasuta auct. non Rchb. f. 1866: sensu Dunst. & Garay, Orchids Venez. Ill. Field Guide 532. 1979; Foldats in Lasser, Fl. Venez. 15(4): 490. 1970, pro parte. Epiphyte or subterrestrial (in the flora area); flowers yellow with a purple or dark orange lip. Lowland forests, 100–200 m; Amazonas (Río Autana). Aragua, Distrito Federal. Specimens from the Coastal Range and from the flora area with obtuse sepals and not-inflated bracts of the inflorescence, previously identified as Maxillaria nasuta, seem to match very well the original description and illustration of M. proboscidea. The material from the flora area has 2-leaved pseudobulbs growing terrestrial on white sand savanna and might represent a different variety or species from typical M. prosbocidea.

ate elevations. Guyana, Ecuador, Brazil (Amazonas, Roraima). ◆Fig. 410.

Maxillaria pterocarpa Barb. Rodr., Gen. Sp. Orchid. 2: 204. 1882. Epiphyte 4–7 cm tall; leaves 3–5 cm long, 5–8 mm wide; sepals and petals greenish yellow, lip pale yellowish brown with pale purple veins, column white; anther yellow; dorsal sepal 7–10 mm long. Tepui-slope forests, ca. 700 m; Bolívar (Cerro Guaiquinima). Amazonian Brazil. This is a rare species in the flora area, from where it is known from a single collection. The plants here referred to Maxillaria pterocarpa are somewhat different from the type material and might represent another, possibly undescribed taxon.

Maxillaria reichenheimiana Endres & Rchb. f., Gard. Chron. 1871: 1678. 1871. Epiphyte, small, cespitose, erect; pseudobulbs 1-foliate, with 1 or 2 leaf-bearing sheaths; leaves white-spotted over a clear green or clear blue-green background; flowers medium-sized for the genus but large for the plant, with subparallel perianth segments; petals and sepals yellowish or orange, sometimes apically flushed with purple or brown; lip yellow or orange. Lower to montane forests, 600–1400 m; Bolívar (Altiplanicie de Nuria, Auyán-tepui, La Escalera to Cerro Venamo region, Río Uairén basin), Amazonas (Cerro Autana). Costa Rica, Colombia, Trinidad, Suriname, Ecuador, Peru.

Maxillaria quelchii Rolfe, Trans. Linn. Soc. London, Bot. 6: 64. 1901. Commonly lithophyte or subterrestrial, more rarely an epiphyte, medium-sized to rather large, with pseudobulbs closely disposed along an ascendent to erect rhizome, sometimes the rhizome very long; pseudobulbs 1-foliate with 1 or 2 leaf-bearing sheaths; flowers medium-sized to relatively large with subparallel perianth segments; sepals and petals red, brownish red or yellowish sometimes with white or yellow bases; lip brownish red. Cloud forests to rather open forests or tepui scrubs on tepui tops, 1600–2000 m; common and somewhat variable species in the flora area at appropri-

Maxillaria ramosa Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 1: 226. 1798. Scaphyglottis tafallae Rchb. f., Linnaea 22: 855. 1849. —Maxillaria tafallae (Rchb. f.) C. Schweinf., Bot. Mus. Leafl. 11: 288. 1945. Epiphyte, medium-sized to rather large, erect to pendulous, rhizomatous; pseudobulbs 1-foliate, with leaf-bearing sheaths at intervals along the rhizome; inflorescences short; flowers very small for the genus, with greenish, creamy, or creamy pink tepals and lip. Cloud forests, 700–1500 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region). Aragua, Carabobo, Falcón, Mérida, Miranda, Zulia; Colombia, Suriname, Ecuador, Peru.

Maxillaria rigida Barb. Rodr., Gen. Spec. Orchid. 2: 206. 1882. Camaridium pendulum Barb. Rodr., Gen. Spec. Orchid. 1: 123. 1877, non Maxillaria pendula (Poepp. & Endl.) C. Schweinf., Bot. Mus. Leaf. 11: 285. 1945. —Maxillaria pendens Pabst, Bradea 1: 176. 1972. Epiphyte or rarely a lithophyte, mediumsized to rather large; pseudobulbs distant on a strong, terete rhizome, 1- or 2-foliate, with 1–3 leaf-bearing sheaths; inflorescences racemose, originating from the axil of the sheaths, fasciculate; flowers small to medium-sized; dorsal sepal arching over col-

Maxillaria 443

umn; lateral sepals widely spreading; petals parallel to column; sepals apically yellowish, basally purplish; petals and column pale cream flushed with pink. Cloud forests, sometimes in rather open places, 1100–1600 m; Amazonas (Cerro Aracamuni, Cerro Autana, Cerro Duida, Cerro Marahuaka, Sierra Tapirapecó). Colombia, Ecuador, Peru, Amazonian Brazil. Reports of Maxillaria rigida from Bolivia at much higher elevations are almost certainly based on misidentifications of M. multicaulis (Poepp. & Endl.) C. Schweinf. Maxillaria rufescens Lindl., Edwards’s Bot. Reg. 21: sub t. 1802. 1836. Epiphyte or rarely a lithophyte, small to medium-sized, cespitose, rather variable; pseudobulbs 1-foliate, without leaf-bearing sheaths; inflorescences short; flowers small to medium-sized with spreading lateral sepals and subparallel dorsal sepal and petals, with a strong vanilla scent; sepals and petals in various shades of yellow or orange-yellow and a red- or purple-spotted lip. Lower to upper montane forests, 50–1300(–2000) m; Delta Amacuro (Río Cuyubini), Bolívar (Altiplanicie de Nuria, La Escalera to Cerro Venamo region, Macizo del Chimantá, Río Cuyuní basin, Río Paragua basin, Serranía de Imataca), Amazonas (Cerro Marahuaka, Río Casiquiare basin). Widespread elsewhere in Venezuela in the Amazon basin; Guyana, Suriname, French Guiana. Populations previously referred in the literature to Maxillaria rufescens from Central America, the Venezuelan Coastal Range, Andean South America above 1000 m elevation, the western side of the Andes, and southeastern Brazil belong to closely related but distinct taxa. Even after removal of the smaller-flowered M. acutifolia and taken in a broader sense, the present circumscription of M. rufescens probably contains several entities in the Venezuelan Guayana, some of which are undoubtedly valid species (e.g., M. cleistogama Brieg. & Bicalho). Flowers of the complex preserve poorly in herbarium specimens, which has obscured our current understanding of the group whose systematics will only be understood after much field work, plus herbarium and greenhouse studies are carried out with live plants and pickled flowers.

Maxillaria santanae Carnevali & I. Ramírez, Ann. Missouri Bot. Gard. 76: 377. 1989. Epiphyte, small, cespitose, shade-loving; pseudobulbs small, with leaf-bearing sheaths, leaves very fleshy, linear; inflorescence short; flowers small, with subparallel perianth segments; perianth segments creamy yellow; lip orange-yellow, often flushed with lilac-red. Lower montane forests 600–1000 m; Bolívar (Auyán-tepui, Perai-tepui), Amazonas (Cerro Marahuaka). Guyana, French Guiana, Ecuador (Amazonian states), northern Brazil. ◆Fig. 408. Maxillaria schlechteri Foldats, Acta Biol. Venez. 31: 403. 1959. —Maxillaria rugosa C. Schweinf., Bot. Mus. Leafl. 20: 24. 1962. Maxillaria rugosa Schltr., Notizbl. Königl. Bot. Gart. Berlin 6: 125. 1914, non Scheidw. 1843. Apparently an epiphyte or a lithophyte, medium-sized, scandent to rhizomatous; pseudobulbs 1-foliate, with leaf-bearing sheaths, remote on rhizome; inflorescences originating from leaf axils; flowers mediumsized, color unknown. Cloud forests on open tepui formations, ca. 2000 m; Bolívar (Roraima-tepui). Guyana, Brazil. Maxillaria schlechteri has not been collected since last century and it may only be a misidentified M. meridensis Lindl. Maxillaria scorpioidea Kraenzl., Kongl. Svenska Vetenskapsakad. Handl. n.s. 46: 71. 1911. Epiphyte, medium-sized, rhizome attached to the substrate only at base; inflorescences shorter than pseudobulb, narrowly ovoid, 2–5 cm long; flowers greenish or yellowish, lip with maroon spots on the margins; dorsal sepal 18–25 × ca. 4 mm. Presumably rain forests. Report based on a cultivated plant known with certainty to have come from the Venezuelan Guayana, probably from Bolívar state. Mexico, western El Salvador, Costa Rica, western Panama, Colombia, Ecuador, Peru, Brazil. Maxillaria setigera Lindl., Edwards’s Bot. Reg. 31: misc. 30. 1845. Epiphyte, medium-sized to rather large,

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cespitose, sun-loving; pseudobulbs compressed, without leaf-bearing sheaths; inflorescences longer than pseudobulbs; flowers very large, 10–15 cm in diameter with widely spreading sepals and petals subparallel to column; tepals cream or white with yellow tips; lip white with a yellow central lobe and purple stripes on disk and lateral lobes. Rain forests or shrubby formations, 50–600 m; Bolívar (Río Aonda on Auyán-tepui), Amazonas (basin of Río Casiquiare, Río Sipapo, Río Ventuari). Barinas, Táchira; Colombia, Guyana, Peru, Brazil. ◆Fig. 413. Maxillaria splendens Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 38. 1836. Maxillaria buchtienii Schltr., Repert. Spec. Nov. Regni Veg. 27: 86. 1929. Epiphyte or lithophyte, sun-loving, medium-sized, cespitose, erect; pseudobulbs 1foliate, with non-leaf-bearing sheaths; inflorescences shorter than leaves, several simultaneously; flowers rather large, with subparallel perianth segments, white with some yellow or orange on lip. Rain or cloud forests or in scrub, (200)–500–1300 m; Bolívar (upper Río Caroní and upper Río Caura basins), Amazonas (Cerro Autana, Cerro Marahuaka, Cerro Sipapo, Río Autana, Río Cataniapo, Río Casiquiare basin). Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia. Maxillaria squamata Barb. Rodr., Gen. Sp. Orchid. 1: 118. 1877. —Ornithidium squamatum (Barb. Rodr.) Barb. Rodr., Gen. Sp. Orchid. 2: 209. 1882. —Camaridium squamatum (Barb. Rodr.) Hoehne, Arq. Bot. Estado Sao Paulo 2: 72. 1947. Maxillaria uncata auct. non Lindl. 1837: sensu Foldats in Lasser, Fl. Venez. 15(4): 542. 1970, pro parte; Dunst. & Garay, Venez. Orchid. Ill. 2: 224. 1961, pro parte. Epiphyte very similar to Maxillaria uncata but with longer rhizomes; usually pendulous to arching; leaves 5–9 cm long; flowers white flushed with pale maroon, particularly on the veins, lip 12–14 mm long. Lowland forests, 100–500 m; Bolívar (Río Icabaru), Amazonas (widespread). Amazonian Colombia, Ecuador, Peru, and Brazil. The limits between this species and

Maxillaria uncata are somewhat diffuse, but there seems to be two entities, one with shorter stems (rhizomes) and leaves, narrow lip base, and rounded apex to the lip (M. uncata) and the other with longer, laxer stems, longer leaves, broader lip base, and retuse or 2-lobed lip apex (M. squamata). Maxillaria uncata has wide distribution, possibly ranging into Central America while M. squamata is restricted to the Amazonian lowlands. Maxillaria stenophylla Rchb. f., Bonplandia (Hanover) 2: 17. 1854. Epiphyte, small to medium-sized, erect to pendulous, anchored over substrate only basally, with a sheath-enveloped rhizome and bearing 2-foliate pseudobulbs at intervals; flowers small, campanulate, originating from the new rhizome sheath axils; tepals white or clear yellow and a basally orangebrown or red-spotted lip; ventral face of the column red-spotted. Cloud forests, ca. 1900 m; Bolívar (Macizo del Chimantá [Apacarátepui]). Aragua, Distrito Federal, Mérida, Miranda); Colombia, Guyana, Suriname, Ecuador. Maxillaria stenophylla occurs at much lower elevations in Guyana, Suriname, and French Guiana. Maxillaria superflua Rchb. f., Cat. Orch.Samml. Schiller 45. 1857. Dicrypta longifolia Barb. Rodr., Gen. Spec. Orchid. 1: 125. 1877. —Maxillaria longifolia (Barb. Rodr.) Cogn. in Mart., Fl. Bras. 3(6): 33, t. 11. 1904, non Lindl. 1832. —Maxillaria tarumaensis Hoehne, Arq. Bot. Estado São Paulo n.s. formato maior 2: 73. 1947. Epiphyte, small to medium-large, cespitose, erect to pendulous; pseudobulbs relatively small with 2–4 leaf-bearing sheaths; leaves fleshy, straight to arched; inflorescences short, erect to horizontal; flowers small to medium-sized with fleshy, yellow or orange perianth segments; lip yellow, orange or deep maroon, purple with a ± well-developed tomentose callus. Moist lowland to lower montane forests, 50–400(–800) m; Bolívar (Altiplanicie de Nuria, Río Caroní, Río Caura, Río Cuyuní), Amazonas (Río Casiquiare, Río Cataniapo, Río Negro, Río

Maxillaria 445

Sipapo, Río Ventuari, Yavita to Maroa road). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. Maxillaria superflua is an amazingly variable species and may contain more than one taxa, but the variation range of each character overlaps. The waxy callus is often lacking or incomplete in herbarium material. Maxillaria tenuis C. Schweinf., Bot. Mus. Leafl. 11: 289. 1945. Epiphyte to 15 cm tall, cespitose to subrepent, erect; pseudobulbs 1-foliate, almost entirely enclosed by verruculosepustulose sheaths of which the innermost 1 or 2 may have leaf blades; inflorescences relatively short; flowers medium-sized for the genus but large for the plant, with subparallel perianth segments; tepals white; lip white with a yellow callus. Moist lowland forests, 50–200 m; Amazonas (Maroa, near San Carlos de Río Negro). Amazonian parts of Colombia, Peru, and Brazil. ◆Fig. 407. Maxillaria uncata Lindl., Edwards’s Bot. Reg. 23: sub t. 1986. 1837. —Camaridium uncatum (Lindl.) Hoehne, Arq. Bot. Estado São Paulo n.s. formato maior 6: 127. 1952. Epiphyte or rarely a lithophyte, minute to medium-sized, cespitose to rhizomatous, erect to pendulous; rhizome clothed with scarious sheaths that envelope the pseudobulbs; pseudobulbs minute, 1-foliate; leaves subterete or very fleshy, involute-conduplicate; flowers originating from the rhizome sheaths, small but medium-sized for the plant; perianth segments white or creamy white, sometimes with purple or brown veins. Lowland to lower montane forests 50–600(–1200) m; widespread species in the flora area, somewhat more common in Amazonas. Miranda; possibly Mexico and Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. Populations from Delta Amacuro and northeastern Bolívar consist of smaller, more dense, thinner-leaved plants with smaller flowers and a somewhat differently shaped lip, and match the typical Maxillaria uncata, while the Amazonian form belongs to M. squamata Barb. Rodr. The Maxillaria uncata complex is much in need of a revision and no

heterotypic synonyms are cited here until this is carried out. Maxillaria villosa (Barb. Rodr.) Cogn. in Mart., Fl. Bras. 3(6): 34, t. 12. 1904. —Dicrypta villosa Barb. Rodr., Gen. Spec. Orchid. 1: 125. 1877. Epiphyte, medium-sized, shade-loving; flowers small to medium-sized, not opening widely; perianth segments yellow or greenish yellow; lip yellow or orange with brownorange or purple specks. Rain forests, 100– 600(–700) m; Bolívar (Sierra de Lema), Amazonas (Brazo Casiquiare, Río Negro, Yavita to Maroa road). Colombia, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 400. Maxillaria villosa is often confused with M. discolor, but M. villosa has longer and narrower leaves, relatively wider petals, and conspicuously 3-lobed lip. The lower surface of the leaves of M. villosa is never purplishtinged as is usually the case in M. discolor. Maxillaria violaceopunctata Rchb. f., Bonplandia (Hanover) 3: 216. 1855. Epiphyte, medium-sized to medium-large, cespitose, erect, usually shade-loving; pseudobulbs 1-foliate, with 2–4 leaf-bearing sheaths; inflorescences short; flowers medium-large for the genus, yellow-cream with a purple-, rose-, or maroon-purple speckled lip. Rain forests, 50–300 m; Delta Amacuro (basin of Río Amacuro, Río Cuyubini), Bolívar (Canaracuni, Kavanayén, lower Río Caura basin, Serranía de Imataca), Amazonas (Río Cataniapo, Río Siapa, Río Sipapo, Yavita to Maroa road). Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Maxillaria xylobiiflora Schltr., Repert. Spec. Nov. Regni Veg. 27: 76. 1929. Epiphyte, minute to small, cespitose; pseudobulbs with 2 or 3 leaf-bearing sheaths; inflorescences erect to horizontal, longer than pseudobulb; flowers small but medium-sized for the plant, with not widely opening, yellow to greenish perianth segments; lip purple-striped or wholly purple in the basal 1/2. Rain forests, 100–1000 m; Bolívar (basin of Río Caroní and Río Paragua), Amazonas (Río Arari, Río Sipapo basin, Sierra de la Neblina). Táchira; Suriname, Ecuador, Peru, Bolivia. ◆Fig. 412.

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Fig. 390. Maxillaria colemanii

1 cm 2 cm

1 cm

Fig. 391. Maxillaria alpestris

Fig. 392. Maxillaria kegelii

Maxillaria 447

Fig. 393. Maxillaria discolor

Fig. 394. Maxillaria aurea

Fig. 395. Maxillaria foldatsiana

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Fig. 396. Maxillaria bolivarensis

Fig. 397. Maxillaria loretoensis

Fig. 398. Maxillaria parkeri

Maxillaria

Fig. 399. Maxillaria brachybulbon

Fig. 400. Maxillaria villosa

Fig. 401. Maxillaria porrecta

449

450

O RCHIDACEAE

Fig. 402. Maxillaria luteoalba

Fig. 403. Maxillaria connellii

Maxillaria 451

Fig. 404. Maxillaria guareimensis

Fig. 405. Maxillaria desvauxiana

Fig. 406. Maxillaria grobyoides

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O RCHIDACEAE

Fig. 407. Maxillaria tenuis

Fig. 408. Maxillaria santanae

Fig. 409. Maxillaria guadalupensis

Maxillaria 453

Fig. 410. Maxillaria quelchii (2 variations)

Fig. 411. Maxillaria nuriensis

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O RCHIDACEAE

Fig. 412. Maxillaria xylobiiflora

Fig. 413. Maxillaria setigera

Mesadenella 455

88. MESADENELLA Pabst & Garay, Arq. Inst. Biol. Veg. 12: 207. 1953. [Subtribe Spiranthinae]. Spiranthes Rich., De Orchid. Eur. 20, 28, 36. Aug.–Sept. 1817, nom. cons., pro parte. by Germán Carnevali, Ivón M. Ramírez-Morillo, and Carlos A. Vargas Humicolous or terrestrial herbs, shade-loving. Roots fasciculate, fleshy, spindleshaped. Leaves convolute, subfleshy, basal, rosulate, petiolate or basally cuneate, present during anthesis. Inflorescences erect, sheathed, terminated by a many-flowered, ± spirally or twisted spike. Flowers small for the subtribe, resupinate, not opening widely; ovary sessile, slightly twisted. Sepals free, dissimilar, subparallel, somewhat diverging toward apex; lateral sepals obliquely decurrent on column foot forming an obtuse mentum; petals connivent with dorsal sepal, forming a galea, inner margins joined with dorsal sepal but apices free. Lip with a distinct claw, with 2 glands at its base; blade conduplicate, simple, constricted about middle forming a basal, oblong to elliptic blade and a broader, elliptic to broadly elliptic, acute to rounded apical blade. Column short, the front pubescent, basally extended into a decurrent incurved foot, together with lateral sepals forming a distinct mentum; anther dorsal, entire, concave, ovate-elliptic, acute to subacuminate; pollinia 4, soft, puberulent, clavate, with a small, roundish viscidium; rostellum rigid, ± cartilaginous, linear-subulate, acuminate; stigmas 2, ventral, approximate, touching each other in middle. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina; 6 species, 1 in Venezuela. Mesadenella angustisegmenta Garay, Bot. Mus. Leafl. 28: 285. 1982. Terrestrial, medium-sized, erect; inflorescences many-flowered, erect; plant autogamous; perianth segments and lip greenish. Montane forests, ca. 800–900 m; Bolívar (Río Yuruaní basin). Zulia; Ecuador. ◆Fig. 414.

Fig. 414. Mesadenella angustisegmenta

456

O RCHIDACEAE

89. MILTONIOPSIS God.-Leb., Orchidophile 9: 63. 1889. [Subtribe Oncidiinae]. Miltonia Lindl., Edwards’s Bot. Reg. 23: sub t. 1976. 1837, pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, cespitose, erect, showy, mostly shade-loving, Roots many, thin; rhizomes short. Pseudobulbs heteroblastic (one internode), 1-leaved apically, strongly laterally compressed (ancipitous), mostly oblong, sometimes ovoid to ± orbicular, enveloped by 1–several imbricate sheaths, the 1 or 2 innermost of which may have foliar blades. Leaves conduplicate, articulate, erect, mostly oblong or linear-oblong, acute, sessile to nearly so. Inflorescences axillary from the sheaths, covering the pseudobulb, 1 or 2 simultaneously, 1-flowered or a several-flowered raceme, erect or nodding; peduncle and rachis terete, remotely sheathed. Flowers large and showy, resupinate, long-lasting, with widely spreading perianth segments, mostly white, pink, or purple, usually with yellow areas on the disk; floral bracts inconspicuous; pedicellate ovary elongate, terete. Sepals free, ± equal, the laterals ± oblique, mostly elliptic or ovate-elliptic, acuminate to obtuse; petals ± equal sepals in length, often broader. Lip flat or ± convex, mostly broadly obovate, longer and broader than the other perianth segments, rigidly affixed to the ventral face of the column through an elevated, keel-like ridge, basally auriculate, without a nectariferous base; disk with several longitudinal keels. Column short, erect, footless, wingless, keeled in the ventral face; anther incumbent, operculate, 1-locular or imperfectly 2-locular; pollinia 2, waxy, ovoid, tegula elongate, linear, viscidium oblong; clinandrium inconspicuous; rostellum transverse, bifid; stigma ventral, large, subquadrate. Capsules ellipsoid. Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, most diverse in the Western slopes of the Andes; 4 species, 1 in Venezuela. Miltoniopsis santanaei Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 276. 1976. Epiphyte 20–30 cm tall, vegetative parts all pale green; flowers showy; sepals and petals white; lip ca. 25 × 25 mm, white with a wide yellow patch in the basal 1/3 and 5 shallowly raised purple keels on the disk; column white. Lower montane forests, ca. 600 m; Bolívar (Altiplanicie de Nuria). Endemic. ◆Fig. 415.

Fig. 415. Miltoniopsis santanaei

Mormodes 457

90. MORMODES Lindl., Intr. Bot. ed. 2, 446. 1835. [Subtribe Catasetinae]. Cyclosia Klotzsch, Allg. Gartenzeitung 6: 305. 1838. by Gustavo A. Romero-González Epiphytes or rarely terrestrial herbs. Stems modified into pseudobulbs of several internodes, slender-ovoid or fusiform, when young entirely concealed by distichous, scarious sheaths. Leaves several, distichous, convolute, plicate, articulate, deciduous at the end of the growing season. Inflorescences 1–several, lateral, arising from the middle internodes of the pseudobulbs, erect, arching or pendent, racemose, few- to many-flowered. Flowers resupinate or not, functionally staminate or bisexual, protandric, sometimes polymorphic, fragrant, often showy; floral bracts lanceolate, funnel-shaped. Sepals and petals fleshy, the lateral sepals strongly reflexed or not; lip fleshy-thickened, clawed, resupinate, entire or 3-lobed, glabrous or variously pubescent, the margins generally strongly reflexed. Column semiterete, elongate, twisted to one side; anther ventral, operculate, imperfectly bilocular to unilocular, fleshy, apiculate; clinandrium elongate, long-acuminate; pollinarium with 2 yellow pollinia, ovoid, cartilaginous, with a tubular tegula, and a large, adhesive, circular or elliptic viscidium. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 50–55 species, 8 in Venezuela, 4 of these in the flora area. Some Mormodes species can produce staminate or usually larger, bisexual, protandric flowers (acting as pollen donors first, then as pollen receptors). Staminate flowers wilt soon after pollinarium removal; in protandric flowers the column straightens up and moves away from the lip after pollinarium removal, and the flowers last many days thereafter. Furthermore, there is evidence of variation in the morphology and pubescence of the lip from at least one Central American species, even in flowers borne in a single inflorescence. Several species display wide variation in color. It is not known whether species throughout the genus can exhibit this wide range of variation or whether this variation is due to sexual dimorphism. Flowers of this genus are pollinated by euglossine bees. Key to the Species of Mormodes 1. 1. 2(1). 2. 3(1). 3. 4(3). 4.

Lip apex, when flattened, repand or shallow to deeply 3-lobed ............... 2 Lip apex entire, not repand or conspicuously 3-lobed .............................. 3 Lip broader than long, apex truncate, obscurely 3-lobed, midlobe apiculate ............................................................................ M. carnevaliana Lip as broad as long, apex repand, acute, midlobe abruptly acuminate ............................................................................................. M. vernixioidea Lip broadly ovate ...................................................................... M. cucumerina Lip transversely elliptic ............................................................................. 4 Lip glabrous ........................................................................... M. carnevaliana Lip densely and shortly pilose ................................................ M. orinocoensis

Mormodes carnevaliana Salazar & G.A. Romero, Lindleyana 9: 255. 1994. Epiphyte; inflorescence arching; flowers red, maroon, orange, or light brown. Wet forests, 50–1200 m; Delta Amacuro (Serranía

de Imataca), Bolívar (Los Pijiguaos, Macizo del Chimantá [Acopán-tepui], Paragua, Río Caranacuni, Río Sotebe), Amazonas (Caño Iguana, Río Cataniapo, Sierra Pakaraima). Brazil (Roraima). ◆Fig. 416.

458

O RCHIDACEAE

Mormodes cucumerina Pabst, Bradea 1: 167. 1972. Epiphyte; inflorescence arching; flower light brown. Evergreen lowland forests, 100– 200 m; Amazonas (San Carlos de Río Negro). Brazil (Amazonas). ◆Fig. 418. Mormodes orinocoensis Salazar & G.A. Romero, Lindleyana 9: 259. 1994. Epiphyte; inflorescence arching; sepals and petals brownish to reddish green, lip pink spotted with lavender, shortly pilose. Evergreen lowland forests, 100–600 m; Bolívar (Los Pijiguaos), Amazonas (San Fernando de Atabapo). Colombia. Mormodes vernixioidea Pabst, Bradea 2: 56. 1975. Colombia, Venezuela, Brazil (Amazonas); 2 subspecies, 1 in Venezuela, restricted to the flora area. M. vernixioidea subsp. autanensis G.A. Romero & Salazar, Lindleyana 9: 260. 1994. Epiphyte; inflorescence arching; flowers brown spotted with maroon-red or sepals and petals red or orange-red with brownish purple veins and lip of a homogeneous red, orange, or white. Evergreen lowland forests, 100–200 m; Amazonas (Cerro Sipapo, Río Autana, Río Cataniapo, San Fernando de Atabapo). Endemic. ◆Fig. 417.

Fig. 417. Fig. 416. Mormodes vernixioidea Mormodes carnevaliana subsp. autanensis

Fig. 418. Mormodes cucumerina

Myoxanthus

459

91. MYOXANTHUS Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 50, t. 88. 1835. [Subtribe Pleurothallidinae]. Duboisia H. Karst., Allg. Gartenzeitung 15: 394. 1847, non R. Br. 1810. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, sometimes lithophytes or subterrestrial herbs, inconspicuous to rather large and showy, 5–50(–150) cm tall, erect to subpendulous, cespitose to longcreeping, sometimes suffrutescent, shade- to sun-loving. Rhizomes very short to long-creeping, appressed to substrate or rarely ascendent, unbranched or branching, mostly covered with hispidulous sheaths. Stems subterete, 3–50 cm tall, apically 1-foliate, wiry to stout, sometimes distinctly swollen at the base, often apically proliferous, enveloped by large, papery, usually hispidulous sheaths that are often easily shed. Leaves conduplicate, thin to rigidly coriaceous, linear to broadly elliptic or oblong. Inflorescences originating without an annulus from or near the apex of stem and then 1-flowered, successive or simultaneous, sometimes in large numbers in a fascicle; in rare cases the inflorescences originating low on the stem or from the rhizome, and then often short, successively flowered racemes. Flowers resupinate, inconspicuous to relatively showy, 5–30 mm diameter, campanulate to widely spreading, floral bracts inconspicuous, often hispidulous; pedicel and ovary smooth or hispidulous to spiny. Perianth segments often fleshy, mostly variously pubescent; sepals free or the laterals variously connate into a synsepal, often fleshier and broader than the petals; petals similar to sepals, to longer and thinner or much shorter, often variously thickened, the apices sometimes developed into scent-producing glands. Lip articulate to column foot, unlobed, 3- or 5-lobed, usually much smaller than the other perianth segments, often convex and recurved, variously callose or ecallose. Column well developed, apically variously toothed or winged, basally produced into a well-developed column foot; anther ventral to subterminal, incumbent, operculate, sometimes the surface covered with fine points; clinandrium entire to variously dentate or lacerate; pollinia 2, attached to a small viscidium; rostellum transverse; stigmatic surface ventral. Capsules mostly hispidulous to spiny. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, southern and Amazonian Brazil, Bolivia; ca. 40 species, 10–12 species in Venezuela, 5 of these in the flora area. Most species of Myoxanthus are easily recognized by the pubescent to hispidulous sheaths on the stems, but this character is not exclusive to Myoxanthus. It is also found in other genera of the Pleurothallidinae. Most Myoxanthus species also have petals that are dissimilar to the sepals, have an apical osmorphore, or are elongate or caudate. Key to the Species of Myoxanthus 1. 1. 2(1).

Rhizome shortly creeping to subscandent; plants small with stems 2.2– 7 cm long ......................................................................... M. trachychlamys Rhizome very short to almost absent, plants cespitose; stems mediumsized to rather large with stems 5–35 cm long ..................................... 2 Petals apically developed into a rounded, well-developed purple or purple-brown osmophore (scent-producing gland); lateral sepals connate for > 1/2 their length ....................................................... M. reymondii

460

2.

3(2).

3.

4(3). 4.

O RCHIDACEAE

Petals attenuated in apical 1/2, never developed into a conspicuous, rounded, purple or purple-brown osmophore; lateral sepals free or connate for < 1/2 their length ....................................................................... 3 Flowers relatively large with dorsal sepal ca. 11 mm long; perianth segments bright reddish yellow with strong longitudinal, narrow, redpurple stripes ........................................................................... M. speciosus Flowers small with dorsal sepal 3–8 mm long; perianth segments white, yellow, or greenish, sometimes purple-tinged or dorsal sepal with a longitudinal purple stripe ........................................................................... 4 Stems apically proliferous; dorsal sepal 7–8 mm long; lip ± unlobed, 1.8– 2 mm wide; lateral sepals abruptly short-acuminate ......... M. parvilabius Stems not apically proliferous; dorsal sepal 3–5 mm long; lip 3-lobed near or below its middle, 1–1.3 mm wide; lateral sepals acute to subacute .......................................................................................... M. simplicicaulis

Myoxanthus parvilabius (C. Schweinf.) Luer, Selbyana 7: 49. 1982. —Pleurothallis parvilabia C. Schweinf., Bot. Mus. Leafl. 20: 14. 1962. Epiphyte 20–100 cm tall; flowers deep purple or translucent and tinged with purple. Cloud forests, 700–800 m; Amazonas (Sierra de la Neblina). Guyana, Suriname, Ecuador, Brazil. ◆Fig. 419. Myoxanthus reymondii (H. Karst.) Luer, Selbyana 7: 49. 1982. —Duboisia reymondii H. Karst., Allg. Gartenzeitung 15: 394. 1847. —Pleurothallis reymondii (H. Karst.) Rchb. f. in Walp., Ann. Bot. Syst. 3: 520. 1852. Epiphyte, rarely a lithophyte; leaves 8–20 cm long; sepals spreading, externally purplebrown, internally pale or dark honey brown; petals parallel to column, basally pale yellow-brown, apically with dark purple-brown scent-producing glands, lip yellow with dark purple-brown patches. Presumably cloud forests, (unspecified elevation); Bolívar (unspecified locality). Aragua, Distrito Federal, Falcón, Lara, Miranda, Mérida, Trujillo, Táchira, Yaracuy; Colombia, Ecuador. Myoxanthus simplicicaulis (C. Schweinf.) Luer, Selbyana 7: 51. 1982. —Pleurothallis scandens var. simplicicaulis C. Schweinf., Bot. Mus. Leafl. 20: 16. 1962. —Pleurothallis simplicicaulis (C. Schweinf.) Luer, Selbyana 3: 388. 1977. Epiphyte; stems 7–25 cm long; leaves 6.5– 14 cm long; flowers campanulate; perianth segments yellowish or whitish, often the dor-

sal sepal with a purple longitudinal stripe; dorsal sepal 3–5 mm long. Locally common in cloud forests, (800–)1100–1900 m; Bolívar (Auyán-tepui, Jaua-Sarisariñama massif, Macizo del Chimantá, upper Río Cuyuní basin, Sororopán-tepui), Amazonas (Río Iguapo, Sierra de la Neblina). Guyana. Myoxanthus speciosus (Luer) Luer, Selbyana 7(1): 51. 1982. —Pleurothallis speciosus Luer, Selbyana 3: 392, fig. 296. 1977. Pleurothallis exasperata auct. non Lindl. 1859: sensu Foldats in Lasser, Fl. Venez. 15(2): 269. 1970. Epiphyte or lithophyte; stems 20–35 cm long; leaves 11–16 cm long; flowers relatively showy for the genus; perianth segments bright yellow or yellow-purple with dark brown-purple longitudinal stripes; dorsal sepal 11–12 mm long. Cloud forests or shrublands on tepui summits, 2200–2600 m; Bolívar (Macizo del Chimantá [Apacarátepui]), Amazonas (Cerro Marahuaka). Distrito Federal; Costa Rica, Panama, Colombia. ◆Fig. 420. Myoxanthus trachychlamys (Schltr.) Luer, Selbyana 7: 51. 1982. —Pleurothallis trachyclamys Schltr., Repert. Spec. Nov. Regni Veg. 17: 23. 1922. Epiphyte; stems 2–7 cm long; leaves 4–7.5 cm long; dorsal sepal 2.7–5 mm long, whitish or yellowish. Rain forests, 400–600 m; Bolívar (Río Canarakuni, Río Churún near Guarimba), Amazonas (Río Putaco, Sierra Parima). Costa Rica, Panama, Ecuador, Peru, Brazil.

Myoxanthus 461

Fig. 419. Myoxanthus parvilabius

Fig. 420. Myoxanthus speciosus

462

O RCHIDACEAE

92. NIDEMA Britton & Millsp., Bahama Fl. 94. 1920. [Subtribe Laeliinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes or lithophytes, small to medium-sized. Rhizomes well developed, creeping; pseudobulbs erect, stipitate, basally enveloped with imbricating sheaths, apically 1-foliate. Leaves rather thin but rigid, oblong or linear, erect. Inflorescence terminal, racemose, emerging from a small spathe, conspicuously bracteate, fewflowered. Flowers small to medium-small; floral bracts conspicuous; pedicellate ovary subterete. Perianth segments narrowly elliptic, acute. Sepals free, parallel or spreading; petals ± parallel to column. Lip articulated with the base of the column, thick, unlobed, ecallose, longitudinally canaliculate, dorsally keeled at apex. Column arched, exauriculate, somewhat triquetrous apically; clinandrium simple; anther apical, operculate, incumbent, 2-locular, each locule longitudinally septate and thus anther appearing 4-locular; pollinia 4, laterally compressed, with short caudicles; rostellum transverse; stigma ventral. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil; 2 species, 1 in Venezuela. This genus is apparently related to the Scaphyglottis complex and is not near Epidendrum L., a genus with which it has been synonimized in many floristic treatments. Nidema ottonis (Rchb. f.) Britton & Millsp., Bahama Fl. 94. 1920. —Epidendrum ottonis Rchb. f., Hamburger GartenBlumenzeitung 14: 213. 1858. Epiphyte, small to medium-sized, creeping to ± cespitose; pseudobulbs apically 1foliate; flowers small, white, creamy, or yellowish, perianth segments not widely opening. Rain forests or rarely tepui slopes, 50– 500(–1000) m; Bolívar (Altiplanicie de Nuria, basins of Río Caroní, Río Caura, and Río Cuyuní), Amazonas (upper Río Orinoco, Río Pavone). Aragua, Miranda, Monagas, Portuguesa, Táchira, Yaracuy; Nicaragua, Panama, West Indies, Colombia, Ecuador, Peru, Brazil. ◆Fig. 421.

Fig. 421. Nidema ottonis

Notylia 463

93. NOTYLIA Lindl., Bot. Reg. 11: sub t. 930. 1825. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, sometimes on twigs, cespitose. Rhizomes short. Stems very short, usually developed into pseudobulbs. Pseudobulbs when present, clustered, small, 1foliate, enveloped with distichous, imbricate or equitant sheaths, at least the inner with foliar limbs. Leaves conduplicate, articulate, coriaceous or fleshy; erect and spreading. Inflorescences lateral, basal, usually arched or recurved, racemose or rarely with 1 or 2 basal branches, few- to many-flowered, originating from the base of pseudobulbs or from the axils of the sheaths. Flowers small, resupinate, fleshy or subfleshy, greenish, yellowish, or whitish; floral bracts generally narrow; pedicellate ovary terete. Sepals similar, mostly narrow, stout, and spreading, free or the laterals variously connate, ± oblique, often revolute; dorsal sepal free, lanceolate, strongly concave; lateral sepals subequal, ± oblique; petals similar to the sepals, generally smaller, oblique. Lip unlobed or slightly lobed, commonly clawed, footless, blade unlobed, usually flat or concave, usually triangular to ovate or cordate, obtuse to acuminate; disk smooth or with an elevated median keel. Column stout or usually slender, subterete, wingless, footless, apex recurved or geniculate; anther dorsal, imperfectly 2-locular, deeply concave; pollinia 2, cartilaginous, tegula very elongate, linear-spatulate, viscidium small; rostellum elongate, acuminate; stigma apical or subventral. Capsule ellipsoid or subspherical. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 40 species, 21 in Venezuela, 11 of these in the flora area. The species of Notylia are poorly known, and careful monographic work will be required to ascertain the actual number of taxa, their delimitations, and distributions. Key to the Species of Notylia 1. 1. 2(1).

2.

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

Adult leaves ca. 3 cm long .......................................................................... 2 Adult leaves > 3.5 cm long ......................................................................... 3 Lateral sepal connate about or above its middle, subparallel to slightly divergent apically; claw of lip glabrous; lip blade acute to acuminate when flattened ..................................................................... N. angustifolia Lateral sepal connate only below its middle, obviously divergent apically; claw of lip papillose; lip blade apically truncate to rounded when flattened .............................................................................. N. longispicata Leaves > 9 times longer than wide ............................................................ 4 Leaves 4–5 times longer than wide ........................................................... 5 Claw of the lip about as long as the blade or only slightly shorter; blade at least 2 times as long as wide .................................................. N. aromatica Claw of the lip 1/3–1/2 as long as blade; blade about as long as wide or slightly longer ......................................................................... N. peruviana Column densely papillose ........................................................... N. pentachne Column glabrous or glabrescent ................................................................ 6 Claw of lip 1/5–1/4 as long as blade .............................................................. 7

464

O RCHIDACEAE

Claw of lip 1/3–1/2 as long as blade to ± equaling it .................................... 9 Sepals 8–12 mm long; petals 4–5 times longer than wide; blade of the lip abruptly broad-acuminate .................................................. N. rhombilabia 7. Sepals ca. 5 mm long; petals ca. 3 times longer than wide; blade of lip obtuse to acute ........................................................................................... 8 8(7). Claw of lip with a well-defined longitudinal keel; lateral sepals fused to 1/2 their length .................................................................................. N. laxa 8. Claw of lip rounded, without a well-defined longitudinal keel; lateral sepals fused to 4/5 their length .............................................. N. platyglossa 9(6). Lip claw papillose ................................................................. N. yauaperyensis 9. Lip claw glabrous ..................................................................................... 10 10(9). Synsepal bifid; blade of lip longer than its claw ....................... N. microchila 10. Synsepal entire; blade of lip shorter than claw, basally conduplicate, about as long as wide ........................................................................... 11 11(10). Plants with conspicuous pseudobulbs, at least 1.5 cm long; leaves 3–4 times longer than wide; claw of lip ca. 1/3–1/2 as long as blade ........ N. aromatica 11. Plants with small pseudobulbs totally enveloped by leaf sheaths, < 1 cm long; leaves at least 5 times longer than wide; claw of lip ca. 3/4 as long as to ± equaling the blade ......................................................... N. fragrans 6. 7(6).

Notylia angustifolia Cogn. in Urb., Symb. Antill. 6: 618. 1910. Notylia nana Cogn. in Urb., Symb. Antill. 6: 619. 1910. Twig epiphyte, minute, erect to pendulous, cespitose; flowers very small, few in a short, nodding inflorescence; sepals and petals green; lip white. Rain forests, 100–700 m; Bolívar (Río Caroní basin), Amazonas (basin of Río Manapiare and Río Ventuari). Trinidad-Tobago, northern Brazil. Notylia aromatica Barker & Lindl., Edwards’s Bot. Reg. 27: misc. 40. 1841. Twig epiphyte; sepals and petals green and lip white, or sepals, petals, and lip yellow and woolly. Rain forests, always as a twig epiphyte, 50–1200 m; Bolívar (Río Caroní basin), Amazonas (Cerro Marahuaka, Río Cataniapo). Mexico, Central America, Amazonian Brazil. Notylia fragrans H. Focke, Bot. Zeitung (Berlin) 11: 342. 1853. Notylia venezuelana Schltr., Repert. Spec. Nov. Regni Veg. 6: 45. 1919. Twig epiphyte; sepals green; petals and lip white. Rain forests, 100–500 m; Amazonas (Río Cataniapo basin). Venezuelan Coastal Cordillera, Táchira, Trujillo; Guyana, Suriname, French Guiana.

Notylia laxa Rchb. f., Gard. Chron. n.s. 16: 620. 1881. Epiphyte; leaves 10–12 × 2–3 cm; sepals and petals pale greenish yellow, lip shiny, yellowish green; dorsal sepal ca. 3.5 mm long. Rain forests, 500–700 m; Bolívar (upper Río Churún gorge near Guarimba). Barinas; Ecuador, Amazonian Brazil, Bolivia, possibly Peru. The flora area plants here referred to Notylia laxa have narrower leaves than is usual for the species, but more material is required to ascertain their taxonomic status. Notylia longispicata Hoehne & Schltr., Arq. Bot. São Paulo 1: 280, pl. 22. 1926. Notylia trullifera Brade, Rodriguésia 20: 43. 1943. Twig epiphyte; inflorescence few-flowered; flowers white. Rain forests, 50–200 m; Amazonas (basin of Río Atabapo, Río Casiquiare, and Río Cataniapo). Amazonian Brazil. Notylia microchila Cogn. in Mart., Fl. Bras. 3(6): 124. 1904. Notylia amesii L.B. Sm. & S.K. Harris, Contr. Gray Herb. 97: 40, figs. H–K. 1937. Epiphyte; sepals yellow-green; petals whitish or green; lip greenish or whitish, gla-

Octomeria 465

brous; column stout, glabrous or minutely pubescent. Rain forests, 100–700 m; Bolívar (Cerro Guaiquinima, upper basins of Río Caroní and Río Paragua), Amazonas (Caño Yureba, Río Mawarinuma). Ecuador, Peru, Amazonian Brazil. Notylia pentachne Rchb. f., Bonplandia (Hanover) 2: 90. 1854. Usually a twig epiphyte, sometimes growing or thick branches, rarely subterrestrial; flowers showy; sepals pale or yellowish green; petals white or very pale green with 2 or 3 orange spots; lip white; column green. Rain forests, 700–800 m; Bolívar (Río Kukenán). Aragua, Carabobo, Distrito Federal, Falcón, Miranda, Yaracuy, Zulia; Panama, Colombia. Notylia peruviana (Schltr.) C. Schweinf., Bot. Mus. Leafl. 12: 205. 1946. —Dipteranthus peruvianus Schltr., Orchis 10: 187, fig. 44. 1916. Twig epiphyte; racemes long, pendulous; sepals and petals greenish yellow or yellow; lip white. Rain forests, 100–700 m; Bolívar (basins of Río Caroní and Río Caura), Amazonas (Río Casiquiare). Ecuador, Peru, Amazonian Brazil, Bolivia. Notylia platyglossa Schltr., Notizbl. Königl. Bot. Gart. Berlin 6: 125. 1914. Epiphyte; sepals and lip very pale green, almost white; petals white with a variable number of tiny to relatively large orange marks. Rain forests, low, open humid associations, 100–800 m; Bolívar (Río Caroní, Río Cuyuní), Amazonas (basin of Río Manapiare, upper Río Orinoco, Río Padamo). Amazonian Brazil.

Notylia rhombilabia C. Schweinf., Bot. Mus. Leafl. 12: 206. 1946. Usually a twig epiphyte, sometimes growing in thicker branches; inflorescences fewto many-flowered; flowers medium-sized to large for the genus; sepals pale green; petals white with 1 or 2 orange spots; lip white. Rain forests, 100–500 m; Bolívar (lower basin of Río Caroní, Río Caura), Amazonas (upper Río Orinoco). Miranda, Monagas, Sucre; Peru, Amazonian and southern Brazil. ◆Fig. 422. Notylia yauaperyensis Barb. Rodr., Vellosia ed. 2, 1: 131. 1891. Twig epiphyte; inflorescences many-flowered; sepals and petals white with 2 or 3 small purple spots on petals or totally pale green. Rain forests, 50–200 m; Bolívar (Altiplanicie de Nuria), Amazonas (basin of Río Cataniapo, upper Río Orinoco, Río Ventuari). Amazonian Brazil.

Fig. 422. Notylia rhombilabia

94. OCTOMERIA R. Br. in W.T. Aiton, Hortus Kew ed. 2, 5: 211. 1813. [Subtribe Pleurothallidinae]. by Germán Carnevali & Ivón M. Ramírez-Morillo Lithophytes, terrestrial herbs, or epiphytes, minute to small, erect to pendulous, rhizomatous to cespitose. Rhizome very short to long-creeping, with ramicauls at long internodes. Ramicauls almost absent, to more commonly well developed, shorter to much longer than leaves, apically 1-foliate, terete or flattened, with 1– many internodes loosely to densely sheathed. Leaves conduplicate to terete, biconvex to hemiterete, thinly coriaceous to thickly fleshy, sessile to subpetiolate, usually oblong-elliptic or elliptic, often narrowly ovate or linear, apex usually tridenticulate.

466

O RCHIDACEAE

Inflorescences fasciculate or solitary, 1-flowered, often emerging from a mass of papery to scarious bracts, without an annulus, sometimes originating new plants. Flowers minute to medium-large, usually yellow, white, purple, or reddish and suffused with other hues, mostly short-leaved; floral bracts minute to small; pedicel articulate with ovary, very short to relatively long. Perianth segments usually membranaceous, rarely fleshy. Sepals usually free, sometimes the lateral ones connate into a short synsepal (1–)3(–5)-veined, subequal; petals ± similar to the sepals, sometimes narrower or wider, 1–3-veined. Lip free from column and hinged to column foot; blade simple to sharply 3-lobed, sometimes basally the margins erect, conduplicate, hence forming a ± 3-lobed lip, sessile to shortly clawed or basally cuneate, apex variable, often emarginate; disk callose to ecallose. Column short to long, straight or arched, wingless, usually subterete, basally produced into a short to long column foot; anther apical to (rarely subventral), incumbent, 4–8-celled, operculate; pollinia 8, exceptionally 6, in 2 sets of 4, clavate, sometimes attached to an ill-defined viscidium; clinandrium usually shallow; stigma narrow, ventral. Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Paraguay, northern Argentina; 150–170 species, 30 or 32 in Venezuela, 28 of these in the flora area. The flowers of the genus are membranous and the subtle details of the lip are difficult to retrieve from rehydrated material. There are probably several additional taxa of the genus in the flora area, particularly in the high tepuis of the Guayana Highlands. There is abundant sterile or difficult-to-determine herbarium material in several relevant herbaria which awaits identification and some may represent additional undescribed or previously unreported taxa. Octomeria has centers of diversity in the Guianas, the Amazon Basin, and southern Brazil. Key to the Species of Octomeria 1.

1.

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

3.

4(2).

4.

Leaves terete or semiterete (the thickness of the leaf about equal to or larger than its width), mostly linear, 0.5–1.5 mm; plants always cespitose ................................................................................................. 2 Leaves conduplicate, thin or very fleshy but with thickness always much smaller than width), linear to broadly elliptic, much wider than 1.8 mm; plants cespitose to creeping ................................................................... 5 Leaves 1/3 or less the length of the stem .................................................... 3 Leaves about as long or longer than the stem .......................................... 4 Stems flattened; leaves ventrally convex in the apical 1/2; dorsal sepal 6–7 mm long; lateral lobes of the lip ca. 1/3 the length of the total lip blade upon flattening ....................................................... O. yauaperyensis Stem terete; leaves at the apical 1/2 flat or somewhat concave; dorsal sepal 3–5 mm long; lateral lobes of the lip, linear-lanceolate, reaching at least 1/2 the length of the total lip blade upon flattening ...... O. scirpoidea Dorsal sepal 3–5 mm long; lip 1.2–1.5 mm long, 3-lobed about its middle; midlobe ovate or subtriangular, apically retuse or emarginate; plants from cloud forests or premontane rain forests, 900–1900 m ..... O. filifolia Dorsal sepal 7–10 mm long; lip 4–5 mm long, 3-lobed in the basal 1/3; midlobe elliptic, apically rounded; plants from tropical rainforests, more rarely from cloud forests, 100–700(–1500) m ............... O. taracuana

Octomeria 467

5(1).

5.

6(5).

6.

7(5). 7.

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

13.

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

Floral peduncle at least half the length of or longer than the subtending leaf; small plants with proportionally short stems and broad leaves; leaves < 1 cm long; flowers large for plant’s size .................................. 6 Floral peduncle much shorter than subtending leaf or almost none; plants variable but usually larger than those of above; leaves > 1.2 cm long; flowers smaller in proportion to plant size (except O. steyermarkii) ................................................................................................................ 7 Stems very short (1–2 mm long); leaves prostrate over substrate and tightly appressed to it, broadly elliptic to suborbicular (confetti-like), green on both faces, much longer than stem; lip with short, retrorse, basal, lateral lobes ................................................................ O. romerorum Stems longer (4–11 mm long); leaves erect or spreading or loosely appressed to the substrate, about as long as the stems, not prostrate over substrate, elliptic, usually purple-tinged on lower surface; lip with porrect, uncinate, lateral lobes borne about midlength on the lip blade .................................................................................................. O. gemmula Mature plants under 5 cm tall; leaves to 2.5 cm long, succulent with thickened cross sections, often purple-tinged ............................................... 8 Mature plants > 5.5 cm tall; leaves > 2.6 cm long, usually coriaceous, more rarely succulent with thickened cross sections, purple-tinged or not .............................................................................................................. 16 Stems remote on rhizome, plants creeping ................................. O. apiculata Stems aggregate on an abbreviated rhizome, plants cespitose ................ 9 Petals 1-veined (or with a partial, faint, second vein) ............................ 10 Petals 3-veined ......................................................................................... 13 Petals and sepals broadly ovate, obtuse at apex, ca. 2 mm long, 1.7 mm wide; lip entire with 1/2 calli on the basal half ............................... O. sp. A Petals and sepals elliptic or ovate, variable at apex, at least 3 mm long; lip 3-lobed or entire, but if entire with an emarginate apex ................... 11 Petals and sepals setose-acuminate; lip entire, apically emarginate ............................................................................................. O. steyermarkii Petals and sepals elliptic to oblong-elliptic, not setose-acuminate, lip sharply 3-lobed, not apically emarginate ............................................ 12 Leaves always much shorter than the stem .............................. O. parvifolia Leaves as long as or longer than the stems ................................... O. parvula Lip ca. 2 times longer than wide, thickened toward apex, surface verruculose; sepals and petals linear-lanceolate; flowers white ....................................................................................................... O. exigua Lip about as long as wide to not more than 1.5 times longer than wide, not thickened toward apex, surface smooth; petals and sepals lanceolate to elliptic; flowers pure yellow, or pink- or purple-tinged ...................... 14 Lip oblong-elliptic, ca. 1.5 times as long as wide, with 2 sharp longitudinal keels or folds, 3-toothed apically, clawed at base .......................... O. minor Lip broadly elliptic to suborbicular, about as long as wide, lacking 2 longitudinal keels or folds, apically emarginate, not clawed at base ......... 15 Lip suborbicular to subquadrangular, apically rounded, emarginate; petals lanceolate; flowers clear yellow, often with some purplish veins or apices, never or rarely cleistogamous; the midvein not impressed on upper surface of leaf (better observed in fresh leaves or good rehy-

468

O RCHIDACEAE

15.

16(7). 16. 17(16). 17. 18(17).

18. 19(17).

19.

20(19).

20.

21(16). 21. 22(21). 22. 23(22).

23.

24(22).

24.

drated material); plants from relatively low elevations at 400– 1400(–1900) m ...................................................................... O. integrilabia Lip subrhombic, apically truncate and very shallowly emarginate; petals elliptic; flowers reddish orange or purplish, rarely yellow, often cleistogamous; midvein on upper surface of leaf sharply impressed (better observed in fresh leaves or good rehydrated material); plants from higher elevations usually above 1400 and more often well above 1700 m ......................................................................................................... O. nana Plants with elongated rhizomes and thus long-creeping, the internodes between stems at least 0.5 cm long ..................................................... 17 Plants cespitose or very shortly creeping ................................................ 21 Mature leaves < 6 cm long ....................................................................... 18 Mature leaves > 8 cm long ....................................................................... 19 Rhizome long-creeping, forming mats around branches or the boles of host trees; lip sharply 3-lobed, ca. 3 mm long; mature leaves 3–6 mm wide .......................................................................................... O. apiculata Rhizome short-creeping, subcespitose; lip somewhat 3-lobed, ca. 5 mm long; mature leaves (8–)12–20 mm wide ............................... O. spathulata Plants short-creeping, subcespitose; leaves on mature (flowering) stems usually much shorter than (1/3–1/2 as long) the stems; petals and sepals > 9 mm; lip lobed at its lowermost 1/4 ........................... O. flaviflora Plants long-creeping, never cespitose; leaves on mature stems longer, about as long or slightly shorter than the stems; petals and sepals < 7 mm long; lip lobed at its lowermost 1/3 .......................................... 20 Flowers with relatively long pedicels that are not totally clothed by sheaths; apical lobe of lip subrhombic, wider at its central portion, apically not 3-lobed or 3-toothed ........................................ O. rhizomatosa Flowers with relatively short pedicels, totally clothed by a series of imbricate sheaths; apical lobe of lip ovate or lanceolate, widest at its base, apically 3-lobed or 3-toothed .................................................. O. erosilabia Leaves elliptic to broadly elliptic, at the most 3 times as long as wide, to 55 × 22 mm ............................................................................. O. spathulata Leaves linear, linear-elliptic, or narrowly elliptic, at least 4 times as long as wide, variable in size ....................................................................... 22 Petals and sepals long-acuminate; lip simple ......................................... 23 Petals and sepals apically rounded; lip obtuse to acute ......................... 24 Petals and sepals setose-acuminate from a narrowly lanceolate base, the tails as long to several times longer than the basal portion; perianth segments widely spreading; plants form eastern Bolívar state at elevations of 1500 m or below ..................................................... O. steyermarkii Petals and sepals long-acuminate from a broadly elliptic base, the tails shorter than to as long as the basal portions; dorsal sepal and petals porrect and subparallel to the lip, the lateral sepals parallel in the lower half, then abruptly reflexed; plants from Amazonas state tepuis or western Bolívar at elevations of 2500 m or above ............... O. anomala Mature leaves under 6 mm wide, usually under 5 mm wide (occasional leaves may reach 7 mm but most leaves in such individual will be within the previous width range) ........................................................ 25 Mature leaves 8 mm or more wide (occasional leaves may be somewhat

Octomeria 469

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

28.

29(24).

29.

30(29).

30.

31(29).

31.

32(31). 32.

33(32). 33.

narrower but most leaves in such an individual will be within the previous width range) ............................................................................... 29 Most mature leaves on a given plant less than 1/2 as long as their stems .............................................................................................................. 26 Most mature leaves on a given plant longer, as long as to 3/4 as long as their stems ........................................................................................... 28 Petals 3-veined; flowers yellow; sepals broadly elliptic to nearly rounded; old mature plants with stems to 20 cm long .......................... O. cordilabia Petals 1-veined; flowers purple or reddish; sepals elliptic to oblongelliptic; old mature stems rarely surpassing 15 cm tall ..................... 27 Lip 3-lobed; sepals 1-veined; petals and sepals similar in width, ca. 3 mm long .......................................................................................... O. parvifolia Lip simple; sepals 3-veined; petals narrower than sepals, ca. 5 mm long ............................................................................................... O. lancipetala Flowers white or very pale yellow, subcampanulate; lip elliptic to spatulate, much longer than wide, verruculose on its upper surface; petals 3-veined; plants from elevations below 1500 m ........................... O. exigua Flowers dark purple, widely opening; lip suborbicular, about as long as wide, smooth on its upper surface; petals 1-veined; plants from elevations above 1800 m ................................................................. O. splendida Large plants with most mature stems usually 15–28 cm long; leaves 1/6–1/3 as long as their stems, foliar blades mostly 9–15 mm wide; flowers yellow; sepals 8–12 mm long ......................................................... 30 Smaller plants with most mature stems under 11 cm long; leaves less than 1/2 as long as their stems or longer, foliar blades 5–11 mm wide; flowers yellow, reddish, or purple; sepals mostly under 7 mm long (to 10 mm in O. surinamensis) ................................................................. 31 Petals broadly elliptic, abruptly acute, 3–3.5 mm wide; leaves elliptic, to 4 times longer than wide, at least 5 times as long as the subtending stem; plants apparently restricted to the Roraima massif ...... O. connellii Petals elliptic, acute, ca. 2 mm wide; leaves oblong-elliptic, > 6 times as long as wide, 3–4 times longer than the subtending stems; plants widespread in the Guayanan tepuis ............................................... O. flaviflora Sepals yellow, 8–11 mm long; lip 3-lobed, the central lobe obovate, much broader toward its apex; plants from lowland rain forests at elevations usually under 500 m ......................................................... O. surinamensis Sepals and petals purple or rarely yellow, white, or cream colored: lip simple or 3-lobed, when 3-lobed the central lobe ovate or oblong; plants from tepuis summits or slopes above 850 m, usually well above 2000 m .............................................................................................................. 32 Lip simple, about as broad as long; flowers purple; sepals and petals elliptic, subacute ..................................................................... O. heleneana Lip sharply 3-lobed, longer than wide; flowers purple, white, yellowish, or cream-colored; sepals and petals lanceolate or narrowly lanceolate, acute or acuminate .............................................................................. 33 Central lobe of lip ovate or broadly lanceolate; flowers yellowish ................................................................................................... O. dentifera Central lobe of lip ovate-quadrate or oblong; flowers pinkish ................................................................................................. O. monticola

470

O RCHIDACEAE

Octomeria anomala Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 284. 1976. Lithophyte, cespitose, sun-loving; leaves basally pseudopetiolate; flowers small, deep wine red or purple, with retrorse lateral sepals; dorsal sepal and petals forming a hood over lip and column; dorsal sepal 10−13 mm long. Tepui summits on exposed places, 2500–2700 m; Bolívar (Serranía Marutaní), Amazonas (Cerro Marahuaka). Endemic. Octomeria apiculata (Lindl.) Garay & Sweet, J. Arnold Arbor. 53: 391. 1972. —Pleurothallis apiculata Lindl., Fol. Orchid., Pleurothallis 17. 1859. Epiphyte, creeping; flowers not widely opening, pale yellow; dorsal sepal 4–6 mm long. Rain or cloud forests, 400–1100 m; Bolívar (Altiplanicie de Nuria, near Cerro Venamo-La Escalera, upper Río Caura basin). Aragua, Carabobo, Sucre, Yaracuy; West Indies. Octomeria apiculata has never been found in flower in the flora area, thus the identity of the Guayanan material currently referred to this taxon must remain doubtful. Octomeria connellii Rolfe, Trans. Linn. Soc. London, Bot. 6: 60. 1901. Epiphyte, lithophyte, or subterrestrial, cespitose, erect to arching, usually sun-loving; leaves very fleshy, much shorter than the stout stem; flowers with subparallel perianth segments; pedicellate ovary wine red; sepals and petals yellow, waxy; lip deep yellow with some orange basally; dorsal sepal 8–12 mm long. Open tepui-summit associations, often as a rock dweller, 2600–2900 m; Bolívar (Ilú-tepui, Matahui-tepui, Roraimatepui, Tramen-tepui). Guyana. Octomeria cordilabia C. Schweinf., Bot. Mus. Leafl. 19: 204. 1961. Terrestrial, lithophyte, or epiphyte, cespitose, erect, sun-loving; stems thin, often very elongate; flowers membranous, yellow; dorsal sepal 4–6 mm long. Cloud forests to, more commonly, open tepui associations, 1300– 1800 m; Bolívar (Jaua-Sarisariñama massif, Kamarkawarai-tepui), Amazonas (Cerro Aracamuni, Cerro Marahuaka, Sierra de la Neblina). Brazil. ◆Fig. 423.

Octomeria dentifera C. Schweinf., Bot. Mus. Leafl. 19: 205. 1961. Lithophyte, erect, cespitose, sun-loving; flowers white, yellowish, or cream-colored; dorsal sepal 5–6 mm long. Tepui summits in exposed situations, 2200–2500 m; Bolívar (Macizo del Chimantá, Roraima-tepui). Endemic but probably in adjacent Guyana. Octomeria erosilabia C. Schweinf., Bot. Mus. Leafl. 3: 85. 1935. Epiphyte or lithophyte, erect to subpendulous, rhizomatose; flowers fasciculate, with a short peduncle completely hidden by scarious bracts, appearing almost sessile; sepals and petals pale yellow to cream-colored, sometimes with some of red on lip. Rain or cloud forests, sometimes in rather exposed situations, sometimes in deep shade, 100–1200 m; Bolívar (Río Caroní, Río Cuyuní, and Río Parguaza basins), Amazonas (throughout). Colombia, Peru. ◆Fig. 424. Sterile specimens of this species are almost indistinguishable from the closely related Octomeria rhizomatosa but the rhizomes tend to be shorter and stouter and the leaves more oblong and shorter. Octomeria exigua C. Schweinf., Bot. Mus. Leafl. 3: 86. 1935. Octomeria exigua var elata C. Schweinf, Bull. Torrey Bot. Club 75: 216. 1948. Octomeria deltoglossa Garay, Bot. Mus. Leafl. 18: 199, t. 37. 1958. Octomeria kestrochila Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 288. 1976. Epiphyte or subterrestrial, erect, cespitose; flowers fasciculate, not opening widely; sepals and petals whitish; lip verruculose; dorsal sepal 5–8 mm long. Rain or cloud forests on tepui summits or slopes, 500–1400 m; Bolívar (Amaraway-tepui, Aparamán-tepui, Jaua-Sarisariñama massif, La Escalera to Cerro Venamo region, Macizo del Chimantá), Amazonas (Cerro Duida, Cerro Yapacana, Cerro Yutajé). Guyana, Suriname. ◆Fig. 429. Octomeria filifolia C. Schweinf., Bot. Mus. Leafl. 19: 206. 1961. Epiphyte, erect to subpendulous, cespitose, shade-loving; leaves terete or subterete, ± equal to or surpassing the stem; flowers

Octomeria 471

fasciculate, flowering 1–4 simultaneously, not opening widely, cream, whitish, or yellowish. Cloud forests, 900–1900 m; Bolívar (La Escalera to Cerro Venamo region, upper Río Caroní, Río Caura basin), Amazonas (Cerro Duida, Cerro Marahuaka). Guyana. Octomeria flaviflora C. Schweinf., Bot. Mus. Leafl. 19: 207. 1961. Terrestrial, lithophyte, or rarely an epiphyte, erect, cespitose, sun-loving; flowers fasciculate, flowering 1–4 simultaneously, yellowish, often with reddish apex at margins. Cloud forests to open tepui associations, (600–)1200–2500 m; Bolívar (Altiplanicie de Nuria, Auyán-tepui, JauaSarisariñama massif, La Escalera to Cerro Venamo region, Macizo del Chimantá), Amazonas (Cerro Aracamuni, Cerro Marahuaka, Sierra de la Neblina). Endemic. Octomeria gemmula Carnevali & I. Ramírez, Ernstia 39: 6. 1986. Low epiphyte, creeping or erect; Stems relatively short; leaves fleshy, the lower surface purple; flowers fasciculate, flowering one at a time, white with 2 red spots on lip base. Open sclerophyllous scrub on whitesand soil, 50–100 m; Amazonas (Río Sipapo basin, Yavita to Maroa road). Adjacent Amazonian Colombia. ◆Fig. 426.

Lithophyte or epiphyte, erect, cespitose; flowers with widely opening sepals and petals, subparallel to column; perianth segments purple. Cloud forests to open associations on tepui summits, 1300–2500 m; Bolívar (Macizo del Chimantá), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú, Sierra de la Neblina). Northern Brazil. Octomeria minor C. Schweinf., Bot. Mus. Leafl. 3: 89. 1935. Epiphyte, erect to pendulous, cespitose; flowers pale yellow with lip pale yellow basally, maroon-purple in the apical 1/2. Rain forests, 300–600 m; Bolívar (La Escalera to Cerro Venamo region, Río Paragua basin), Amazonas (Río Mawarinuma). Guyana. Octomeria monticola C. Schweinf., Bot. Mus. Leafl. 9: 43. 1941. Terrestrial or lithophyte, erect, cespitose, sun-loving; flowers apparently with subparallel segments; perianth segments and lip yellow with dull maroon marks or salmon, or pink with maroon marks. Open high-tepui associations, 2600–2800 m; Bolívar (Kukenán-tepui, Roraima-tepui). Guyana.

Octomeria heleneana Carnevali & Delascio, Ernstia 45: 11. 1987. Epiphyte, erect, cespitose; flowers widely opening, sepals and petals purple or bronzy with magenta tips; lip maroon-magenta; dorsal sepal ca. 3 mm long. Rain or cloud forests, 800(–2500?) m; Amazonas (Cerro Huachamacari). Endemic. ◆Fig. 425.

Octomeria nana C. Schweinf., Bot. Mus. Leafl. 19: 211, t. 30, fig. 2. 1961. Lithophyte or epiphyte, erect, cespitose; flowers fasciculate, not widely opening, yellowish or reddish orange. Open tepui associations to cloud forests, (1400–)1700–2100 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region, Murisipán-tepui, Macizo del Chimantá, Tereké-yurén-tepui), Amazonas (Sierra de la Neblina). Endemic.

Octomeria integrilabia C. Schweinf., Bot. Mus. Leafl. 3: 87. 1935. Epiphyte, erect to subpendulous, cespitose; flowers opening widely or not; sepals and petals pink or translucent yellow, often with purple-red veins and apices; lip yellow or reddish. Rain or cloud forests, 400–1400 (–1900) m; Bolívar (Río Caroní basin, Río Caura basin, Río Cuyuní basin), Amazonas (Cerro Aracamuni). Guyana.

Octomeria parvifolia Rolfe, Trans. Linn. Soc. London, Bot. 6: 60. 1901. Lithophyte, subterrestrial, or epiphyte, variable, cespitose; flowers very small, with reddish, yellowish, yellow-white, or lavender-maroon perianth segments. Open tepui associations to cloud forests, (1300–)1700– 2800 m; Bolívar (widespread above 1400 m), Amazonas (Cerro Marahuaka, Sierra de la Neblina). Sucre; Guyana. Brazil.

Octomeria lancipetala C. Schweinf., Bot. Mus. Leafl. 19: 210. 1961.

Octomeria parvula C. Schweinf., Bot. Mus. Leafl. 3: 90. 1935.

472

O RCHIDACEAE

Epiphyte, erect, cespitose; flowers very small, perianth segments yellow, lip purple. Cloud forests, ca. 1000 m; Bolívar (Macizo del Chimantá). Colombia, Guyana.

keels. Rain or cloud forests, usually in rather open places, 500–1200 m; Bolívar (Río Caroní basin), Amazonas (Cerro Marahuaka). Amazonian Ecuador.

Octomeria rhizomatosa C. Schweinf., Fieldiana, Bot. 28: 188, fig. 36. 1951. Epiphyte, rarely a lithophyte, creeping, erect to pendulous; flowers long-pedunculate with whitish or pale yellow perianth segments and lip. Rain or cloud forests, (400–) 700–1800 m; Bolívar (Altiplanicie de Nuria, Río Caroní and Río Caura basins, upper Río Paragua basin), Amazonas (Cerro Huachamacari, Cerro Marahuaka, Sierra Parima). Barinas, Táchira.

Octomeria splendida Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 292. 1976. Lithophyte or epiphyte, cespitose, erect; flowers shiny dark maroon or reddish purple with maroon, margins of golden brown, perianth segments reflexed. Tepui-summit cloud forests, open, rocky associations, always in exposed situation, 1800–2100 m; Bolívar (Jaua-Sarisariñama massif). Endemic.

Octomeria romerorum Carnevali & I. Ramírez, Ann. Missouri Bot. Gard. 77: 551. 1990. Epiphyte, creeping, minute; Stems almost absent; leaves prostrate over substrate; flowers medium-sized for the genus but very large for the plant, long-pedicellate, opening widely, entirely white. Rain forests, 50–100 m; Amazonas (Río Cataniapo basin). Endemic. ◆Fig. 428. Octomeria scirpoidea (Poepp. & Endl.) Rchb. f., Bot. Zeitung (Berlin) 10: 856. 1852. —Aspegrenia scirpoidea Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 12, t. 116. 1838. Octomeria brevifolia Cogn. in Mart., Fl. Bras. 3(4): 643. 1896. Epiphyte, erect to pendulous, cespitose, usually shade-loving; leaves terete, much shorter than the elongate stem; flowers with widely opening perianth segments, yellow with purple at apex, sometimes with reddish purple veins; lip yellow, often with purple at apex; dorsal sepal 4–6 mm long. Rain forests, 100–500(–1100) m; Bolívar (Cerro Venamo, Río Carrao), Amazonas (Río Atabapo, Río Casiquiare, Río Cataniapo, Río Negro basin, Río Sipapo basin). Colombia, Guyana, Ecuador, Peru, Brazil. ◆Fig. 430. Octomeria spathulata Rchb. f., Allg. Gartenzeitung 16: 424. 1860. Epiphyte, usually sun-loving, small to medium-sized, cespitose to creeping, erect; flowers widely opening or some populations cleistogamous, with white-red or pink-tinged perianth segments; lip yellow with purple

Octomeria steyermarkii Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 3: 204. 1965. Epiphyte, erect to subpendulous, cespitose; flowers small to very large for the genus due to the long-setose, acuminate perianth segments, widely spreading; perianth segments pink or pale pink at base grading to cream apex and a cream lip; dorsal sepal 10– 55 mm long. Cloud forests, 1100–1500 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region). Endemic. ◆Fig. 427. Octomeria steyermarkii usually grows low in trees in shady spots, the whole plant embedded in bryophytes. Octomeria surinamensis H. Focke, Tijdschr. Wis- Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 2: 200. 1849. Epiphyte, erect to pendulous, cespitose to subcreeping, sometimes the whole plant entirely red- or purple-tinged; flowers opening widely, 1 or 2 contemporaneously, wholly yellow, cream, or subhyaline; dorsal sepal 7–10 mm long. Rain forests, 100–700 m; Bolívar (Río Caroní basin), Amazonas (Río Casiquiare, Río Cataniapo, Río Negro, Río Sipapo basin). Venezuelan Coastal Range and Andean foothills; Guyana, Suriname, Peru, Brazil. Octomeria taracuana Schltr., Repert. Spec. Nov. Regni Veg. 42: 93. 1925. Epiphyte, erect to pendulous, cespitose; leaves terete or subterete; flowers not opening widely; perianth segments pale yellow at apex, grading to yellow-white at apex, some-

Octomeria 473

times with purple-maroon veins on a pinkish base with golden brown apices; lip yellow or pale brownish yellow with purple lateral lobes. Rain or cloud forests, 100–1500 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region), Amazonas (Río Temi, Tobogán de la Selva, road between Yavita and Maroa). Táchira; Brazil. Octomeria yauaperyensis Barb. Rodr., Vellosia ed. 2, 1: 120. 1891. Octomeria complanata C. Schweinf., Bot. Mus. Leafl. 10: 195. 1942. Epiphyte, shade-loving, small, erect to subpendulous, cespitose; leaves subterete, much shorter than the elongate stem; flowers small, with widely spreading perianth segments, yellow suffused with red toward apex, veins red or purplish, lip yellow suffused with some red; dorsal sepal 8−10 mm long. Rain forests, 50–200 m; Amazonas (Río Casiquiare basin, Río Cataniapo, Río Negro basin). Guyana, Peru, Brazil.

Fig. 423. Octomeria cordilabia

Octomeria sp. A Minute cespitose epiphytes; leaves 1–1.8 cm, long-elliptic or oblong-elliptic, obtuse, fleshy, longer than to as long as the stems; flowers minute, widely spreading, apparently produced singly in succession, translucent yellow-orange, lip with a deep red-orange zone on the disk; anther apparently white; dorsal sepal 2 mm long, 1.8 mm wide. Cloud forests or tepui shrublands, 1500–1600 m; Amazonas (Cuao-Sipapo massif). Endemic. This species resembles a miniature version of Octomeria cordilabia but besides the differences in plant and flower size, the lip of this new species differs in having a proportionally larger disk while the portion of the lip that flexes downward is proportionally smaller. Furthermore, the smaller petals of the new species feature only a single vein plus half of a second vein on the side of the structure that is contiguous to the dorsal sepal. The leaves of O. cordilabia are usually acute while they are obtuse in this taxon.

Fig. 424. Octomeria erosilabia

Fig. 425. Octomeria heleneana

474

O RCHIDACEAE

Fig. 426. Octomeria gemmula

Fig. 428. Octomeria romerorum

Fig. 427. Octomeria steyermarkii

Fig. 429. Octomeria exigua

Fig. 430. Octomeria scirpoidea

95. ODONTOGLOSSUM H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 350. [1815] 1816. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Mostly epiphytes, often lithophytes or subterrestrial herbs, inconspicuous to large and showy, usually erect but sometimes pendulous, cespitose to creeping to scandent. Rhizomes short to elongate, prostrate to ascendent, naked or covered with sheaths, bearing pseudobulbs at variable distances; pseudobulbs heteroblastic (one

Odontoglossum 475

internode), usually ovoid to subspheroid, sometimes oblong, apically 1–3(4)-foliate, usually enveloped by 1–several, imbricate sheaths, the innermost 1–3 may sometimes with foliar blades. Leaves conduplicate, articulate, coriaceous to fleshy, sessile to shortly pseudopetiolate, erect or spreading. Inflorescences lateral, originating from the base of the pseudobulb or from the axils of the pseudobulb sheaths, usually racemose or a variously branched panicle, rarely 1-flowered, erect, arching or pendulous, sometimes twining; peduncle, rachis, and bracts various. Flowers inconspicuous to very large and showy, resupinate, spreading to nodding, usually longlasting, mostly white, yellowish, or pink, usually spotted, striped, or blotched with red, purple, maroon, pink, or brown, sometimes the whole flower purple or maroon; pedicellate ovary mostly terete. Perianth segments spreading to campanulate, flat or undulate; sepals usually ± equal, free or rarely the laterals connate in the basal 1/3, usually ovate-elliptic or elliptic, acute to acuminate, sometimes basally clawed; petals usually similar to the sepals, rarely different. Lip basally ± parallel to column, then usually deflexed and forming an angle of ca. 90° to the column, margins entire or denticulate to fimbriate, unlobed or 3-lobed; lateral lobes erect or spreading; disk variously callose, the callus denticulate or crestate, or composed of several longitudinal ridges or plates, very rarely ecallose. Column usually elongate and straight, footless, usually with a pair of entire or denticulate to lacerate or setiform, subapical wings; anther terminal, operculate, incumbent, 1-locular or imperfectly 2-locular; pollinia 2, waxy, mostly ovoid, tegula narrowly oblong, viscidium small; clinandrium usually entire, shallow; rostellum transverse; stigma ventral. Capsules various. Colombia, Venezuela, Guyana, Ecuador, Peru, Bolivia; ca. 60 species, ca. 25 in Venezuela, 1 of these in the flora area. There is not a clear line between this genus and Oncidium Sw. Several groups within Odontoglossum are closer to groups within Oncidium than what they are within them. Some recent authors have included Odontoglossum in Oncidium, but it is being maintained separately here until the relationship with Oncidium is resolved. Odontoglossum naevium Lindl. in Lindl. & Paxton, Paxt. Fl. Gard. 1: 87, t. 18. 1853 [1850]. Epiphyte, cespitose; pseudobulbs 4–8 cm long; leaves 15–40 cm long; inflorescences racemose or rarely with 1 or 2 branches; flowers showy; sepals 3–4 cm long, white with purple spots or blotches; lip with bright yellow callus. Cloud forests, 1200–1400 m; Bolívar (La Escalera to Cerro Venamo region). Aragua, Carabobo, Falcón, Trujillo; Colombia, Guyana. ◆Fig. 431.

Fig. 431. Odontoglossum naevium

476

O RCHIDACEAE

96. OECEOCLADES Lindl., Edwards’s Bot. Reg. 18: sub t. 1522. 1832. [Subtribe Cyrtopodiinae]. Eulophidium Pfitzer, Entwurf Anordn. Orch. 87. 1887. by Gustavo A. Romero-González Cespitose or rhizomatous herbs, terrestrial or rarely epiphytes. Stem modified into aggregate or approximate pseudobulbs, ovoid to cylindrical or globose to pearshaped, of one elongate internode. Leaves 1–3, coriaceous, linear, narrowly ovate, narrowly elliptic, broadly ovate, or elliptic, acute, obtuse, or acuminate, conduplicate, most commonly petiolate, basally cordate or cuneate, rarely sessile, articulate. Inflorescences lateral, arising from the base of the pseudobulbs, erect, racemose or paniculate, few-flowered, usually much longer than the leaves. Flowers membranous, resupinate, often showy; floral bracts narrowly to very narrowly ovate, acuminate, concave, often inconspicuous. Sepals and petals membranous, variously spreading; sepals spatulate, narrowly ovate, narrowly obovate, or obovate, obtuse, acute, or subacuminate; petals < 1/2 as long as to longer than the sepals, elliptic to subcircular, acute or obtuse. Lip sessile, basally produced into a conspicuous spur, blade 3-lobed, the midlobe 2-lobed or shallowly emarginate; disk with a pair of variously shaped calli or with 3 axial, variously thickened, sparsely papillose or hirsute ridges. Column erect, short, slightly arched, semiterete; anther terminal, operculate, incumbent, 2-locular; pollinia 2, yellow, cartilaginous, subtriangular in outline, sulcate, attached to a short or rudimentary tegula. U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina, tropical Africa, Seychelles, Madagascar, Mascarene Islands; 31 species, 1 in Venezuela. Oeceoclades maculata (Lindl.) Lindl., Gen. Sp. Orchid. Pl. 237. 1833. —Angreacum maculatum Lindl., Coll. Bot. t. 15. 1821. —Eulophidium maculatum (Lindl.) Pfitzer, Entwurf Anordn. Orch. 88. 1887. Terrestrial, often weedy; pseudobulbs small (to 2.5 cm); leaves dark green-mottled; inflorescence basal, racemose; flowers 15, pink, showy. Densely shaded wet forests, 50– 300 m; Bolívar (Represa Guri, Río Yuruán, San Martín de Turumbán southwest of Tumeremo, around Upata), Amazonas (around Puerto Ayacucho, Río Cataniapo basin). Coastal Cordillera, portions of the Venezuelan Andes, Anzoátegui, Falcón, Zulia, locally frequent elsewhere; U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Guyana, French Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina, tropical Africa. ◆Fig. 432.

Fig. 432. Oeceoclades maculata

Oncidium 477

97. ONCIDIUM Sw., Kongl. Vetensk. Acad. Nya Handl. 21: 239. 1800, nom. cons. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Mostly epiphytes, but often muscicolous on the soil or on rocks, sympodials, very variable. Rhizome abbreviate or long and creeping to scandent; stems mostly pseubulbous-thickened but often very abbreviate and hidden among sheaths; pseudobulbs heteroblastic (one internode), usually oblong, ovoid, or ellipsoid, mostly ancipitous (compressed laterally), sometimes cylindrical or fusiform, basally clothed with several sheaths of which the innermost 1–6 may have foliar blades, often the sheaths bladeless, apically 1–3(–5)-foliate, sometimes with an aborted leaf at apex. Leaves conduplicate, articulate or very rarely marcescent, thinly coriaceous to fleshy-coriaceous. Inflorescences lateral, originating from base of the pseudobulb or from the axils of the leaf sheaths, 1-flowered (rarely), racemes, or variously branched panicles, sometimes very elongate and vining, in several cases proliferous and originating plantlets; peduncle and rachis usually terete but often 3-edged, rachis often zigzag; floral bracts mostly inconspicuous but sometimes ± showy; pedicellate ovary conspicuous, usually terete. Flowers mostly resupinate, inconspicuous to large and showy, usually with spreading perianth segments, erect or nodding, often long lasting; in some groups there are abortive flowers formed by 3–6 linear, undifferentiated segments and no fertile structures; perianth segments subfleshy to thickly fleshy, mostly in shades of yellow, variously spotted, blotched, or barred with brown, maroon, or reddish, sometimes the background color of the flower pink, purple, or white; sepals usually subequal, free or less frequently the lateral variously connate; petals mostly subequal to the sepals, in a few cases larger and showier. Lip free, spreading, usually larger and showier or differently colored than sepals and petals; the base sessile or shortly clawed and adnate to column base, often forming a nearly right angle to it, usually 3-lobed, often pandurate, rarely nearly entire, the central portion usually with a claw or isthmus; lateral lobes porrect (horizontally extended), erect to reflexed, sometimes very reduced; central lobe spreading, usually larger than the laterals, often transversely dilated, rarely narrow, frequently apically emarginate or bifid; disk conspicuously callose, the callus composed of 1–several variously developed and combined crests, keels, tubercles, teeth, or platforms, very rarely the lip ecallose. Column relatively short, footless, erect, sometimes geniculate, lateral margins near the stigma usually with a pair of variously developed, ± petaloid or auriculate wings or projections, very rarely column wingless; anther mostly terminal, less often subventral or dorsal, operculate, incumbent, strongly convex, semiglobose or cucullate to long-rostrate, 1-locular or imperfectly 2locular, rarely 2-locular; pollinia 2, waxy, usually deeply sulcate, mostly ovoid or subspheroid, tegula variously developed, usually elongate and strap-like or linearoblanceolate, viscidium small; clinandrium short and truncate or ovate and obliquely erect, entire or with the apex shortly bidentate; rostellum transverse; stigma ventral, variously shaped. U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina, Uruguay; ca. 400 species, ca. 40 species in Venezuela, 9 of these in the flora area.

478

O RCHIDACEAE

The circumscription of the genus has been problematic and it has been used as a catch-all genus for Oncidioid taxa with (mostly yellow) flowers mimicking Malpighiaceae. Recent phylogenetic studies have shown that as circumscribed until recently, it was grossly polyphyletic. The genus is restricted here to include only taxa with well-developed, usually somewhat laterally-compressed pseudobulbs, conduplicate leaves, a lip with well-developed calli and lateral lobes. With the exclusion of taxa formerly included in such segregates as Cyrtochilum H.B.K. (this treated as a distinct genus and absent from the Guayana Region) and Cotiniella Rchb. f. and Lophiaris Raf. (this now included in a broadly defined Trichocentrum), the genus becomes a more natural, morphologically and florally coherent assemblage of taxa growing mostly at lowland to intermediate elevations, centered in the Andean region but with distinct secondary centers of diversity in central Mexico, southern Central America, and southeastern Brazil. The boundaries, if any, between Oncidium and the high-Andean clade traditionally treated as Odontoglossum remains controversial and the latter may be eventually included in the former, despite a long tradition of horticultural recognition. Oncidium in the strict sense is poorly represented in the combined Guianas-Amazonas regions. Key to the Species of Oncidium 1.

1.

2(1).

2.

3(2).

3.

4(3).

4.

Rhizome very elongate, the distance between pseudobulbs longer than to ± equal to the length of the pseudobulb; pseudobulbs pyriform, cylindrical to ovoid, not or little compressed; lip white or cream with purple spots ........................................................................................ O. warmingii Rhizome very short, when somewhat elongate the distance between pseudobulbs several times less than the length of the pseudobulbs; pseudobulbs ovoid to oblong, always somewhat to strongly compressed; lip yellow ................................................................................................ 2 Central lobe of lip much narrower than the basal blade, narrowly oblong or narrowly elliptic, acute; perianth segments white with red or maroon blotches ............................................................................ O. nigratum Central lobe of lip as wide or wider than basal lamina, suborbicular or transversely elliptic or oblong, rounded or emarginate apically; perianth segments yellow with brown or maroon blotches to concolorous ................................................................................................................ 3 Central lobe of lip 16–36 mm wide (usually 20 mm), 4–5 times wider than lateral lobes or lateral lobes almost absent; pseudobulbs only slightly if at all compressed when fresh; leaves strongly coriaceous; plants chiefly terrestrial over sandy or sandstone substrate on places of heavy rainfall or over granitic outcrops in places with a well-marked dry seasons ................................................................................................ O. orthostates Central lobe of lip rarely to 15 mm wide, ± equal in width to the basal blade or lobes; pseudobulbs usually strongly compressed when fresh; leaves thinly coriaceous; plants chiefly epiphytic ................................. 4 Pseudobulbs 1-foliate; central lobe of lip separated from basal blade or lobes by a long, narrow, oblong claw that is < 1/5 as wide as the central lobe ....................................................................................... O. boothianum Pseudobulbs 2- or 3-foliate (rarely 1-foliate); central lobe of lip separated from basal lobes or blade by a broad, usually triangular claw, when the

Oncidium 479

5(4).

5.

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

claw is oblong it is always at least 1/3 as wide as the central lobe ....... 5 Floral bracts 2–3 mm long; central lobe of lip 6–7 mm wide; callus of lip with 9 long narrow teeth; column with relatively narrow and long, bifurcate auricles at each side ............................................... O. dactyliferum Floral bracts 5–12 mm long; central lobe of lip 8–16 mm wide; callus of lip with 3–5 broad triangular teeth; column with one triangular or dolabriform auricle at each side ....................................................................... 6 Central lobe of lip conspicuously broader than basal blade, almost sessile on it ...................................................................................... O. sphacelatum Central lobe of lip narrower than or ± equal to basal blade, conspicuously clawed ..................................................................................................... 7 Column with 2 small frontal teeth below stigma and auricles; inflorescences usually > 150 cm ................................................................ O. baueri Column without frontal teeth below stigma; inflorescences usually < 100 cm ................................................................................................. 8 Midlobe of lip about as broad as or somewhat broader than the expanded basal lateral lobes ..................................................................... O. citrinum Midlobe of lip conspicuously broader than the expanded lateral lobes .................................................................................................... O. pardalis

Oncidium baueri Lindl. in F.A. Bauer & Lindl., Ill. Orch. Pl. t. 7. 1833. Oncidium kappleri Rchb. f. ex Lindl., Fol. Orchid., Oncidium 47. 1855. Epiphyte or muscicolous herb, on rocks or on the soil, mostly sun-loving; rhizome creeping; leaves 20–50 cm long; flowers pale or bright yellow with brown blotches on petals, sepals, and base of claw of the lip, lip 12–17 × 7–12 mm. Rain or deciduous forests, near sea level to 200 m; Delta Amacuro (Caño Atoiba, Río Amacuro), Bolívar (lower Río Caroní). Aragua, Carabobo, Mérida, Monagas; West Indies, widespread in tropical South America. Oncidium boothianum Rchb. f., Bonplandia (Hanover) 2: 14. 1854. Epiphyte, mostly sun-loving; leaves 14–25 cm long; flowers pale yellow with brown blotches on sepals, petals, and claw and disk of the lip, lip 7–12.5 × 5–11 mm. Cloud forests, 1000–1200 m; Amazonas (Sierra de la Neblina). Aragua, Carabobo, Distrito Federal, Falcón, Yaracuy. Despite the striking disjunction, the single collection from Sierra de la Neblina is apparently conspecific with the Coastal Range populations. Oncidium citrinum Lindl., Edwards’s Bot. Reg. 21: t. 1758. 1836. Epiphyte, mostly shade-loving; leaves 10– 20 cm long; flowers yellow with brown or maroon blotches, the lip almost entirely yellow,

15–18 × 13–15 mm. Rain forests, 400–700 (–1000) m; Bolívar (Altiplanicie de Nuria, Gran Sabana). Distrito Federal, Miranda, Monagas; Trinidad-Tobago. Colombia. This taxon is rare but widespread in the eastern Venezuelan Guayana. Oncidium dactyliferum Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 5: 224. 1972. Epiphyte; leaves 8–55 cm long, narrowly oblong or linear; inflorescences shorter than or ± equal to the leaves; flowers with yellowish, greenish, or green-brown sepals and petals, lip pale yellow with a transverse, browngreen band on the isthmus, 9–11 × 7–9 mm. Rain forests, 100–300 m; Bolívar (Río Canaracuni), Amazonas (Río Siapa). Amazonian Ecuador. Oncidium dactyliferum is a rare lowland entity with a most distinctive callus on the lip. Oncidium nigratum Lindl. in Lindl. & Paxton, Paxt. Fl. Gard. 1: 78. 1853 [1850]. Oncidium uaipanense Schnee, Revista Fac. Agron. (Maracay) 1(1): 118, t. s.n. 1952. Usually terrestrial on sandy soil, a lithophyte on sandstone, or a muscicolous herb, rarely an epiphyte, sun-loving; leaves 10–33 cm long; sepals and petals white or cream with magenta or purple spots; lip 7–13 × 6–

480

O RCHIDACEAE

10 mm, yellow with a transverse magenta or brownish band close to the claw. Cloud forests or shrublands on tepui summits, where locally common, (1300–)1600–2200 m; Bolívar (widespread). Colombia, Guyana. ◆Fig. 434. Oncidium orthostates Ridl. ex Thurn, Timehri 5: 204. 1886. Terrestrial on sandy soil, humicolous over sandstone or igneous outcrops, or rarely a low epiphyte, sun-loving; inflorescences often viviparous; leaves 5–30 cm long; sepals and petals yellow heavily blotched or spotted with brown, or more commonly, completely dark, shiny brown or bronzy, with a dark yellow-green base; lip 17.5–28.5 × 25–36 mm, much larger than the other perianth segments, bright, waxy, lustrous yellow. Locally common in open rocky outcrops, shrublands, or forest margins, rarely inside forests, ca. 50–1200(–1600) m; Bolívar (Cerro Bolívar, widespread in upper Río Caroní basin, rarer in Río Cuyuní basin), Amazonas (Cerro Coro Coro, Cerro Duida, near Puerto Ayacucho, Río Mapa, Salto Yureba). Guyana, Amazonian Brazil. ◆Fig. 433. Populations of Oncidium orthostates from the granitic outcrops in northern Amazonas state feature consistently smaller flowers and might represent a distinct variety or subspecies.

Oncidium warmingii Rchb. f., Otia Bot. Hamburg. 2: 86. 1881. Oncidium vagans C. Schweinf., Bot. Mus. Leafl. 18: 223, t. 45. 1958. Lithophyte or humicolous herb on sandstone or sandy soil, rarely an epiphyte, sunloving; leaves 3.5–15 cm long; sepals and petals cream or yellowish with brown, red, or maroon spots or blotches; lip 10–13 × 8–12 mm, callus bright yellow. Locally common in open, rocky places and shrublands, rarely in forests, 1600–2700 m; Bolívar (Auyán-tepui, Macizo del Chimantá, Sierra de Maigualida, Uaipán-tepui), Amazonas (Cerro Duida, Cerro Marahuaka). Southern Brazil. ◆Fig. 435. Oncidium warmingii is another species with a strikingly disjunct distribution.

Ondidium pardalis Rchb. f., Bonplandia 2: 13. 1854. Epiphyte, vegetatively similar to Oncidium citrinum; leaves to 25 cm long; petals and sepals yellow with dense brown or maroon blotches, appearing chestnut-colored. Rain forests, 400–700(–1000); Bolívar (Gran Sabana). Distrito Federal. Oncidium sphacelatum Lindl., Sert. Orchid. sub t. 48. 1841. Epiphyte, rarely a humicolous herb, sunloving; leaves 25–100(–130) cm long; flowers bright yellow with brown or maroon spots on lower 1/2 of sepals and petals, and on lip claw; lip 12–17 × 12–16 mm. Rain forests, 50–200 m; Bolívar (lower Río Caroní). Mexico, Central America. Despite the disjunction, the Venezuelan Guayana plants appear conspecific with the Mexican and Central American populations.

Fig. 433. Oncidium orthostates

Oncidium 481

Fig. 434. Oncidium nigratum

Fig. 435. Oncidium warmingii

482

O RCHIDACEAE

98. OPHIDION Luer, Selbyana 7: 79. 1982. [Subtribe Pleurothallidinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, rarely lithophytes, small to medium-sized, cespitose, shade-loving, erect to subpendulous. Ramicauls relatively short, subterete, erect to horizontal, enveloped by 2 or 3 tubular, papery, relatively large sheaths, 1 of them much larger than the basal ones. Leaves conduplicate, elliptic to narrowly elliptic, thin to relatively coriaceous, basally narrowed to a pseudopetiole. Inflorescence originating from ramicaul apex with an annulus, a successively 1–few-flowered raceme, peduncle filiform, shorter to rarely ± equal to the leaves, erect to nodding or even subcreeping; rachis filiform, straight to somewhat zigzag. Flowers small to mediumsized for the subtribe, resupinate, horizontal. Lateral sepals variously connate into a synsepal; synsepal connivent at base and apex with dorsal sepal apex so the flower only opens by means of 2 longitudinal, lateral slits; petals much smaller than sepals, basally attenuate. Lip rigidly attached to column foot and held inside the synsepal, nearly filling it, divided into a hypochile and an epichile, 3-lobed; lateral lobes thick, hypochile concave, epichile usually flat. Column relatively long, terete, arching, apically thickened; anther ventral, incumbent, operculate, 2-locular; pollinia 2, unappendaged; rostellum transverse; stigma ventral. Panama, Colombia, Venezuela, Ecuador; 4 species, 1 in Venezuela. Ophidion pleurothallopsis (Kraenzl.) Luer, Selbyana 7: 80. 1982. —Cryptophoranthus pleurothallopsis Kraenzl., Bull. Misc. Inform. Kew 1925: 11. 1925. Epiphyte or muscicolous herb, cespitose; leaves relatively thin; inflorescences 1- or 2flowered; flowers pale cream-colored with wine purple spots. Cloud forests, 1200–1500 m; Bolívar (La Escalera to Cerro Venamo region, Macizo del Chimantá). Guyana, Ecuador. ◆Fig. 436.

Fig. 436. Ophidion pleurothallopsis

99. ORLEANESIA Barb. Rodr., Gen. Spec. Orchid. 1: 63. 1877. [Subtribe Laeliinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes or lithophytic herbs. Rhizome abbreviate. Secondary stems erect or suberect, cylindric or fusiform (rarely pear-shaped), basally stipitate or not, distichously pauci- to plurifoliate. Leaves conduplicate, fleshy or coriaceous, articulated with their sheaths, usually linear to oblong, rarely narrowly elliptic. Inflorescences terminal, usually laxly flowered, successively panicles, rarely abbreviate racemes. Flowers medium-sized to small, resupinate or nearly so, usually green or yellowish green, variously tinged or spotted with purple, brown, maroon, or orange, opening well or with ± spreading perianth segments. Sepals free, spreading, usually elliptic or oblong-elliptic; lateral sepals usually somewhat falcate or oblique and broader than the dorsal sepal; petals generally linear-oblong or narrowly spatulate, always narrower than sepals. Lip articulate with the column foot, always simple, usually oblong-obovate or obovate with a truncate or retuse apex, minutely apiculate or not; base somewhat attenuate; blade of the lip flat or, more typically S-

Orleanesia

483

shaped, recurved; disk ecallose. Column arched or (rarely) straight, semiterete or subtrigonous, ventrally concave and with winged ventral margins, basally produced into a short but conspicuous foot; anther incumbent, operculate, subglobose, 2-locular and each locule longitudinally chambered; pollinia 4, waxy, uniseriate, obovoid and with caudicles, somewhat laterally compressed, generally the outermost slightly larger; clinandrium somewhat 3-dentate; stigmatic surface ventral; rostellum transverse. Capsules fusiform to subspherical. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 12 species, 4 in Venezuela, 3 of these in the flora area.

Fig. 437. Orleanesia yauaperyensis

484

O RCHIDACEAE

Key to the Species of Orleanesia 1. 1. 2(l).

2.

Inflorescences shorter or only slightly longer than leaves; petals > 1/2 as wide as lateral sepals ............................................................. O. amazonica Inflorescences much longer than upper leaves; petals very narrow, < 1/3 as wide as lateral sepals ............................................................................. 2 Leaves very fleshy, widely channeled above; pseudobulbs conspicuously stipitate; petals linear spatulate; plants from the dryer northern part of the the flora area ................................................................. O. maculata Leaves coriaceous, flat or almost so; pseudobulbs not stipitate; petals linear; plants from low forests or shrublands on humid, white-sand soils .......................................................................................... O. yauaperyensis

Orleanesia amazonica Barb. Rodr., Gen. Spec. Orchid. 1: 64. 1877. Epiphyte, erect, cespitose; inflorescences abbreviate; sepals and petals green or greenish yellow; column with maroon streaks on its ventral face. Semideciduous forests to rain forests, often growing high on trees or in exposed positions, 100–300 m; Bolívar (Río Cuyuní basin), Amazonas (Río Ocamo). Guyana, Suriname, Brazil. Orleanesia maculata Garay, Canad. J. Bot. 34: 256. 1956. Epiphyte or rarely a lithophyte, erect, cespitose; inflorescences elongate with successive flowers on branches of panicles; sepals and petals greenish yellow, sometimes with maroon zones or blotches; ventral face of column green with purple or maroon margins; lip greenish yellow or sometimes

brownish with brick red. Deciduous or rain forests edges, 50–300 m; Bolívar (Cerro Arimagua, Represa Guri, Río Caura, near Villa Lola), Amazonas (near Puerto Ayacucho). Apure, Aragua, Guárico, Monagas, Portuguesa, Yaracuy; Colombia. Orleanesia yauaperyensis (Schltr.) Barb. Rodr., Vellosia ed. 2, 1: 124. 1891. —Huebneria yauaperyensis Schltr., Beih. Bot. Centralbl. 42(2): 97. 1925. Epiphyte 20–60 cm tall, erect, cespitose, sun-loving; inflorescences elongate, with short lateral branches bearing successive, small flowers; sepals and petals greenish with purplish or maroon zones; margins of column with deep maroon streaks. Open, low, humid scrublands over white sand, in rather exposed positions, 50–200 m; Amazonas (Río Sipapo basin). Peru, Amazonian Brazil. ◆Fig. 437.

100. ORNITHOCEPHALUS Hook., Exot. Fl. 2, t. 127. 1825. [Subtribe Ornithocephalinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, rarely twig epiphytes, minute to medium-small, cespitose, erect to pendulous, usually shade-loving. Roots thin, long, often shortly woolly. Stem much reduced. Leaves fleshy or coriaceous, laterally compressed, equitant, articulated with their sheaths, forming a fan-like plant. Inflorescences lateral, originating from leaf sheath axils, racemose, shorter to longer than leaves, erect, arched to nodding; peduncle glabrous to lanuginose, relatively short; floral bracts cordate, stem-clasping, glabrous to pubescent. Flowers small, resupinate or not, usually green, white or yellowish. Perianth segments widely spreading; sepals free, subequal, concave; petals similar to sepals but broader or very different, being broadly cuneate, fan-shaped to suborbicular. Lip subsessile, fleshier than other perianth segments, unlobed or lobed, basally callose, the basal callosities occasionally extending laterally and considered as basal lateral lobes. Column usually short, footless, wingless; anther sub-

Ornithocelphalus 485

dorsal or subapical, operculate, rostrate, the long rostrate apex is superimposed on the long rostellum, operculate, incumbent; pollinia 4, cartilaginous, attached to a very long, variously curved or arched tegula, and a small, ovoid to ellipsoid viscidium; rostellum very long and narrow, subparallel or not to lip; stigma basal. Capsules broadly ellipsoid to obovoid, smooth to with sharp edges (erinaceous). Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 25 species, 5 or 6 in Venezuela, 4 of these in the flora area. Key to the Species of Ornithocephalus 1.

1.

2(1).

2.

3(1). 3.

Plants and inflorescences glabrous; roots lanuginose; leaf sheaths inconspicuous, < 1/5 the length of the leaf blade; petals obovate, minutely apiculate, margins entire or at most erose at the apex ............................. 2 Plants with inflorescences and external face of perianth segments pubescent to lanuginose; roots glabrous; leaf sheaths inconspicuous, at least 1/3 the length of the leaf blade; petals suborbicular to broadly obovate to subquadrate, not apiculate, margins ciliate ......................................... 3 Lip base suborbicular when flattened, deeply concave and with erect lateral lobes when fresh, the concavity with white hairs, the dorsal face bearing a massive, Y-shaped retrorse projection, the section of lip immediately distal to the base flat or convex on the ventral face; apical margin of petals denticulate to finely erose .............................. O. falcatus Lip base transversely oblong when flattened, flat or nearly so when fresh, the dorsal face lacking massive, Y-shaped retrorse projections, the section of lip immediately distal to the base with 2 parallel keels or calli on the ventral face; apical margin of petals glabrous ................. O. gladiatus Lip 4–5 times as long as wide, with 2 transverse, lobe-like callus basally; dorsal sepal suborbicular ..................................................... O. brachyceras Lip to 2.5 times as long as wide, with 4 low, broad, rounded, longitudinal keels; dorsal sepal obovate ........................................................ O. kruegeri

Ornithocephalus brachyceras G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1140. 2000. Ornithocephalus bicornis auct. non Lindl. 1846: sensu Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 2: 262, fig. 436. 1961; sensu Foldats in Lasser, Fl. Venez. 15(5): 18. 1970. Twig epiphyte to 12 cm tall; inflorescences shorter than leaves; peduncle pale green; perianth segments greenish or whitish. Rain forests, 200–300 m; Bolívar (lower Río Caura basin). Miranda, Monagas; Guatemala, Honduras, Nicaragua, Costa Rica, Panama. Ornithocephalus brachyceras is a recently described taxon that has been confused with O. bicornis. Ornithocephalus bicornis, however, is a Mexican species that features much longer lateral lobes on the lip.

Ornithocephalus falcatus H. Focke, Tijdschr. Wis- Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 1: 211. 1848. Epiphyte; leaves to 3 × 0.4 cm; perianth segments white, basal half of the lip with 2 involute yellow calli; dorsal sepal ca. 3 × 2 mm. Rain forests, ca. 900 m; Bolívar (near Betania along the road from Santa Elena to Perai-Tepui). Guyana, Suriname, French Guyana, Ecuador, Brazil. ◆Fig. 439. Ornithocephalus falcatus is frequently treated as a synonym of O. gladiatus Hook., which has a different lip (see key) and entire petals. Ornithocephalus falcatus was cited by Foldats [Flora de Venezuela 15(5): 24. 1970] as likely to occur in Venezuela, a statement that is confirmed by this report. The species appears to be fairly widespread but rare and

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variable. The white hairs on the base of the lip seem diagnostic for the species.

Nuria), Amazonas (near Gavilán). Miranda, Monagas; Trinidad-Tobago, Suriname, Ecuador, Peru, Amazonian Brazil.

Ornithocephalus gladiatus Hook., Exot. Fl. 2: pl. 127. 1824. Epiphyte; perianth segments translucent; lateral lobes and base of lip bright green. Rain forests, 50–200 m; Amazonas (basin of Río Casiquiare and Negro). Anzoátegui, Miranda, Táchira; southern Mexico, Central America, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guyana, Peru, Brazil, Bolivia. ◆Fig. 438. Ornithocephalus kruegeri Rchb. f. in Walp., Ann. Bot. Syst. 6: 495. 1863. Epiphyte; perianth segments whitish; lip whitish with pale green at apex. Rain forests, 200–500 m; Bolívar (Altiplanicie de

Fig. 438. Ornithocephalus gladiatus

5 mm

5 mm

5 cm

5 mm Fig. 439. Ornithocephalus falcatus

Otoglossum 487

101. OTOGLOSSUM (Schltr.) Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 41. 1976. —Odontoglossum subgen. Otoglossum Schltr., Repert. Spec. Nov. Regni Veg. Beih. 27: 109. 1924. [Subtribe Oncidiinae]. Oncidium sect. Varicosa subsect. Serpentia Kraenzl. in Engl., Pflanzenr. IV 50(Heft 80): 167. 1922. —Oncidium sect. Serpentia (Kraenzl.) Garay, Taxon 19: 455. 1970. Odontoglossum H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 350. 1815 [1816], pro parte by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, lithophytes, or subterrestrial herbs, 20–100 cm tall, erect, mostly sun-loving. Rhizome creeping, well developed, stout, appressed to substrate or ascendent, or very elongate, twining, wiry, with pseudobulbs distant (ontogenetically a modified inflorescence). Pseudobulbs heteroblastic (one internode), widely spaced on the rhizome, 1- or 2-foliate apically, mostly oblongoid or ovoid to flattenedsubcircular, enveloped by several imbricate sheaths, the 1 or 2(3) innermost with or without foliar blades. Leaves conduplicate, articulate, thin to fleshy or thickly coriaceous, flat or concave, erect or spreading, mostly oblong to elliptic, apically rounded or obtuse, often emarginate, basally sessile or with a ± developed pseudopetiole. Inflorescences originating from the base of the pseudobulb or among the sheaths that envelope it, racemose, (1)2–15(–25)-flowered, erect and dense, or apparently 1-flowered, or laxly multiflowered on a twining rachis (in O. scansor and related taxa); peduncle when obvious (in O. arminii and related taxa) stout, straight, terete, remotely sheathed, much longer than leaves; rachis straight or ± zigzag; floral bracts often conspicuous but always shorter than pedicellate ovary, boat-shaped, often papery. Flowers resupinate, showy, widely spreading, pedicellate ovary terete, relatively elongate. Perianth segments bright yellow with red-purple spots on sepals and petals (in O. scansor and related taxa) or dull yellow, or yellow-brown with darker brown or orange-brown blotches, to totally dark red-brown (in O. arminii and related taxa); sepals ± equal or the laterals somewhat oblique, free, mostly obovate or elliptic-obovate, apically broadly obtuse to rounded, basally sometimes ± clawed; petals subequal to the sepals or broader, the apex obtuse, rounded, or truncate; lip deflexed, the base replicate and adnate to column base; pandurate with the constriction about or below its middle, a pair of spreading or retrorse basal lobes below the constriction; central lobe obovate to suborbicular, rounded to truncate; disk with a complex callosity composed of several lamellae, tubercles, and teeth, usually bright yellow or purple-spotted; column short, semiterete, ascendent, often forming a small, obtuse mentum with the ovary, footless, apically with 2 conspicuous, axshaped (dolabriform) wings; anther dorsal or subventral, operculate, incumbent; pollinia 2, waxy, tegula subquadrate, psygmoid (fan-like with laterally compressed leaves), viscidium small; clinandrium hooded or not, margin entire or erose; rostellum transverse; stigma ventral, subquadrate to subcircular. Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, Amazonian Brazil, Bolivia; ca. 10 species, 4 in Venezuela, 3 of these in the flora area. The genus is composed of two apparently unrelated clades, whose affinity was not hinted at until recently with the advent of molecular techniques to perform phylogeny reconstructions. The group of taxa around Otoglossum scansor (Oncidium Sect. Serpentia) is characterized by vegetatively reduced plants (small, flattened pseudobulbs) featuring long, twining inflorescences which produce plantlets,

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(“keikis,” vegetative offsets) along its length. These plantlets originate 1- or 2-flowered inflorescences with flowers that are “Oncidium”-like in appearance. Plants of taxa of this clade are restricted to the upper canopy, where they often form large, vining clumps among twigs or thinner branches that flower infrequently but all at once put out a conspicuous display. On the other hand, the Otoglossum brevifolium (Lindl.) Garay & Dunst. complex (Otoglossum sensu stricto) features larger plants with thick-textured leaves and stout, erect inflorescences with larger flowers that look more like the flowers of members of the Andean genus Odontoglossum. These plants occupy different habitats from those where the former group occurs, such as larger branches, open rocky places, or even subterrestrial or marshy environments. Key to the Species of Otoglossum 1. 1.

2(1). 2.

Rhizome thick, shortly creeping; leaves thickly coriaceous; inflorescence erect, stiff, multiflowered raceme; petals broader than lip ....... O. arminii Rhizome slender, wiry, vining, most probably a modified inflorescence with alternating flowers and pseudobulbs, these pseudobulbs originating new inflorescences that in time become also wiry; leaves coriaceous; inflorescences apparently 1- or 2-flowered (but eventually becoming rhizome-like); petals much narrower than lip ......................... 2 Central lobe of lip sessile or subsessile, overlapping lateral sepals ..................................................................................................... O. scansor Central lobe of lip forming an elongate isthmus, exposing lateral sepals ............................................................................................ O. globuliferum

Otoglossum arminii (Rchb. f.) Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 41. 1976. —Odontoglossum arminii Rchb. f., Bonplandia (Hanover) 3: 66. 1855. Odontoglossum brevifolium auct. non. Lindl. 1844: sensu Foldats in Lasser, Fl. Venez. 15(5): 202. 1970. Usually subterrestrial, rarely an epiphyte, 20–50 cm tall; inflorescences (1)2–7flowered; petals and sepals light to medium yellow-brown, bronze, or reddish orange, the margins mostly paler, often with some pale brown, yellow, or chestnut blotching; lip bright yellow with a darker transverse band in the constriction. Open associations on tepui summits, in tepui meadows or on rocky outcrops, rarely as a low epiphyte in shrublands, 1700–2600 m; Bolívar (JauaSarisariñama massif), Amazonas (Cerro Marahuaka, Sierra de la Neblina). Mérida; Colombia, Amazonian Brazil. ◆Fig. 440. The Guayanan plants here referred to Otoglossum arminii might represent a different species from the Andean populations.

Otoglossum globuliferum (H.B.K.) N.H. Williams & M.W. Chase, Lindleyana 16(2): 138. 2001. —Oncidium globuliferum H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 347. 1815 [1816]. Oncidium sancti-pauli auct. non Kaenzl. 1922: sensu Foldats in Lasser, Fl. Venez. 15(5): 387. 1970. Oncidium convolvulaceum Lindl. in Lindl. & Paxton, Paxt. Fl. Gard. 1: 102. 1853 [1850]. Epiphyte; leaves 3–11 cm long; flowers yellow with brownish red spots near the base of petals and sepals; lip 19–28 × 19–35 mm. Cloud forests, 1000–1500 m; Bolívar (La Escalera to Cerro Venamo region), Amazonas (Cerro Marahuaka). Mérida; Costa Rica, Panama, Colombia, Ecuador, Peru, Bolivia. Otoglossum scansor (Rchb.f.) Carnevali & I. Ramirez, comb. nov. —Oncidium scansor Rchb. f., Linnaea 22: 844. 1850. Oncidium globuliferum auct. non H.B.K. 1815 [1816]: sensu Dunst. & Garay, Ven.

Otostylis 489

Orch. Ill. 2: 249. 1961; Foldats in Lasser, Fl. Venez. 15(5): 323. 1970. Epiphyte; leaves 2.5–10 cm long; flowers yellow with some red-brown spotting on the lower 1/2 of sepals and petals, and on lip claw; lip 17–38 × 20–50 mm. Cloud forests, Bolívar (La Escalera to Cerro Venamo region), Amazonas (Cerro Marahuaka). Widespread and locally common in northern Venezuela; Colombia.

Fig. 440. Otoglossum arminii

102. OTOSTYLIS Schltr., Orchis 12: 38. 1918. [Subtribe Zygopetalinae]. by Gustavo A. Romero-González Cespitose epiphytes. Stem abbreviate, modified into small pseudobulbs, concealed by distichous, scarious sheaths. Leaves 1–several, membranous to coriaceous, narrowly linear-ovate, rarely ovate, acute to acuminate, attenuate toward the base to petiolate, plicate, articulate. Inflorescences lateral, arising from the base of the pseudobulbs, erect to slightly arching, racemose, few- to many-flowered, generally longer than the leaves. Flowers resupinate; floral bracts concave, appressed, narrowly ovate to ovate, obtuse to acute, < 1/2 as long as the pedicellate ovary. Sepals and petals similar, spreading, membranous, oblong, ovate-elliptic, or obovate, obtuse, acute, or acuminate, the lateral sepals and petals sometimes slightly oblique. Lip sessile to clawed, fleshy, concave, basally truncate to slightly cuneate, 3-lobed,

490

O RCHIDACEAE

Fig. 441. Otostylis brachystalix

lateral lobes small, triangular or auriculate, midlobe ovate, subrhombic, subcircular, or obovate, apically truncate, apiculate or shallowly emarginate; disk with a transverse, semicircular, or W-shaped callus, slightly to strongly retrorse, entire, irregularly denticulate or verrucose, sometimes 2- to 3-keeled in front. Column short, arched, semiterete, apically auriculate; anther terminal; pollinia 4, yellow, cartilaginous, dorsoventrally flattened, subequal, obovate, attached in 2 pairs to a small, subtriangular tegula. Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Peru, Brazil; 2 species, both in the flora area.

Palmorchis 491

Key to the Species Otostylis 1. 1.

Leaves stiff, coriaceous; lip callus smooth to at most dentate along the edge ..................................................................................... O. brachystalix Leaves soft, membranaceous; lip callus densely tuberculate throughout ....................................................................................................... O. lepida

Otostylis brachystalix (Rchb. f.) Schltr., Orchis 12: 39. 1918. —Zygopetalum brachystalix Rchb. f. in Walp., Ann. Bot. Syst. 6: 660. 1863. Zygopetalum paludosum Cogn., Comiss. Linhas Telegr. Estratég. Mato Grosso Amazonas, anexo 5, Bot. 3: 12. 1912. —Otostylis paludosa (Cogn.) Schltr., Repert. Spec. Nov. Regni Veg. 15: 214. 1918. Terrestrial; inflorescences erect, to 20flowered; flowers white, showy; lip white, often with purple spots at the base; callus semicircular, smooth to at most dentate along the edge, yellow; column white, sometimes ventrally and basally purple. Seasonally moist savannas, morichales, moist, tall forests, 800–1700 m; Bolívar (Auyán-tepui, Guayaraca, Ilú-tepui, Río Apacará, Río Uaiparú, Santa Elena de Uairén), Amazonas (Caño Ucata southeast of Síquita, base of Cerro Yapacana, south of Puerto Ayacucho, Río Casiquiare, Río Siapa). Apure; Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Peru, Brazil (Amazonas, Pará, Mato Grosso). ◆Fig. 441.

Otostylis lepida (Linden & Rchb. f.) Schltr., Orchis 12: 40. 1918. —Aganisia lepida Linden & Rchb. f., Beitr. Orchid.-K. C. Amer. 15, t. 5. 1866 [1867]. Aganisia alba Ridl. ex Thurn, Timehri 5: 204. 1886. —Koellensteinia alba (Ridl. ex Thurn) Schltr., Orchis 9: 32. 1915. —Otostylis alba (Ridl. ex Thurn) Summerh., Bull. Misc. Inform. Kew 1951: 293. 1951. Zygopetalum venustum Ridl. ex Oliv., Trans. Linn. Soc. London, Bot. 2: 283. 1887. —Aganisia venusta (Ridl. ex Oliv.) Rolfe, Bot. Mag. 118: sub t. 7270. 1892. —Otostylis venusta (Ridl. ex Oliv.) Schltr., Orchis 12: 41. 1918. Terrestrial; inflorescences erect to 8-flowered; flowers white, showy; lip white, often with purple spots at the base; callus roughly W-shaped, densely tuberculate throughout, yellow; column white, ventrally and basally purple. Moist savannas and semi-dwarf forests, 50–1700 m; Bolívar (Cerro Venamo, km 125 south of El Dorado, banks of Río Kukenán, Río Yuruán, base of Salto Angel). Brazil (Amazonas).

103. PALMORCHIS Barb. Rodr., Gen. Spec. Orchid 1: 169. 1877. [Subtribe Palmorchidinae]. Neobartlettia Schltr., Repert. Spec. Nov. Regni Veg. 16: 440. 1920. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial herbs, erect, cespitose, shade-loving. Roots fibrous or somewhat fleshy, elongate, few. Rhizomes underground, short, horizontal or somewhat ascendent. Stems terete, elongate, unbranched, rarely branching in the lower 1/5, enclosed in a series of imbricate, tubular to somewhat inflated sheaths, sometimes the basal 1/3 naked, foliose in the upper 1/2–2/3. Leaves nonarticulate, folded, convolute, spiral or subdistichous, suberect, horizontal to arched-spreading, subsessile to conspicuously petiolate; petiole grooved; blade linear-elliptic to broadly elliptic or broadly ovate-elliptic, with conspicuous longitudinal veins, obtuse to shortly acuminate. Inflorescences terminal or lateral, or both, when lateral arising from internodes at different levels on the stem, often several inflorescences simultaneously at the same or different internodes, racemose or rarely few-branched, always shorter than leaves; peduncle terete, laxly or densely covered with several to many, loose, scarious, or

492

O RCHIDACEAE

subfoliaceous bracts; rachis spiraled, straight or ± zigzag; floral bracts conspicuous, similar to peduncle bracts, ± equal to or longer than pedicellate ovary. Flowers inconspicuous, resupinate, spreading, fugacious, membranous, successive. Perianth segments white, greenish, or yellowish; sepals free, connivent, similar or the lateral somewhat oblique, mostly oblanceolate; petals similar to the sepals but often shorter and narrower. Lip greenish, purplish, or bluish, free or connate with the ventral face of column in the basal 1/4, broader than the other perianth segments, parallel to column, concave, ± 3-lobed; lateral lobes erect and ± enfolding the column; central lobe shorter than the lateral lobes, triangular, triangular-ovate to suborbicular or transversely pandurate, erect or somewhat deflexed; disk pilose or glabrous, variously tuberculate or lamellate. Column same color as the sepals, erect or somewhat arched, footless, slender, semiterete; anther terminal, incumbent, operculate, glabrous; pollinia 4, soft but coherent; clinandrium short, concave; rostellum transverse; stigma ventral, emergent. Capsules often drying black, subspheric to linearoblong. Central America, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil; ca. 12 species, 3 or 4 in Venezuela, 3 of these in the flora area. Some recent authors recognize the genus Neobartlettia based on the lip ± free from the column, in contrast with typical Palmorchis in which the lip is connate with the ventral face of the column in the basal fourth. Key to the Species of Palmorchis 1. 1. 2(1). 2.

Inflorescences lateral, terminal, or both; leaves 20–50 × (3–)5–7 cm; sepals ca. 18 mm long; central lobe of lip suborbicular .......... P. pubescens Inflorescence always terminal; leaves 8–27 × 1.1–10 cm; sepals < 14 mm long; central lobe of lip triangular to triangular ovate . ....................... 2 Inflorescence 7–12 mm long; central lobe of lip ca. 1/3 of the total lip length ...................................................................................... P. guianensis Inflorescence 10–30 mm long; central lobe of lip < 1/6 the total lip length ......................................................................................................... P. puber

Palmorchis guianensis (Schltr.) C. Schweinf. & Correll, Bot. Mus. Leafl. 8: 113. 1940. —Neobartlettia guianensis Schltr., Repert. Spec. Nov. Regni Veg. 16: 441. 1920. Terrestrial 30–70 cm tall; flowers greenish or yellowish with some purple or lilac on the central lobe of lip; sepals 10–13 mm long. Rain forests, consisting of widely dispersed individuals, 100–600 m; Bolívar (Altiplanicie de Nuria, La Escalera to Cerro Venamo region), widespread in southern Amazonas. Guyana, Amazonian Brazil. ◆Fig. 443. Palmorchis puber (Cogn.) Garay, Caldasia 8: 518. 1962. —Elleanthus puber Cogn. in Mart., Fl. Bras. 3(5): 333. 1898.

Terrestrial to 60 cm tall; flowers greenish white or yellowish; lip white with purple central lobe; sepals 6–12 mm long. Rain forests, 100–200 m; Amazonas (Río Casiquiare, Río Pasimoni, Río Siapa, near San Carlos de Río Negro). Endemic. ◆Fig. 444. Palmorchis pubescens Barb. Rodr., Gen. Spec. Orchid 1: 170. 1877. Terrestrial 40–100 cm tall; flowers greenish. Rain forests, 100–700 m; Bolívar (La Escalera to Cerro Venamo region, Río Ayaicha, Río Chicanán). Trinidad-Tobago, Amazonian Brazil. ◆Fig. 442. The Venezuelan material is somewhat different from the Brazilian and might represent an undescribed variety of Palmorchis pubescens or another species.

Palmorchis

Fig. 442. Palmorchis pubescens

Fig. 443. Palmorchis guianensis

Fig. 444. Palmorchis puber

493

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O RCHIDACEAE

104. PAPHINIA Lindl., Edwards’s Bot. Reg. 29: misc. 14. 1843. [Subtribe Stanhopeinae]. by Gustavo A. Romero-González Epiphytes or rarely terrestrial herbs, cespitose. Stem modified into ovoid, compressed, sulcate and/or angulate pseudobulbs, of 1 internode, when young entirely concealed by distichous, scarious sheaths, eventually naked. Leaves 1–3, narrowly ovate, acute, plicate, articulate, sometimes subpetiolate. Inflorescences 1–several, lateral, arising from the base of the pseudobulbs, erect to pendent, racemose, fewflowered. Flowers resupinate or not, short-lived, showy; floral bracts concave, narrowly ovate to obovate, shorter than the pedicellate ovary. Sepals and petals membranous, spreading or erect-spreading; sepals concave, narrowly ovate, acuminate, lateral ones basally connate or not, laterally adnate to the column foot, forming a short mentum; petals similar to the sepals but generally shorter and narrower. Lip clawed, 3-lobed, adnate to the apex of the column foot, the lateral lobes erect, narrowly ovate, falcate, sometimes with a pilose tooth on the inner margin, the midlobe triangular to sagittate, sometimes apically concave; disk with an axial, fleshy callus and glandular trichomes between and often extending to the midlobe margins. Column elongate, semiterete, clavate, apically auriculate, sometimes produced into a short to long foot; anther terminal, operculate, incumbent, 1-locular or imperfectly 2-locular; pollinarium with 2 yellow pollinia, narrowly obovate, cleft, cartilaginous, and dorsoventrally flattened, the tegula long, linear, the viscidium subtriangular. Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 10 species, 3 in Venezuela, all in the flora area. Some specimens collected in the upper Río Cataniapo, Amazonas state, are intermediate between Paphinia cristata and P. lindeniana, and probably represent natural hybrids between these 2 species. Key to the Species of Paphinia 1. 1. 2(1). 2.

Plants terrestrial, inflorescence erect ..................................... P. dunstervillei Plants epiphytic, inflorescence pendent .................................................... 2 The anterior margin of the lip lateral lobes without a pilose, triangular tooth, base of the column wings sagittate ................................... P. cristata The anterior margin of the lip lateral lobes with a pilose, triangular tooth, base of the column wings rounded ........................................ P. lindeniana

Paphinia cristata (Lindl.) Lindl., Edwards’s Bot. Reg. 29: misc. 14. 1843. —Maxillaria cristata Lindl., Edwards’s Bot. Reg. 21: t. 1811. 1836. Epiphyte; inflorescence short, pendent, 1– 3-flowered; flowers reddish brown with white markings. Wet forests, 50–1000 m; Delta Amacuro (Río Barima), Bolívar (Cerro Venamo, lower Río Churún near Guarimba, Río Erebato, Uei-tepui), Amazonas (Río Cataniapo, Río Sipapo). Coastal Cordillera of

Miranda; Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana. Paphinia dunstervillei Dodson & G.A. Romero, Amer. Orchid Soc. Bull. 62: 899. 1993. Terrestrial; inflorescence erect, to 6-flowered; flowers purple-brown. Caatinga forests, ca. 100 m; Amazonas (Caño Ucata southeast of Síquita, near Cerro Autana, road between Yavita and Maroa). Endemic.

Pelexia 495

Paphinia lindeniana Rchb. f., Flora 70: 497. 1887. —Lycaste lindeniana (Rchb. f.) G. Nicholson, Dict. Gard. 2: 304. 1888. Epiphyte; inflorescence pendent, to 8flowered; flowers purple-brown with white markings. Wet forests, 50–600 m; Amazonas (Caño Ucata southeast of Síquita, Cerro Yapacana, Río Cataniapo, Río Putaco, Río Siapa, Sierra de la Neblina, road between Yavita and Maroa). Colombia, Peru, Brazil (Amazonas). ◆Fig. 445.

Fig. 445. Paphinia lindeniana

105. PELEXIA Poit. ex Lindl., Bot. Reg. 12: sub t. 985. June 1826, nom. cons. [Subtribe Spiranthinae]. Spiranthes Rich., De Orchid. Eur. 20, 28, 36. Aug.–Sept. 1817, nom. cons., pro parte. by Germán Carnevali, Ivón M. Ramírez-Morillo, and Carlos A. Vargas Terrestrial, humicolous, rarely subepiphytes or subaquatic. Roots fasciculate, fleshy, stipitate to fusiform. Leaves convolute, subfleshy, basal, petiolate, rarely withered during anthesis. Inflorescence erect, variously bracteate, terminated by a loosely to densely, many-flowered spike; floral bracts small to large; ovary cylindric to ± fusiform, sessile to subsessile, somewhat twisted. Flowers small to more often medium-sized to medium-large, resupinate. Sepals unequal, usually spreading; dorsal sepal concave to cucullate, together with petals form a distinct helmet-like structure (galea); lateral sepals ± porrect, decurrent on the protruding column foot, basally connate into a ventricose, saccate or spur-like vesicle which is often manifested in a pronounced mentum; petals connivent with dorsal sepal, decurrent or variously oblique at base. Lip usually fleshy, conspicuously clawed, with 2 glands at its base, rarely auriculate, lip lateral margins joined with sides of the column, blade entire or more frequently constructed about apical 1/4–1/3. Column rather stout, elongate, puberulent or pilose in front, basally produced in a decurrent foot which often protrudes from the wall of the ovary; anther ovate-cordate, obtuse; pollinia 4, soft, clavate with a thick, ovate to suborbicular viscidium; rostellum rather soft, pliable laminar, strap-shaped, linear-oblong to ligulate, obtuse or truncate; stigmas 2, free to confluent. U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Para-

496

O RCHIDACEAE

guay, northern Argentina, Uruguay; ca. 70 species, 4 in Venezuela, 2 of these in the flora area. This genus contains usually forest-floor, shade-loving plants. Key to the Species of Pelexia 1. 1.

Basal leaves many; leaves ca. 10 × 3.5 cm; lip with a pair of calli at base of terminal lobe ............................................................................... P. callifera Basal leaves 2 or 3; leaves ca. 20 × 10 cm; lip without a pair of calli at base of terminal lobe ................................................................................. P. laxa

Pelexia callifera (C. Schweinf.) Garay, Bot. Mus. Leafl. 28: 343. 1980. —Spiranthes callifera C. Schweinf., Bot. Mus. Leafl. 20: 1. 1962. Terrestrial or muscicolous herb, erect; inflorescences to 25 cm tall, 10–15-flowered in the apical 1/2; flowers not widely opening, perianth segments tan. Lower montane forests, 200–500 m; Bolívar (near Apoypué, lower slopes of Macizo del Chimantá). Suriname, Ecuador. ◆Fig. 446. Pelexia laxa (Poepp. & Endl.) Lindl., Gen. Sp. Orchid. Pl. 482. 1840. —Stenorrhynchos laxus Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 7, t. 109. 1837. —Spiranthes laxa (Poepp. & Endl.) C. Schweinf., Bot. Mus. Leafl. 10: 29. 1941. Pelexia maculata Rolfe, Kew Bull. 1893: 7. 1893. —Spiranthes maculata (Rolfe) C. Schweinf., Bot. Mus. Leafl. 10: 30. 1941. Terrestrial to 65 cm tall, shade-loving; leaves often variegated with white spots; flowers greenish or whitish, sepals often with purplish tips. Rain forests, 200–300 m; Bolívar (Serranía de Imataca). Aragua, Carabobo, Distrito Federal, Falcón, Monagas. Ecuador, Peru.

Fig. 446. Pelexia callifera

Peristeria 497

106. PERISTERIA Hook., Bot. Mag. 58: t. 3116. 1831. [Subtribe Stanhopeinae]. by Gustavo A. Romero-González Cespitose, robust epiphytes, or terrestrial (outside the Venezuelan Guayana) herbs. Stem modified into ovoid to subcylindric pseudobulbs, of 1 internode (often second and third internodes are also conspicuous). Leaves 1–5, narrowly to widely ovate, acute, plicate, articulate. Inflorescences 1–several, lateral, arising from the base of the pseudobulbs, erect or pendent (short and pendulous in species from the flora area), racemose, few- to many-flowered. Flowers globose, fleshy, waxy, resupinate, often showy; floral bracts concave, ovate, shorter than the pedicellate ovary. Sepals fleshy, concave, spreading, elliptic to very broadly ovate, the lateral sepals partially connate or not; petals similar to but generally smaller than the sepals. Lip fleshy, subsessile, adnate to the apex of the column foot, conspicuously clawed, divided into a hypochile and an epichile; hypochile deeply concave or not, with or without a central, fleshy callus, winged or auriculate or not, the margins keeled or not; the epichile articulate to the apex of the hypochile, inflexed or incumbent, the margins erect, spreading or reflexed, the upper surface smooth or often with 2 lateral keels and/or a low, fleshy, sulcate callus. Column short, erect, semiterete, apically winged to auriculate, sometimes produced into a conspicuous foot; anther terminal, operculate, incumbent, imperfectly 2-locular; pollinarium with 2 yellow pollinia, very narrowly obovate, cleft, cartilaginous, and dorsoventrally flattened, the tegula very short, transversely elliptic, the viscidium subcircular, sometimes hooked. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; 7 species, 4 in Venezuela, 3 of these in the flora area. A male euglossine bee (Euglossa analis Westwood group) captured in Amazonas state (Río Cataniapo) carried a pollinarium of a Peristeria species as yet unknown from the flora area. The pollinarium indicates flowers smaller than those of the species presently known from the flora area. Key to the Species of Peristeria 1. 1. 2(1). 2.

Column not winged or auriculate, margin of mesochile finely serrate, epichile covered with minute tubercles ........................................ P. cerina Column winged or auriculate, margin of lip mesochile entire, epichile smooth .................................................................................................... 2 Flowers yellowish orange spotted with maroon; column auriculate, auricles < 2 mm long ........................................................................ P. aspersa Flowers pale greenish white spotted with maroon; column winged, wings > 4 mm long ................................................................................. P. pendula

Peristeria aspersa Rolfe, Lindenia 6: 57, t. 267. 1891. Epiphyte; inflorescences to 5-flowered; flowers showy, yellow or yellowish orange spotted with maroon. Wet forests, 50–1500 m; Bolívar (Cerro Guaiquinima), Amazonas (Caño Coromoto, Cerro Marahuaka, upper

Río Cataniapo, Río Siapa). Brazil (Amazonas). ◆Fig. 447. Peristeria cerina Lindl., Edwards’s Bot. Reg. 23: t. 1953. 1837. Peristeria guttata Knowles & Westc., Fl. Cab. 2: 99, t. 70. 1838.

498

O RCHIDACEAE

Epiphyte; inflorescences to 10-flowered; flowers showy, pale salmon spotted with maroon. Wet forests, 200–1000; Bolívar (upper Río Cuyuní, Sierra de Lema). Coastal Cordillera of Miranda, Apure (Reserva Forestal San Camilo); Colombia, Guyana, Peru. Peristeria pendula Hook., Bot. Mag. 63: t. 3479. 1836. Epiphyte; inflorescences to 6-flowered; flowers pale greenish white spotted with maroon. Wet forests, ca. 500 m; Bolívar (Río Apacará), Amazonas (near Maroa). TrinidadTobago, Guyana, Ecuador, Peru, Brazil (Amazonas, Pará).

Fig. 447. Peristeria aspersa

107. PHRAGMIPEDIUM (Pfitzer) Rolfe, Orchid. Rev. 4: 330. Nov. 1896, nom. cons. —Paphiopedilum sect. Phragmopedilum Pfitzer, Bot. Jahrb. Syst. 19: 41. 1894. [Subfamily Cypripedioideae]. Uropedium Lindl., Orchid. Linden. 28. Nov.–Dec. 1846, nom. rej. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrials, muscicolous, epiphytes, lithophytes, or semiaquatics, erect. Rhizome short or creeping. Roots fibrous, very elongate. Stem reduced or absent. Leaves conduplicate, nonarticulate with the sheathing, imbricate base, distichous; blades coriaceous, linear to strap-shaped. Inflorescences terminal, racemes or panicles, usually successively several-flowered; peduncle and rachis terete, mostly pubescent, distantly or imbricately bracted; floral bracts conspicuous, shorter to ± equal to the ovaries, rarely longer; pedicels short; ovary cylindric, 3-locular, placentation axillary. Flowers resupinate, showy, lasting several days. Perianth articulate with ovary; segments spreading, subfleshy, usually variously pubescent; dorsal sepal free, ± concave; lateral sepals connate into a synsepal subsimilar to dorsal sepal, usually hidden by the lip; petals free, narrower than sepals and frequently with a meristematic apex and elongating with age (to 75 cm in one species). Lip sessile, calceolate, inflated with involute margins, with basal and apical openings due to differences in the degree of involution of medial margins, in one species undifferentiated and similar to the petals (peloric form). Column short, stout, with 2 laterally fertile globose, 2-celled anthers, the third, medial anther modified into a variously shaped, shield-like staminode, flat or ± convex; pollen in monads, united in soft pollinia; filaments largely united with style; fertile stigmas 3, confluent into a suborbicular or dome-like stigmatic surface. Capsules fusiform. Widespread in Neotropics (excluding the West Indies), most diverse in the Andean countries; ca. 12 species, 3 in Venezuela, 2 of these in the flora area.

Phragmipedium

499

Key to the Species of Phragmipedium 1. 1.

Plants long-creeping; leaves 4–12 mm wide; petals acuminate, 4.7–10 cm long ................................................................................... P. klotzschianum Plants cespitose; leaves 3.5–8 cm wide; petals rounded, 3.5–6 cm long .............................................................................................. P. lindleyanum

Phragmipedium klotzschianum (Rchb. f.) Rolfe, Orchid Rev. 4: 332. 1896, “Phragmopedilum klotzschianum.” — Cypripedium klotzschianum Rchb. f., Linnaea 22: 811. 1848. Terrestrial or more often semiaquatic; leaves 10–35 cm long; inflorescences racemes, successively few-flowered; flowers with dorsal sepal 3–3.5 cm long, pale magenta striped with dark magenta; synsepal greenish with dull maroon tinges, petals streaked with dull magenta; lip greenish, inner margin white with dull lavender spots; column greenish with dark lavender on each side. Locally common, forming colonies in rocky stream margins, often partially submerged several months a year, (400–)900– 1400 m; Bolívar (basins of the upper Río Caroní and Río Paragua). Amazonian Brazil, Guyana. ◆Fig. 448.

Fig. 448. Phragmipedium klotzschianum

Phragmipedium lindleyanum (R.H. Schomb. ex Lindl.) Rolfe, Orchid Rev. 4: 330. 1896. —Cypripedium lindleyanum R.H. Schomb. ex Lindl., Gen. Sp. Orchid. Pl. 53. 1830. Selenipedium kaieteurum N.E. Br., Gard. Chron. n.s. 2: 262. 1885. —Phragmipedium kaieteurum (N.E. Br.) Garay, Orchid Digest. 43: 136. 1979. Humicolous on soil or rocks; leaves 25–70 cm long, often with yellow margins; inflorescences panicles or racemes, many-flowered; flowers with dorsal sepal 2.8–4 cm long, greenish with pale brownish veins; petals pale olive green with brown-magenta or maroon veins; lip olive or brown-green with tawny brown veins. Rare or locally common in ± open places in rain or cloud forests, especially on igneous rock outcrops, (300–)800– 2400 m; Bolívar (basins of Río Caroní and Río Cuyuní), Amazonas (Cerro Sipapo, Serranía Tapirapecó). Brazil (Roraima, Amazonas, Bahia), Guyana. ◆Fig. 449.

Fig. 449. Phragmipedium lindleyanum

500

O RCHIDACEAE

108. PINELIANTHE S. Rauschert, Feddes Repert. 94(7/8): 465. 1983. —Pinelia Lindl., Fol. Orchid., Pinelia. 1853, non Pinellia Tenore 1830. [Subtribe Laeliinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Minute epiphytes, sometimes twig epiphytes, 3–10 cm tall, erect, cespitose or shortly creeping. Rhizome abbreviate, thin; pseudobulbs ellipsoid, pear-shaped to subspheric, heteroblastic (one internode), apically 1-foliate. Leaves conduplicate, fleshy, 1-veined, sessile, erect or spreading, ± equal to or a little longer than the pseudobulb. Inflorescence terminal, solitary, erect, racemose, 1- or successively fewflowered, remotely few-bracted, much longer than subtending leaf; peduncle elongate, terete; rachis straight, much shorter than the peduncle; floral bracts shorter than pedicellate ovary, tubular; pedicellate ovary clavate, glabrous. Flowers resupinate, campanulate or widely spreading, spreading, large for the size of the plant. Perianth segments subfleshy or membranous; sepals free, subsimilar, the laterals slightly oblique; petals shorter and narrower than the sepals. Lip erect, simple, larger than the other perianth segments, the basal margins connate with the base of the column wings; disk basally callose. Column relatively short, along all its length flanked by a pair of petaloid wings; anther terminal or subventral, operculate, incumbent, imperfectly 4-locular; clinandrium winged; pollinia 4, laterally compressed, caudicles poorly developed; rostellum transverse; stigma ventral. Colombia, Venezuela, Brazil; 3 or 4 species, 1 in Venezuela. Pinelianthe alticola (Garay & Dunst.) S. Rauschert, Feddes Repert. 94(7/8): 465. 1983. —Pinelia alticola Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 3: 242. 1965. Twig epiphyte 2–5 cm long, including the inflorescence; pseudobulbs pear-shaped to subspheric; leaves broadly ovate; inflorescence 1-flowered; flowers hyaline or pinkish. Cloud forests or open shrublands, 1200–1800 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region), Amazonas (Cerro Marahuaka). Endemic. ◆Fig. 450.

Fig. 450. Pinelianthe alticola

109. PLATYSTELE Schltr., Repert. Spec. Nov. Regni Veg. 8: 565. 1910. [Subtribe Pleurothallidinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes or rarely lithophytes, minute to small, cespitose to creeping, erect, usually shade-loving. Rhizome from almost absent to creeping and branching and then plants forming dense mats. Ramicauls 1-foliate, very short to relatively elongate, covered by sheaths. Leaves fleshy to thinly coriaceous, erect to prostrate, narrowly obovate to broadly ovate, basally subsessile to conspicuously pseudopetiolate. Inflorescences originating from near ramicaul apex with an annulus, raceme shortto long-pedunculate, shorter to much longer than subtending leaf; rachis usually zigzag, more rarely straight. Flowers minute to small, usually resupinate, opening in slow succession or, rarely, ± simultaneously, often with widely spreading perianth

Platystele 501

segments. Perianth segments membranaceous, often translucent, purple-tinged, yellowish to deep purple, usually 1-veined, free. Sepals usually similar, rarely dorsal much longer, often acuminate. Petals similar to sepals but often narrower, sometimes margins ciliolate. Lip simple to rarely 3-lobed, hinged to the base of the column, often ovate to narrowly ovate, margins frequently erose or ciliolate. Column short, broad, ± membranaceous, dilated toward apex. Anther apical, incumbent, operculate. Pollinia 2, pear-shaped to almost subspherical. Stigma transverse, 2lobed, subapical. Capsules ellipsoid, crowned by the persistent perianth. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, southern Brazil, Bolivia; ca. 50 species, 7 in Venezuela, 3 of these in flora area. Platystele is closely related to Teagueia, a recent segregate. Key to the Species of Platystele 1. 1. 2(1). 2.

Plants cespitose; flowers often purple-tinged; sometimes lip wholly purple ................................................................................................... P. oxyglossa Plants creeping; rhizome branching or not, sometimes forming dense mats; flowers translucent, cream-yellow, greenish, white, or green ... 2 Dorsal sepal filiform, long-acuminate, 3–4 times longer than lateral sepals; petals acuminate .......................................................... P. johnstonii Dorsal sepal not attenuated into a filiform tails subequal to slightly longer than lateral sepals; petals obtuse to acute ................... P. ovalifolia

Platystele johnstonii (Ames) Garay, Caldasia 10: 233. 1968. —Pleurothallis johnstonii Ames, Orchidaceae 2: 271. 1908. Epiphyte, minute, creeping, shade-loving; rhizome branching so plants form dense mats; inflorescence erect, 1–3-flowered; flowers small, resupinate, successive; dorsal sepal long-filiform attenuate, perianth segments and lip greenish white or creamy yellow. Rain or cloud forests, 600–800 m; Bolívar (Altiplanicie de Nuria). Coastal Range (Miranda, Nueva Esparta). Ecuador. ◆Fig. 451. Platystele ovalifolia (H. Focke) Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 2: 268. 1961. —Stelis ovalifolia H. Focke, Natuurw. Tijdschr. 2: 202. 1849. —Pleurothallis ovalifolia (H. Focke) Rchb. f. in Walp., Ann. Bot. Syst. 6: 188. 1861. Epiphyte, minute, creeping, shade-loving; rhizome branching and forming dense mats; inflorescence erect, 2–4-flowered; flowers minute, opening widely, successive, green or

greenish white. Rain or cloud forests, 200– 600 m; Bolívar (Altiplanicie de Nuria, Serranía de Imataca). Coastal Range (Falcón, Miranda); West Indies, Guyana, Suriname, Brazil. Platystele oxyglossa (Schltr.) Garay, Orquideología 9: 120. 1974. —Pleurothallis oxyglossa Schltr., Repert. Spec. Nov. Regni Veg. 10: 354. 1912. —Pleurothallis lancilabris var. oxyglossa (Schltr.) C. Schweinf., Bot. Mus. Lefl. 6: 200. 1938.

Fig. 451. Platystele johnstonii

502

O RCHIDACEAE

Platystele lancilabris auct. non (Rchb. f.) Schltr. 1923: sensu Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 1: 299. 1959. Epiphytic, minute, cespitose, erect, shadeloving; inflorescences erect to arching; flowers minute, with widely spreading perianth segments, perianth segments translucent,

dull yellow or whitish, tinged with lavender or purple, lip often dark purple. Rain forests, 300–800 m; Bolívar (upper Río Caroní and Río Caura basins). Coastal Range (Carabobo, Miranda, Sucre), Andes (Mérida); Mexico, Guatemala, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Bravil, Bolivia.

110. PLECTROPHORA H. Focke, Tijdschr. Wis- Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 1: 212. 1848. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Twig epiphytes, erect to subpendulous, cespitose, mostly sun-loving. Rhizome very short. Pseudobulbs 1-foliate, small, laterally flattened, suborbicular to ovoid, enveloped by 2–4 imbricating sheaths of which innermost 1 or 2 have foliar blades. Leaves laterally flattened, articulate, ensiform, usually linear-oblong or somewhat falcate. Inflorescences 1–4, lateral, originating from the axils of the sheaths which enfold the pseudobulb, 1–3-flowered, much shorter than the leaves; floral bracts acute, inconspicuous; pedicellate ovary terete or obtusely 3-edged, longer than sepals. Flowers relatively large for the plant size. Perianth segments fleshy, greenish, yellowish, or white. Sepals subequal, erect to subparallel to the column; dorsal sepal free, the laterals basally produced into a very long, narrow spur, straight to somewhat falcate; petals broader than sepals. Lip decurrent with column base, simple to shallowly lobed, broadly suborbicular to suborbicular-ovate, somewhat cucullate with the basal part funnel-shaped and enfolding the column, basally produced into 2 short to relatively long spurs included into the sepal cup and connate with it. Column short, erect, thick, semiterete, ventrally sulcate, footless, exauriculate or minutely 2-auriculate; anther terminal, operculate, incumbent, 1-locular to imperfectly 2-locular; pollinia 2, waxy, oblong to obovoid, laterally compressed; tegula linear, relatively long; viscidum small, ovate; clinandrium small, truncate; rostellum transverse; stigma ventral. Capsules 3-edged. Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 85 species, 1 in Venezuela. Plectrophora iridifolia H. Focke, Tijdschr. Wis-Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 1: 212. 1848. Plectrophora cultrifolia auct. non (Barb. Rodr.) Cogn. 1904: sensu Garay & Dunst. in Dunst & Garay, Venez. Orchid. Ill. 1: 303, fig. 793. 1959; Foldats in Lasser, Fl. Venez. 15(5):83. 1970. Twig epiphyte, sun-loving; leaves 4–7 cm long; flowers relatively showy; sepals 9–13 mm long, not widely spreading, white or greenish white; petals parallel to column, white; lip 14–18 mm long, funnel-shaped, white with yellow or orange stripes or longitudinally arranged dots, spur white or greenish, 20–25 mm long. Rain forests or open shrublands, often growing close to rivers or

Fig. 452. Plectrophora iridifolia

Pleurothallis 503

streams, 50–500 m; Bolívar (near Río Canaracuni, Río Uaiparú), Amazonas (Río Sipapo,

Yavita to Maroa road). Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 452.

111. PLEUROTHALLIS R. Br. in W.T. Aiton, Hortus Kew. ed. 2, 5: 211. 1813. [Subtribe Pleurothallidinae]. Acronia C. Presl, Reliq. Haenk. 1: 104. 1827. Specklinia Lindl., Gen. Sp. Orchid. Pl. 8. 1830. Physosiphon Lindl., Edwards’s Bot. Reg. 21: sub t. 1797. 1836. Acianthera Scheidw., Allg. Gartenzeitung 10: 292. 1842. Rhynchopera Klotzsch in Link, Klotzsch & Otto, Icon. Pl. Rar. 2: 103. 1844. Crocodeilanthe Rchb. f. & Warsc., Bonplandia (Hanover) 2: 114. 1854. Anathallis Barb. Rodr., Gen. Spec. Orchid. 1: 23. 1877. Cryptophoranthus Barb. Rodr., Gen. Spec. Orchid. 2: 80. 1882. Otopetalum F. Lehm. & Kraenzl., Bot. Jahrb. Syst. 26: 257. 1899, non Miq. 1856. Kraenzlinella Kuntze, Lex. Gen. Phan. 310. 1903. Phloeophila Hoehne & Schltr., Arq. Bot. Sâo Paulo 1: 199. 1926. Chamelophyton Garay, Orquideología 9: 115. 1974. Anthereon Pridgeon & M.W. Chase, Lindleyana 16(4): 252. 2001. Echinosepala Pridgeon & M.W. Chase, 17(2): 100. 2002. —Echinella Pridgeon & M.W. Chase, Lindleyana 16(4): 253. 2001. by Germán Carnevali and Ivón M. Ramírez-Morillo Usually epiphytes, but often lithophytes or subterrestrial herbs, minute to relatively large. Rhizome or primary stem abbreviate to long-creeping, usually clothed by tubular, variously ornamented sheaths; roots few to many, slender to coarse or fleshy. Ramicauls erect to pendent, sometimes prolific (stems are borne on top of other stems), terete, triquetrous, laterally flattened or winged, very rarely tetragonous, unifoliate, partially or totally enclosed by tubular sheaths, sheaths rarely with trichomes; leaf erect or often spreading in relation to the ramicaul, thinly or thickly coriaceous, to terete or laterally compressed, glabrous, sometimes glaucous, light to dark green, rarely purplish, linear, orbicular, ovate, cordate, or obovate, the apex acute, obtuse to rounded, notched with a mucro in the sinus, the base petiolate or sessile, cuneate, rounded, or cordate, or decurrent on the ramicaul, the petiole sometimes twisted. Inflorescence emerging laterally with or without an annular ring (annulus) below or at the apex of the ramicaul (the leaf-stem abscission layer), or from the apex of the ramicaul without an annulus, 1-flowered, single or fasciculate, or racemose, single or fasciculate, flowering simultaneously or successively, longer or shorter than the leaf, the peduncle slender or stout, short or long, round or laterally compressed, with few to many bracts; floral bracts short or long, tubular to cucullate; pedicels slender to stout, long or short, sometimes verrucose; ovary 3-valved, deciduous, smooth, carinate to crested, verrucose, papillose to spiculate; flowers small to relatively large and showy, resupinate or not, usually lasting a few days; sepals membranous to thickly fleshy, smooth, verrucose, or pubescent, ciliate or fringed, narrow to transverse, petals usually smaller than the sepals, always free, parallel or widely spreading to recurved; lip usually fleshier than the other perianth segments, 1–5-lobed, acute, acuminate, or obtuse, the base variously articulated with the base of the column or apex of the column foot, sometimes inflexibly adnate; column semiterete, long or short, slender to stout, winged or wingless, toothed or without teeth, the anther apical to ventral, hooded or exposed, pollinia 2,

504

O RCHIDACEAE

rarely 6, pear-shaped to spherical, naked or with caudicles, with or without a detachable viscidium, the stigma apical to ventral, 1- or 2-lobed, the base of column developed or not developed into a column foot with the apex of the ovary, the tip of the foot sometimes elongated beyond the ovary. U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina; ca. 2000 species, ca 160 in Venezuela, 70 of these in the flora area. As here circumscribed, Pleurothallis is probably the largest of the neotropical genera of the Orchidaceae. It is most diverse in the Andean countries and in southern Central America. As presently understood, this extremely large and heterogeneous assemblage of species is probably para- or polyphyletic, even after the segregation of some discordant groups during the late 1970s and early 1980s (e.g., Platystele Schltr., Myoxanthus Poepp. & Endl., Trichosalpinx Luer; C. Luer. Monogr. Syst. Bot. Missouri Bot. Gard. 15. 1986). The genus has been subdivided into many subgenera (C. Luer. Monogr. Syst. Bot. Missouri Bot. Gard. 20. 1986), some of which will probably be recognized as distinct genera after phylogenetic studies. The subtribe Pleurothallidinae has recently been under intense phylogenetic scrutiny. We have long been aware of the diffuse boundaries between some of the genera recognized until recently and that new, carefully conducted phylogenetic analysis would require a generic realignment of the subtribe. Such phylogenetic studies, both morphological (Neyland, R.L., E. Urbatsch, and A. M. Pridgeon. Bot. J. Linn. Soc. 117: 13–28. 1995), and more recently, DNA-based (Pridgeon, A.M., and M. W. Chase. Lindleyana 16: 235–271. 2001; Pridgeon, A.M., R. Solano, and M.W. Chase. Amer. J. Bot. 88: 2286–2308. 2001) have recently been published. The results of these DNA-based studies provide a new framework of hypotheses of relationships and are congruent with the expectations of a necessity for generic realignments. However, the authors of the above mentioned papers hastened to produce long lists of new nomenclature, involving both new genera and new combinations. In doing so, several clusters of species, previously referred to other genera (e.g., some of the subgenera of Pleurothallis sensu Luer. Monogr. Syst. Bot. Missouri Bot. Gard. 20. 1986), were lumped into large, morphologically undiagnosable generic concepts (e.g., Stelis sensu Pridgeon & Chase 2001). In most of the cases, alternative classifications, involving the circumscription of still monophyletic but smaller, diagnosable generic entities were possible and desirable for floristic and identification purposes. Since only a fraction of this huge subtribe has been sampled and no total evidence analyses (involving molecular, morphological, and other characters) have yet been conducted, it is likely that some of the fine details of the phylogeny topology will change, which would result in yet more nomenclatural changes. Thus, we have decided not to follow this system for the time being, albeit recognizing that it has its merits and that it could provide the foundation for a future system that would hopefully recognize diagnosable generic entities. The synonymies under the several species of Pleurothallis treated in this account indicate their generic placement in the Pridgeon and Chase system. Some of the generic taxa proposed in the Pridgeon and Chase system should eventually be recognized (e.g., Acianthera, Anathallis, Specklinia, etc.), and a future publication on the Pleurothallidinae of the Venezuelan Guayana will deal with this problem since many new combinations would be required.

Pleurothallis 505

Key to the Species of Pleurothallis 1.

Stems either triquetrous (triangular in cross section) or sharply compressed, at least in the apical half, always conspicuous ...................... 2 1. Stems neither strongly flattened nor triquetrous, either conspicuous or almost absent ......................................................................................... 9 2(1). Rhizome abbreviate; stems aggregate; leaves 1/2 or less the length of the stem, cuneate to rounded basally; petals elliptic, truncate ............................................................................................... P. sarcosepala 2. Rhizome elongate; stem distanced; leaves as long as or longer than the stem, when 1/2 or less the length of the stems, then basally cordate; petals various .............................................................................................. 3 3(2). Stems sharply compressed in the apical half, rigidly arched at the apex; lip with fimbriate margins and lateral lobes of the lip ......................... 4 3. Stems triquetrous, not arched apically; lip with ± entire margins, either without lobes or these are not fimbriate ............................................... 5 4(3). Inflorescences 1-flowered; stems about as long as or shorter than the leaves; leaves deeply concave; dorsal sepal linear; lip with 2 basal truncate, retrorse lobes ......................................................P. phoenicoptera 4. Inflorescences several-flowered; stems conspicuously longer than leaves; leaves flat or shallowly concave; dorsal sepal elliptic; lip with 2 basal fimbriate lobes ........................................................................... P. prolifera 5(3). Petals truncate or rounded; stem 3.5–10 cm long; leaves 4–5 cm long; plants usually from lowlands below 300 m, rarely reaching 1200 m ................................................................................................................ 6 5. Petals ovate-elliptic or narrowly obovate, acute to rounded; stem to 3.5 cm long; leaves to 4 cm long; plants usually from rain or cloud forests above 850 m ........................................................................................... 7 6(5). Lip obovate, totally lobeless; synsepal 9–11 mm long .............. P. discophylla 6. Lip elliptic, basally with very small, erect, rounded lobes; synsepal 6– 6.5 mm long ......................................................................... P. sandaliorum 7(5). Lip simple; flowers entirely greenish yellow; petals 1-veined ....... P. morilloi 7. Lip 3-lobed; flowers with purple or maroon stripes; petals 3-veined. ...... 8 8(7). Stems sharply triquetrous, i.e., with dorsal face keeled, thickened toward apex; leaves cordate to subcordate basally, usually concave; dorsal sepal at least 8 mm long; central lobe of lip ovate, broader toward base ...................................................................................................... P. garciae 8. Stems subtriquetrous, i.e., with dorsal face rounded or at least not keeled; leaves basally rounded to subtruncate, flat; dorsal sepal to 7 mm long; central lobe of lip elliptic or elliptic-obovate; broader toward middle or apex ................................................................................... P. yauaperyensis 9(1). Inflorescences strictly 1-flowered, either solitary or fasciculate ............ 10 9. Inflorescences 2–several-flowered; flowers can be successive and appearing one at a time but scars on rachis will reveal the multiflowered nature of inflorescence ............................................................................. 27 10(9). Stems somewhat distanced on a creeping rhizome ................................ 11 10. Stems approximate; plants cespitose ...................................................... 18 11(10). Flowers originating from the rhizome; dorsal sepal apically connivent

506

O RCHIDACEAE

11.

12(11). 12.

13(12).

13. 14(13).

14. 15(13).

15. 16(15). 16. 17(16). 17. 18(10). 18. 19(18). 19. 20(18).

20. 21(20).

21.

22(20).

with the apex of the synsepal; capsules red, hairy ............. P. lappiformis Flowers originating at or near the apex of the stem; dorsal sepal not connivent with the apex of the synsepal, usually spreading; capsules usually green, not hairy ............................................................................. 12 Leaves rugulose on the upper surface, appressed to substrate, much longer than the inconspicuous stems; pollinia 6 ...................... P. hexandra Leaves smooth on the upper surface, erect or pendulous, not appressed to substrate, stems longer or about as long as the leaves, never inconspicuous; pollinia 2 .............................................................................. 13 Plants large, coarse, with stems 8–20 cm long; external surface of the sepals, pedicel, ovary, and floral bract hairy or scabrous with short, thick hairs in small mounds ......................................................................... 14 Plants smaller, more delicate, stems 2–6 cm long; external surface of sepals, pedicel, ovary, and floral bract glabrous .................................... 15 Leaves conspicuously shorter than the stems, basally rounded or broadly obtuse, very rarely subacute; plants terrestrial over sandy soils or lithophytic over sandstone in tepui shrublands ...................... P. arenicola Leaves as long or longer than the stem, basally acute or obtuse; plants epiphytic in cloud forests ..................................................... P. aspasicensis Leaves 10–14 mm wide, often pendulous; plants growing on shaded granitic rocks, rarely epiphytic; lip basally with 2 small, retrorse auricles ................................................................................................... P. granitica Leaves 4–10 mm wide, usually erect; plants growing as epiphytes, rarely on rocks; lip without basal auricles ..................................................... 16 Dorsal sepal about 3 times as long as wide; surface of lip ± smooth; flowers cleistogamous, developing into triquetrous fruits ..................... P. wilsonii Dorsal sepal at least 5 times as long as wide; surface of lip verruculose or pubescent; flowers not cleistogamous ................................................. 17 Lip covered with narrow, flattened hairs .................................. P. longisepala Lip tuberculate ........................................................................... P. miqueliana Leaves basally attenuated into a pseudopetiole ..................................... 19 Leaves basally rounded to cordate; rarely broadly cuneate ................... 20 Lip basally with 2 linear, falcate, antrorse lobes; dorsal sepal 14–22 mm long .......................................................................................... P. hitchcockii Lip without basal lobes; dorsal sepal 5–7 mm long ..................... P. ruscifolia Stems 2.7–9(–11.5) cm long; dorsal sepal to 6 mm long, usually 2.5–5 mm long; leaves basally rounded to shallowly cordate, even in the mature stages .................................................................................................... 21 Stems (10–)15–40 cm long; dorsal sepal (4.5–)7–12 mm long; leaves deeply cordate at base when fully mature .......................................... 22 Inflorescences usually fasciculate, several at a time; dorsal sepal 1.2– 1.4 times as long as wide; lip acute, deeply cordate basally; petals about 3 times as long as wide .............................................................. P. ionantha Inflorescences usually solitary; dorsal sepal at least 2 times as long as wide; lip apically truncate to rounded, broadly cordate to subtruncate basally; petals 4–6 times as long as wide ................................ P. tridentata Dorsal sepal 6–9 mm long, acute or shortly acuminate; lip oblong to ovateoblong, with a papillose or subscabrous surface ................... P. coriacardia

Pleurothallis 507

22. 23(22).

23.

24(23).

24.

25(24).

25.

26(25).

26.

27(9). 27. 28(27).

28.

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

31.

Dorsal sepal to 7 mm long, usually 4–5.5 mm long, obtuse to acute; lip ovate with a smooth to rugulose surface ............................................. 23 Lip very fleshy with a coarsely rugose surface, suggesting a geographic surface, 1.8–2 mm long; petals 0.3–0.5 mm wide, dorsal sepal 4–4.5 mm long, leaves strongly deflexed on the stem ......................... P. geographica Lip coriaceous with a smooth or minutely verruculose surface, petals 0.8– 1.2 mm wide, dorsal sepal 5–6(–7) mm long; leaves perpendicular to erect on the stem, rarely deflexed ....................................................... 24 Base of lip with a well-developed, semilunate or ovate glenion with a truncate callus in its center, basally deflexed and dorsally developed into a mentum; petals straight ................................................... P. callifera Base of lip ecallose with a concavity surrounded apically by a pair of flaps, the dorsal surface not produced into a mentum; petals usually decurved ................................................................................................... 25 Lip oblong-lanceolate or oblong in general outline, surface minutely but densely and conspicuously scabrous or hispid, the dorsal surface with a prominent subapical keel; dorsal sepal (8–)10–15(–23) mm long, narrowly lanceolate or narrowly elliptic, acute to subacuminate ................................................................................................ P. coriacardia Lip ovate or ovate-triangular, surface smooth or microscopically verruculose, the dorsal surface without a subapical keel; dorsal sepal 4– 8 mm long, elliptic or broadly elliptic, obtuse to subacute ................. 26 Lip flat or somewhat convex, apically obtuse to rounded, surface microscopically verruculose; petals with erose or denticulate margins; dorsal sepal 6–8 mm long; flowers resupinate ............................... P. archidiaconi Lip concave, apically acute, surface smooth; petals with smooth, entire margins; dorsal sepal 4–5.5 mm long; flowers apparently nonresupinate ............................................................................................ P. stenocardium Inflorescence subtended by a relatively large spathe, 1–7 cm long ....... 28 Inflorescences naked or subtended by a small spathe, < 0.9 cm ............ 32 Spathe relatively long, 2–7 cm, linear, about 1/2 the length of the subtending leaf; lateral sepals totally connate into an acuminate or arched synsepal; petals acuminate ............................................. P. loranthophylla Spathe relatively short, 1–2.5(–3) cm long, < 1/2 the length of the subtending leaf; lateral sepals not connate into a synsepal; petals obtuse to rounded ................................................................................................ 29 Petals broadly elliptic to suborbicular, 3-veined; lip simple; plants from elevations above 1800 m, usually above 2200 m ....................... P. moritzii Petals oblong to narrowly elliptic, 1-veined; lip 3-lobed; plants from lower elevations between 800–1800 m .......................................................... 30 Flowers erect to horizontal; dorsal sepal 1-veined; lateral lobes of the lip acute or obtuse; lip basally with a pulvinate callus .............. P. floribunda Flowers nodding; dorsal sepal 3-veined; lateral lobes of lip rounded, lip basally with a transverse blade-like callus ......................................... 31 Plants 35–70 cm tall; leaves elliptic to broadly ovate-elliptic, 2–3 times as long as wide; inflorescences several in anthesis simultaneously; synsepal narrower to about as broad as the dorsal sepal .......... P. galeata Plants 20–30 cm tall; leaves narrowly oblong elliptic to linear-elliptic, 6–

508

O RCHIDACEAE

32(27).

32.

33(32). 33.

34(33). 34. 35(34). 35.

36(35). 36. 37(34). 37. 38(37). 38. 39(32). 39. 40(39).

40. 41(40).

41.

42(40).

18 times as long as wide; inflorescences usually, solitary, successive; synsepal as broad as the dorsal sepal ..................................... P. tepuiensis Plants very small, long to shortly creeping, with broadly elliptic, ovateelliptic, or suborbicular leaves, 5–10 mm long, appressed to substrate; stem very short, 1–3 mm long ............................................................. 33 Plants small to large, cespitose to creeping but leaves never appressed to substrate; leaves usually > 10 mm; stem usually > 15 mm (shorter in P. parvifolia and a few others) ................................................................ 39 Inflorescences 5–10 times longer than subtending leaf; peduncle filiform; rhizome long-creeping, internodes usually longer than leaves ..... P. kerrii Inflorescences to 2 times longer than subtending leaf, usually shorter; peduncle ± stout; rhizome shortly but definitely creeping, internodes shorter to about as long as the leaves ................................................. 34 Lateral sepals connate for 3/4 or more of their length ............................. 35 Lateral sepals free or connate in their basalmost 1/4; inflorescences shorter or about as long as the leaves ................................................. 37 Leaves rugulose on the upper surface; inflorescences shorter than subtending leaves, apparently 1-flowered; lip 3-lobed .................. P. hexandra Leaves with smooth upper surface but with thickened margin (marginate); inflorescences longer than subtending leaves, successively several-flowered; lip simple ...................................................................... 36 Lip to 2.6 mm long, 4–4.5 times as long as wide, margins ciliate, apex acute .......................................................................................... P. hilariana Lip to 3–3.2 mm long, 2.9–3.7 times as long as wide, margins glabrous, apex obtuse to rounded ........................................................ P. steinbuchiae Petals obovate to elliptic-obovate, broader than sepals; apical portion of lip semiterete ............................................................................ P. nanifolia Petals elliptic or ovate-elliptic, acuminate, about as broad as the sepals; apical portion of lip flat, recurved ....................................................... 38 Lip narrowly obovate, apex rounded and recurved ..................... P. deborana Lip narrowly elliptic or narrowly ovate-elliptic, apex acute, flat ......... .................................................................................................. P. pemonum Rhizome elongated; plants creeping ........................................................ 40 Rhizome abbreviate; plants cespitose ..................................................... 43 Leaves elliptic, narrowly elliptic, or oblanceolate, at least 8 mm wide; inflorescences much shorter than subtending leaf; lateral sepals connate for > 1/2 their length into a concave synsepal ...................................... 41 Leaves linear-oblong or linear-elliptic, to 5 mm wide (usually narrower); inflorescences longer than subtending leaf; lateral sepals free ......... 42 Leaves 5.5–9.5 cm long; inflorescences simultaneously 2–7-flowered; lip broader in the apical 1/2, its anterior margin erose-laciniate; plants epiphytes ................................................................................ P. erebatensis Leaves 3–6 cm long; inflorescences successively 1-flowered; lip broader in the basal 1/2, its anterior margin smooth or finely erose; plants lithophytes on granitic boulders ...................................................... P. granitica Inflorescences at least 4 times longer than subtending leaf (generally 5 times or more); petals 0.15 mm wide; lip only slightly 3-lobed ................................................................................................. P. spiculifera

Pleurothallis 509

42. 43(39). 43. 44(43). 44. 45(44). 45.

46(44). 46. 47(46). 47. 48(47).

48.

49(48).

49.

50(49). 50. 51(50). 51.

52(47). 52. 53(52).

53.

Inflorescences to 3 times longer (usually shorter) than subtending leaf; petals 0.5–1 mm wide; lip sharply 3-lobed .............................. P. trinitensis Inflorescences shorter than stem ............................................................ 44 Inflorescences longer or about as long as subtending stem ................... 54 Lateral sepals totally free or connate at their very base ........................ 45 Lateral sepals connate for > 1/2 their length ........................................... 46 Adult leaves 2–5 × 1–1.5 cm; petals narrowly ovate-elliptic, acuminate; plants from elevations below 300 m ........................................... P. brevipes Adult leaves 5.5–15 × 2.5–9 cm; petals cuneate-spathulate, subtruncate to broadly rounded; plants from elevations of (700–)1200–2000 m ........................................................................................................ P. imraei Leaves 1–5 × 0.4–0.9 cm; stem wiry .............................................. P. pruinosa Leaves 4–27 × 1.4–9 cm; stem stout ........................................................ 47 Synsepal apically bifid or connate to 3/4–4/5 of its length ........................ 48 Synsepal entire (it can become bifid if forcefully flattened) ................... 52 Lip 3-lobed; petals apically rounded; flowers totally creamy-yellow; plants coming from elevations above 1,500 m; stem basally enveloped by several, large, loose, papery sheaths, which sometimes enclosed more than one stem .................................................................................P. velaticaulis Lip simple or subtrilobed; petals acute or obtuse; flowers orange or purplish with dots, zones, or blotches of others colors or entirely purple; stem not enveloped by large papery sheaths; plants coming from elevations below 850 m (at least in the flora area) ..................................... 49 Inflorescences about as long as the subtending leaf or longer, erect or horizontal; flowers dull to bright orange or yellowish or greenish orange; petals 3-veined, irregularly dentate to laciniate; dorsal sepal 8– 15 times as long as wide ............................................................ P. lanceana Inflorescences about 1/2 the length of subtending leaf, nodding to arching; flowers basically purple with orange at apex of sepals; petals 1-veined, entire or minutely erose; dorsal sepal 3–4 times as long as wide ...... 50 Lip not clawed, elliptic-obovate; peduncle of inflorescence almost null; apices of free portions of synsepal acute ............................... P. erebatensis Lip clawed, ovate above the claw; peduncle of inflorescence conspicuous; apices of free portions of synsepal rounded to obtuse ........................ 51 Dorsal sepal acute, shorter than the synsepal; lip acute at apex, with erect, broadly rounded sides below the middle .............................. P. fockei Dorsal sepal obtuse to subacute, longer than the synsepal; lip broadly rounded at apex, with retrorse marginal lobes below the middle ....................................................................................................... P. omissa Petals 3-veined, narrowly ovate-elliptic; inflorescences 5–8-flowered, pendulous .................................................................................... P. elvirana Petals 1-veined, linear or linear-obovate; inflorescences (5–)8–30-flowered, erect to arching ........................................................................... 53 Inflorescences erect, multiflowered; flowers yellow or yellowish cream to greenish, sometimes with red or purple spots; petals linear-obovate; lip subsimple; synsepal obtuse ........................................................ P. revoluta Inflorescences arching, (5–)7–9-flowered; flowers deep maroon or wine red; petals linear; lip 3-lobed; synsepal acute ........................... P. suspensa

510

O RCHIDACEAE

54(43). Lateral sepals totally free ........................................................................ 55 54. Lateral sepals totally or partially connate into a synsepal .................... 62 55(54). Leaves 90–120 mm long; ovary densely muricate; petals 7.6–9 mm long, basally auriculate ....................................................................... P. erinacea 55. Leaves to 45 mm long (rarely to 95 mm in P. sclerophylla); ovary smooth; petals to 6.5 mm long, without basal auricles .................................... 56 56(55). Stems 30–150 mm long; leaves (10–)15–38 mm wide; petals 1/3 or less the length of the sepals; inflorescences 10–35(–50)-flowered; flowers all opening simultaneously ....................................................................... 57 56. Stems to 20 mm long; leaves 5–16 mm wide; petals 2/3 as long as or about equal to sepals; inflorescences 2–8(–15)-flowered; flowers opening in slow succession or few opening at a time ............................................ 58 57(56). Sepals oblong, obtuse; lip with well-defined lateral lobes below the middle; column acute, lacking a conspicuous beak .................. P. maguirei 57. Sepals narrowly elliptic, acute to acuminate; lip lacking well-defined lateral lobes below the middle; column long cucullate at apex ............................................................................................... P. sclerophylla 58(56). Sepals caudate-attenuate; petals fimbriate to laciniate ......................... 59 58. Sepals acute to acuminate; petals entire to finely ciliolate .................... 60 59(58). Sepals externally glandular-ciliate; petals with few but large laciniae; inflorescences perennial, continually growing, at first erect, then arching and afterward creeping over substrate to subpendulous, becoming very elongate in age ....................................................................... P. samacensis 59. Sepals externally glabrous; petals finely and densely fimbriate at base; inflorescences erect, not perennial .......................................... P. zephyrina 60(58). Peduncle and rachis finely glandular-pubescent; petals 3–4 times as long as wide .......................................................................................... P. funerea 60. Peduncle and rachis glabrous; petals 5–8 times as long as wide ........... 61 61(60). Leaves 1.5–2 times as long as wide, concolorous; inflorescences shorter than to about as long as the leaves; petals finely erose apically; lip without any trace of lobes .............................................................. P. holstii 61. Leaves 3–8 times as long as wide, purple-spotted; inflorescences usually longer than subtending leaf; petals apically smooth; lip with 2 small erect, triangular to subrounded, lobes at the basal 1/3 ............... P. humilis 62(54). Leaves at least 20 mm wide ..................................................................... 63 62. Leaves to 15 mm wide, usually much less .............................................. 67 63(62). Synsepal basally produced into a conspicuous mentum; column foot very long, at least half the length of column; lip narrowly deltoid, broadly truncate apically ......................................................................... P. mentosa 63. Synsepal produced into a small, inconspicuous mentum or no mentum; column footless or foot < 1/3 the length of column; lip variously shaped but never narrowly deltoid nor broadly truncate apically ................. 64 64(63). Petals as long as dorsal sepal or slightly shorter; lip convex, geniculate, wider than long ........................................................................... P. revoluta 64. Petals to 1/4 the length of dorsal sepal or shorter; lip concave or flat, longer than broad ............................................................................................ 65 65(64). Synsepal at least 2 times broader than dorsal sepal; petals with margins dentate to deeply fimbriate ....................................................... P. lanceana

Pleurothallis 511

65. 66(65). 66.

67(62). 67. 68(67).

68.

69(67). 69. 70(69). 70. 71(70). 71. 72(71). 72. 73(72).

73.

74(71). 74. 75(74).

75.

76(75).

Synsepal as broad as dorsal sepal or slightly broader; petals with entire margins ................................................................................................ 66 Lateral sepals totally united into a synsepal; petals rhombic; flowers successive ........................................................................... P. rhombipetala Lateral sepals united into a synsepal for 1/2 or less of their length; petals elliptic to obovate-elliptic; flowers all opening simultaneously or at least many at a time ............................................................. P. sclerophylla Stems at least 30 mm long ....................................................................... 68 Stems to 17 mm long, usually much shorter ........................................... 69 Synsepal at least 2 times broader than dorsal sepal, totally connate; petals with dentate to deeply fimbriate margins; flowers usually orange or yellow-orange .............................................................................P. lanceana Synsepal as broad as dorsal sepal or slightly broader, united only in its basal third; petals with smooth margins; flowers usually cream, white, yellowish, or greenish, very rarely orange ........................... P. sclerophylla Sepals caudate-attenuate; petals with fimbriate to laciniate margin ..................................................................................................... P. aristata Sepals obtuse to acuminate but never caudate-attenuate; petals with entire to erose margins ............................................................................ 70 Leaves narrowly linear, < 1.5 mm wide; peduncle and rachis pubescent ............................................................................................... P. vittariifolia Leaves various but rarely narrowly linear, when so, at least 2 mm wide; peduncle and rachis glabrous .............................................................. 71 Petals 1.3–2 times as long as wide, rounded to broadly obtuse ............. 72 Petals (2.5–)3–5 times as long as wide, obtuse to acuminate ................. 74 Rachis zigzag, multiflowered; dorsal sepal obovate, rounded; lip basally very fleshy, spheroid thickened .................................................... P. seriata Rachis straight or slightly zigzag, flowers densely spaced; dorsal sepal elliptic or oblong-elliptic; lip not spheroid-thickened basally ............... 73 Inflorescences about as long as leaves; leaves elliptic to narrowly elliptic, acute to acuminate, (10–)15–60 mm long; flowers entirely yellow or orange ..................................................................................... P. corniculata Inflorescences 2.5–4 times as long as leaves; leaves broadly spatulate, obovate, rounded to broadly obtuse, 6–10 mm long; flowers yellowish with purple stripes ................................................................... P. parvifolia Leaves linear-obovate, 8–15 times as long as wide; dorsal sepal acuminate .......................................................................................................... P. picta Leaves obovate to elliptic, 3–5 times as long as wide; dorsal sepal acute to obtuse ................................................................................................... 75 Stem 3–12 mm long; leaves 12–70 mm long, marginate; dorsal sepal as broad as synsepal, about 3 times as broad as petals; lip glabrous ....................................................................................................... P. grobyii Stem 1–2 mm long; leaves 5–12(–16) mm long, not marginate; dorsal sepal narrower than synsepal, about as broad or slightly broader than petals; lip pubescent or glabrous ......................................................... 76 Rachis conspicuously abbreviate, all flowers emerging from one or a few cup-like bracts at top of peduncle; synsepal reflexed in natural position .............................................................................................................. 77

512

O RCHIDACEAE

76.

Rachis short but conspicuous; flowers not emerging from a cup-like bract at top of the peduncle; synsepal concave, or only slightly reflexed in natural position .................................................................................... 78 77(76). Petals elliptic, obtusely acute, glabrous; lip oblong-elliptic, marginally glabrous ......................................................................................... P. aondae 77. Petals lanceolate, acute, marginally ciliate; lip acute, marginally ciliate throughout ............................................................................... P. barbulata 78(76). Inflorescences as long or a little longer than subtending leaves; petals straight; lip simple, apically ciliate ............................................ P. minima 78. Inflorescences 4–5 times longer than subtending leaves; petals obliquely deflexed, somewhat falcate; lip basally with somewhat porrect, truncate lateral lobes, apically glabrous ................................................. P. sp A Pleurothallis aondae Carnevali & G.A. Romero in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1141. 2000. Epiphyte 2.5–3 cm tall, cespitose; leaves obovate, rounded, 1.5–2.5 cm long; inflorescences a little shorter than subtending leaves; flowers very dark wine-maroon; dorsal sepal ca. 5 mm long. Rain forests, ca. 600 m; Bolívar (Río Aonda on Auyán-tepui). Endemic. Pleurothallis aondae is one of the members of the P. barbulata complex. It is easily recognized by its rachis entirely clothed by floral bracts and glabrous lip and petals. Pleurothallis archidiaconi Ames, Sched. Orchid. 9: 29. 1925. Pleurothallis monocardia auct. non Rchb. f. 1855: sensu Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 1: 337, fig. 312. 1959. Epiphyte or subterrestrial, cespitose, shade-loving; leaves 7–10 cm long; flowers resupinate, brownish maroon, tending to yellow-green at margins, lip red-maroon with yellow center; dorsal sepal 5–7 mm long. Cloud forests, 900–1400 m; Bolívar (south of Aparamán-tepui, Kavanayén, La Escalera to Cerro Venamo region), Aragua, Distrito Federal, Falcón, Miranda; Trinidad-Tobago, Guyana, Suriname, Ecuador, Amazonian Brazil. Pleurothallis curvifructa H. G. Jones [Bradea 1(23): 263. 1972], known from Guyana, might belong here but the type of this concept was not available for confirmation. The eastern Andean taxon, Pleurothallis omoglossa Luer, may also be conspecific. Pleurothallis arenicola (Carnevali & I. Ramírez) Carnevali & I. Ramírez in G.A.

Romero & Carnevali, Orchids Venez. Ill. Field Guide 1141. 2000. ––Myoxanthus aspasicensis subsp. arenicola Carnevali & I. Ramírez, Novon 3: 117. 1993. Pleurothallis uncinata auct. non Fawc. 1895: sensu Foldats in Fl. Venez. 15(2): 437. 1970, pro parte; Dunsterv. & Garay, Venez. Orchid. Ill. 1: 355. 1959, pro parte. Lithophyte, terrestrial, or rarely an epiphyte; leaves 7.5–13 × 1.5–3.3 cm; flowers brown or reddish with purple markings; sepals externally pubescent with the hairs growing in clumps; lip yellow-brown, heavily covered with dark purple-brown. In sand or on sandstone, locally common in shrublands, rarer in rain forests or edges of cloud forests, (300–)600–1400(–2000) m; Bolívar (Auyántepui, Cerro Guaiquinima, Gran Sabana, Jaua-Sarisariñama massif, La Escalera to Cerro Venamo region, Río Paragua), Amazonas (Cerro Duida, Cerro Yapacana, Sierra de la Neblina). Guyana, Brazil (Amazonas). ◆Fig. 458. Pleurothallis arenicola is related to the mainly Central American and Andean P. aspasicensis Rchb. f., also known from the flora area from one collection, but has proportionally shorter, broader leaves, smaller flowers, and a consistent terrestrial habit over sandstone or sandy soils. Pleurothallis aristata Hook., Ann. Mag. Nat. Hist. 2: 229. 1839. —Specklinia aristata, Pridgeon & M.W. Chase, Lindleyana 16: 256. 2001. Pleurothallis barberiana Rchb. f., Gard. Chron. n.s. 16: 6. 1881. Epiphyte, cespitose; leaves 6–25 mm long; flowers large for the plant, resupinate; hyaline with deep maroon streaks, lip very dark

Pleurothallis 513

purple, almost black; dorsal sepal 4.5–11 mm long. Rain to cloud forests, often growing among bryophytes, 100–1100 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region), Amazonas (Cerro Yutajé). Aragua, Carabobo, Miranda, Yaracuy; Costa Rica, Panama, West Indies, Colombia, Guyana, Ecuador. Pleurothallis aspasicensis Rchb. f., Bonplandia (Hanover) 3: 73. 1855. —Humboldtia aspasicensis (Rchb. f.) Kuntze, Revis. Gen. Pl. 2: 667. 1891. —Myoxanthus aspasicensis (Rchb. f.) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 1: 38. 1986. —Echinella aspasicensis (Rchb. f.) Pridgeon & M.W. Chase, Lindleyana 16: 253. 2001. —Echinosepala aspasicensis (Rchb. f.) Pridgeon & M.W. Chase, Lindleyana 17: 101. 2002. Pleurothallis uncinata auct. non Fawc. 1895: sensu Foldats in Fl. Venez. 15(2): 437. 1970, pro parte. Epiphyte, shortly creeping or cespitose; stems 6–10 cm long; leaves 13–20 cm long, 2.5–4.5 cm wide, obovate or obtuse, basally attenuate; lateral sepals ca. 12 mm long, 6 mm wide, lip 5 mm long, 2.6–2.7 mm wide. Cloud forests, 1500–1600 m; Bolívar (Ptari-tepui). Costa Rica, Panama, Colombia, Ecuador. The flora area material of Pleurothallis aspasicensis has smaller flowers than the Andean populations. Pleurothallis barbulata Lindl., Fol. Orchid., Pleurothallis 40. 1859. —Pleurothallis barbata H. Focke, Bot. Zeitung (Berlin) 11: 227. 1853, non Westc. 1841. —Anathallis barbulata, Pridgeon & M.W. Chase, Lindleyana 16: 247. 2001. Epiphyte, minute; leaves 10–16 mm long, obovate; flowers large for the plant, entirely purple or greenish; sepals reflexed; dorsal sepal 2.8–3.5 mm long. Rain forests, frequently growing on thin branches, 50–800(–1200) m; Delta Amacuro (Caño Simoina west of Isla Cocuina), Bolívar (Altiplanicie de Nuria, Auyán-tepui, La Escalera to Cerro Venamo region, Río Icabarú), Amazonas (Río Casiquiare, Río Cataniapo, Sierra Parima). Aragua, Falcón, Miranda, Yaracuy; Guyana, Suriname, Ecuador, Brazil. Sometimes Pleurothallis abjecta Ames, P. nubensis Fold., and P. minima C. Schweinf. are considered to belong within P. barbulata,

in which case its range would extend from Mexico to Ecuador. All three species, however, seem to be amply distinct so as to guarantee specific status. Pleurothallis barbulata is the “core” taxon of a complex that includes many closely related taxa differing in details of lip shape, petal shape, pubescence, and whether the rachis of the inflorescence is elongated or not. This complex includes the following species in the flora area: P. aondae, P. barbulata, P. hillariana, P. minima, P. steinbuchiae, and P. sp. A. Pleurothallis brevipes H. Focke, Tijdschr. Wis- Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 2: 198. 1849. —Anathallis brevipes, Pridgeon & M.W. Chase, Lindleyana 16: 247. 2001. Epiphyte, cespitose, erect; leaves 2.5–5.5 cm long; inflorescence continuously flowering, flowers resupinate, opening well, sepals pale yellow or yellow-orange, petals yellow, often striped with red, lip marked with dark orange or red; dorsal sepal 5.5–6-6 mm long. Rain forests, 100–200 m; Amazonas (Paso de Ganado, San Antonio). Táchira; Guyana, Suriname, Ecuador, Bolivia. Pleurothallis callifera C. Schweinf., Fieldiana, Bot. 28: 180. 1951. Epiphyte, cespitose; leaves 5–11 cm long; flowers widely spreading, apparently nonresupinate, sepals translucent with dull lavender stripes, petals dull lavender, lip reddish purple; dorsal sepal 7–7.5 mm long. Cloud forests; 2100–2300 m; Bolívar (Macizo del Chimantá, Ptari-tepui). Endemic. The closely related Pleurothallis stenocardium has smaller flowers, no mentum in the synsepal, and the lip has a smaller, deeper glenion and is basally not flexed. The scant material of cordate-leaved members of the genus Pleurothallis need urgent revision. Pleurothallis coriacardia Rchb. f., Bonplandia (Hanover) 2: 26. 1854. Pleurothallis linguifera auct. non Lindl. 1859: sensu Foldats in Fl. Venez. 15(2): 329. 1970. Pleurothallis lansbergii auct. non Regel 1856: sensu Dunst. & Garay, Orchids Venez. Ill. Field Guide 783. 1979. Epiphyte, cespitose, shade-loving; leaves (5–)10–14 cm long; flowers resupinate, large for the genus, yellowish with red-brown

514

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veins, or completely red-brown or purplish; dorsal sepal (8–)10–15(–23) mm long. Cloud forests, 1200–1900 m; Bolívar (Cerro Jaua), Amazonas (Cerro Huachamacari, Cerro Aracamuni, Sierr de la Neblina). Aragua, Distrito Federal, Mérida, Miranda, Táchira, Trujillo, Yaracuy; Colombia, Ecuador. Outside the flora area, Pleurothallis coriacardia tends to grow at higher elevations. Most collections of it from the flora area have been reported as having purplish flowers, while yellowish flowers are more common in Andean populations. Flora area populations also have much smaller flowers with dorsal sepals 6–7 mm long and are referred to Pleurothallis coriacardia with some hesitation. Pleurothallis corniculata Lindl., Bot. Reg. 1842: misc. 83. 1842. Pleurothallis uniflora auct non Lindl. 1836: sensu Dunst. & Garay, Orch. Venez. 1: 844. 1979. Epiphyte, cespitose, shade-loving; leaves 2.5–7 cm long; flowers resupinate, campanulate, all segments bright or dull orange, or dark yellow; dorsal sepal 6–6.6 mm long. Locally common in rain forests, near sea level to 600 m; Delta Amacuro (Punta Barima, Río Amacuro, Río Cuyubini), Bolívar (Altiplanicie de Nuria, Auyán-tepui, Icabarú, Río Tonoro, Sierra Imataca). Apure, Barinas, Lara, Portuguesa, Zulia; Colombia, Guyana, Suriname, Brazil. ◆Fig. 460. Pleurothallis deborana Carnevali & I. Ramírez, Ann. Missouri Bot. Gard. 77: 553. 1990. Epiphyte, minute, creeping, shade-loving; leaves 5.5–8 mm long; flowers clear greenish yellow, maroon-tinged within segments, petals with a dark purple longitudinal stripe, lip dark red-purple with a yellow longitudinal streak; dorsal sepal ca. 3.2 mm long. Rain forests, ca. 100 m; Bolívar (close to Río Parguaza). Endemic. ◆Fig. 456. Pleurothallis discophylla Luer & Carnevali, Novon 3: 158. 1993. Pleurothallis coffeicola auct. non Schltr. 1929: sensu Foldats in Lasser, Fl. Venez. 15(2): 245. 1970. Epiphyte, short-creeping, shade-loving; leaves 4–7 cm long; flowers subcampanulate, resupinate, perianth segments yellow-or-

ange, brownish at apex of synsepal, lip purple-brown with a brownish yellow apex; dorsal sepal 9.2–12 mm long. Rain forests 100–400(–1200) m; southern Amazonas (Cerro Marahuaka, Río Casiquiare, Río Negro). Amazonian Colombia, Ecuador, Peru, Brazil, and Bolivia. Pleurothallis elvirana Carnevali & I. Ramírez, Novon 3: 118. 1993. Epiphyte, erect; leaves 14–17 cm long; flowers not resupinate, widely spreading, yellow; dorsal sepal 6.5–7.5 mm long. Cloud forests, 1200–1300 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 470. Pleurothallis elvirana is the only known member of the Pleurothallis lindenii Lindl. complex in the Guayana Region. Pleurothallis erebatensis Carnevali & G.A. Romero, Novon 3: 18. 1993. Epiphyte, cespitose or shortly creeping, erect; leaves 5.5–9.5 cm long; flowers resupinate or not, dark brownish red; dorsal sepal 4.8–5 mm long. Rain forests, 300–400 m; Bolívar (near Santa María de Erebato). Táchira; possibly Bolivia. Pleurothallis erinacea Rchb. f., Bonplandia (Hanover) 3: 72. 1855. Epiphyte, shortly creeping; leaves 9–12 cm long; flowers relatively large for the genus, resupinate, clear yellow or brownish; dorsal sepal 13–16 mm long. Rain forests, 1000–1100 m; Bolívar (near Santa Elena de Uairén). Aragua, Miranda; Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Suriname, Ecuador, Peru, Bolivia. Pleurothallis floribunda Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 48, t. 84. 1836. —Stelis floribunda (Poepp. & Endl.) Pridgeon & M.W. Chase, Lindleyana 16: 263. 2001, non H.B.K. 1815 [1816]. —Stelis vulgaris Pridgeon & M.W. Chase, Lindleyana 17: 100. 2002. Pleurothallis pluriracemosa Garay, Arch. Jard. Bot. Rio de Janeiro 12: 174. 1952. Pleurothallis velaticaulis auct. non Rchb. f. 1849: sensu Foldats in Lasser, Fl. Venez. 15(2): 440. 1970. Epiphyte or lithophyte, erect, usually sunloving; leaves 4–30 cm long; flowers resupinate, campanulate, yellow-green, pale yel-

Pleurothallis 515

low, or pale orange; dorsal sepal 3–6 mm long. Rain or cloud forests, 900–1400(–2100) m; Bolívar (Aparamán-tepui, Auyán-tepui, La Escalera to Cerro Venamo region), Amazonas (Cerro Aracamuni, Sierra de la Neblina). Anzoátegui, Aragua, Miranda, Monagas, Nueva Esparta, Sucre, Yaracuy; Costa Rica, Panama, Colombia, Trinidad-Tobago, Ecuador, Peru, Bolivia. Pleurothallis fockei Lindl., Fol. Orchid., Pleurothallis 23. 1859. —Humboldtia fockei (Lindl.) Kuntze, Revis. Gen. Pl. 2: 667. 1891. —Pleurothallis tricarinata H. Focke, Bot. Zeitung (Berlin) 11: 339. 1853, non Poepp. & Endl. 1853. —Acianthera fockei, Pridgeon & M.W. Chase, Lindleyana 16: 243. 2001. Pleurothallis myrmecophila Hoehne, Comiss. Linhas Telegr. Estratég. Mato Grosso Amazonas, anexo 5, Bot. 5: 47. 1915. Epiphyte, erect or subpendent, usually purple-tinged, shade-loving; leaves 4–10 cm long; flowers resupinate, maroon or brownish, petals whitish or hyaline, lip dark maroon; dorsal sepal 4.2–6 mm long. Rain forests, 100–400 m; Bolívar (Río Canaracuní, Río Paragua, Río Parguaza), Amazonas (Cerro Yutajé, near Puerto Ayacucho, San Carlos de Río Negro, Santa Rosa de Amanadona). Apure; Brazil, Suriname. The sterile collection, Liesner 17881 (MO, VEN), from the vicinity of Culebra (Amazonas) resembles this species but the leaf is much narrower. It probably represents a previously unreported species of Pleurothallis for the flora area. Pleurothallis funerea (Barb. Rodr.) Cogn. in Mart., Fl. Bras. 3(4): 567. 1896. ––Lepanthes funerea Barb. Rodr., Vellosia ed. 2, 1: 118. 1891. Pleurothallis breviscapa C. Schweinf., Bot. Mus. Leafl. 3: 79. 1935. Pleurothallis ciliolata auct. non Schltr. 1926: sensu Foldats in Lasser, Fl. Venez. 15(2): 240. 1970; Dunst. & Garay, Orchids Venez. Ill. Field Guide 747. 1979. Epiphyte, cespitose, erect or prostrate; leaves 7–20 mm long; flowers campanulate, resupinate, sepals and petals maroon, lip of a darker magenta hue. Rain forests, 200–500 m; Delta Amacuro (Serranía de Imataca), Bolívar (Serranía de Imataca, Río Icabarú).

Colombia, Guyana, Ecuador, Brazil (Amazonas), Bolivia. The entity previously called Pleurothallis breviscapa from the Venezuelan Coastal Range and the Andes from Venezuela through Ecuador is Pleurothallis rabei Foldats. Pleurothallis galeata Lindl., Ann. Mag. Nat. Hist. ser. 1, 15: 107. 1845. —Stelis galeata, Pridgeon & M.W. Chase, Lindleyana 16: 263. 2001. Epiphyte or subterrestrial, cespitose, erect; leaves 13–22 cm long; flowers resupinate, nodding, yellowish or pale lavender marked with purplish along veins; dorsal sepal 4.5–6 mm long. Cloud forests, 1800–2000 m; Bolívar (Cerro Jaua), Amazonas (Sierra de la Neblina). Lara, Mérida, Táchira, Trujillo; Colombia, Ecuador, Peru. Pleurothallis garciae Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 76: 173. 1999. —Acianthera garciae, Pridgeon & M.W. Chase, Lindleyana 16: 243. 2001. Pleurothallis consimilis auct. non Ames 1922: sensu Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 2: 272. 1961; Foldats in Lasser, Fl. Venez. 15(2): 247. 1970. Epiphyte, creeping, shade-loving; leaves 2.5–4 cm long; flowers resupinate, sepals and petals white or hyaline with red-purple streaks and dots, lip red-purple; dorsal sepal 8–9 mm long. Rain or cloud forests, 800– 1000 m; Bolívar (collection by Dunsterville without locality). Falcón, Nueva Esparta; Guyana. Pleurothallis geographica Luer, Selbyana 3: 310. 1977. Epiphyte or lithophyte, cespitose; leaves 5–8 cm long, deflexed on the stem; flowers apparently campanulate, described as magenta; dorsal sepal 4–4.5 mm long. Rock outcrops along streams, ca. 2200 m; Bolívar (Macizo del Chimantá). Andean Colombia and Ecuador. The Andean populations of Pleurothallis geographica have somewhat larger flowers (dorsal sepal 6.5–8 mm long) with yellow sepals and petals and a red-purple lip. Pleurothallis granitica Luer & G.A. Romero in G.A. Romero & Carnevali,

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Orchids Venez. Ill. Field Guide 1142. 2000. Pleurothallis papillosa auct. non. Lindl. 1836: sensu Dunst. & Garay, Orchids Venez. Ill. Field Guide 801. 1979. Lithophyte, creeping, shade-loving; leaves often green-purple, 3–6 cm long; flowers resupinate, campanulate, dark red-maroon or dark purple, lip darker maroon; dorsal sepal 4–5 mm long. Shady granitic outcrops, 50– 100(–400) m; Bolívar (Río Maniapure, Río Ore, Río Parguaza), Amazonas (Laja Los Oripopos, Laja Garcitas, Rio Cataniapo). Brazil. ◆Fig. 471. Pleurothallis grobyi Bateman ex. Lindl., Edwards’s Bot. Reg. 21: t. 1797. 1835. —Specklinia grobyi, Pridgeon & M.W. Chase, Lindleyana 16: 258. 2001. Epiphyte, cespitose, erect, usually shadeloving; leaves 1.2–7 cm long; flowers resupinate, white, yellowish, greenish yellow, or hyaline with maroon or red stripes, lip frequently entirely reddish; dorsal sepal 3–6 mm long. Rain forests, 100–700(–1000) m; widespread in Bolívar and Amazonas. Neotropics. ◆Fig. 469. Pleurothallis hexandra Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 3: 252. 1965. —Garayella hexandra (Garay & Dunst.) Brieger, Orchideen 425. 1975, nom. inval. Restrepia kegelii Rchb. f., Linnaea 41: 133. 1877, non Pleurothallis kegelii Rchb. f. 1877. —Barbosella kegelii (Rchb. f.) Schltr., Repert. Spec. Nov. Regni Veg. 15: 262. 1918. —Chamelophyton kegelii (Rchb. f.) Garay, Orquideología 9: 115. 1974. Epiphyte, growing low on rough-barked shrubs, often in shade; leaves purplish on the lower surface; flowers lasting 3 or 4 days; synsepal greenish or brownish, purpletinged; dorsal sepal, petals, and lateral lobes of lip apple green with purple-red apex, blade of lip pale brownish orange, purpletinged. Rain forests, sclerophyllous scrub, 400–1300 m; Bolívar (Gran Sabana, Río Paragua). Guyana. ◆Fig. 454. Pleurothallis hilariana Carnevali & G.A. Romero in G.A. Romero & Carnevali,

Orchids Venez. Ill. Field Guide 1143. 2000. Epiphyte 0.5–1 cm tall, shortly creeping; leaves ca. 3.5 mm long, 2.5 mm wide; flowers apparently not widely spreading, dorsal sepal 3.5 mm long, 1.3 mm wide; lip ca. 2.6 mm long, 0.6 mm wide. Rain forests, ca. 1000 m; Bolívar (Ilú-tepui). Endemic. Pleurothallis hitchcockii Ames, Orchidaceae 7: 117. 1922. Pleurothallis ringens C. Schweinf., Bot. Mus. Leafl. 10: 184. 1942. Epiphyte, cespitose, erect, shade-loving; leaves 4–8 mm long; flowers resupinate, bilabiate, pale green with maroon veins; dorsal sepal 14–18 mm long. Rain forests, 400– 1000 m; Bolívar (La Escalera to Cerro Venamo region, Río Akaruai, Río Betanía in Gran Sabana, Río Icabarú, Río Paragua), Amazonas (Sierra Parima). Falcón; Guyana, Ecuador. Pleurothallis holstii Carnevali & I. Ramírez, Ernstia 39: 18. 1986. Epiphyte, minute, cespitose, erect; leaves 8–10 mm long; flowers green-yellow. Rain forests, 100–200 m; Bolívar (Río Parguaza). Endemic. ◆Fig. 457. Pleurothallis humilis C. Schweinf., Fieldiana, Bot. 28: 185. 1951. Epiphyte, cespitose, erect; leaves usually purple-spotted, 3–4.5 cm long; flowers orange-brown, lip red or maroon with a yellow central stripe and white apex and lateral lobes; dorsal sepal 5–6 mm long. Rain forests, (100–)600–1600 m; Bolívar (Auyántepui, Kavanayén, Macizo del Chimantá [Amurí-tepui], Río Pacairao), Amazonas (Río Autana). Endemic. Pleurothallis imraei Lindl., Fol. Orchid., Pleurothallis 9. 1859. —Stelis imraei, Pridgeon & M.W. Chase, Lindleyana 16: 263. 2001. Pleurothallis vaginata Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19: 197. 1923. Epiphyte, cespitose; leaves 5.5–15 cm long; flowers greenish, yellowish, brownish, or purplish, usually cleistogamous; dorsal sepal 5.5– 9 mm long. Cloud forests, 700–1600(–2300) m;

Pleurothallis

Bolívar (Auyán-tepui, Cerro Uroi, La Escalera to Cerro Venamo region, Macizo del Chimantá, Ptari-tepui, Sierra de Maigualida), Amazonas (Cerro Marahuaka). Lara, Yaracuy; Costa Rica, Panama, West Indies, Guyana, Colombia, Ecuador, Peru, Bolivia. Pleurothallis ionantha Rchb. f., Linnaea 22: 830. 1850. Epiphyte, cespitose, erect; leaves 3–7 cm long; flowers pale yellow, resupinate, campanulate; dorsal sepal 4–5 mm long. Rain forests, 700–900 m; Bolívar (Río Carrao, Río Churún near Guarimba). Aragua, Miranda. Pleurothallis kerrii Braga, Bradea 3: 172. 1981. Pleurothallis delascioi Carnevali & I. Ramírez, Ernstia 31: 6. 1985. Epiphyte, long-creeping, usually growing low on trees among bryophytes, shade-loving; leaves 3.5–4.3 mm long; flowers frequently cleistogamous, purple with darker veins, lip dark purple; dorsal sepal ca. 2 mm long. Rain forests, 50–600 m; Amazonas (Río Ararí, Río Cataniapo, Salto Yureba, San Carlos de Río Negro-Solano area). Amazonian Peru and Brazil. ◆Fig. 453. Pleurothallis lanceana Lodd., Bot. Cab. 18: t. 1767. 1831 [1832]. —Acianthera lanceana, Pridgeon & M.W. Chase, Lindleyana 16: 244. 2001. Pleurothallis ciliata Knowl. & Westc., Flor. Cab. 1: 89. 1837. Pleurothallis plumosa Lindl., Bot. Reg. 1842: misc. 72. 1842. Pleurothallis kegelii Rchb. f., Linnaea 41: 132. 1877. Pleurothallis huebneri Schltr., Beih. Bot. Centralbl. 42(2): 90. 1925. Pleurothallis ciliata var. elongata C. Schweinf., Bot. Mus. Leafl. 16: 47. 1953. Epiphyte, shortly creeping, usually growing in ant nests; leaves 5–14 cm long, elliptic to obovate-elliptic; flowers orange or orangeyellow with darker orange veins, resupinate; dorsal sepal 9–17 mm long. Rain forests, 50– 300(–800); Bolívar (Altiplanicie de Nuria, La Escalera to Cerro Venamo region, upper Rio Caroní), Amazonas (widespread). Aragua, Distrito Federal, Miranda, Monagas; Costa Rica, Panama, Colombia, Guyana, Suri-

517

name, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 462. As here circumscribed, Pleurothallis lanceana is a variable entity that may encompass several subspecific or specific taxa. Amazonian plants are larger with less coriaceous leaves and laxer, longer inflorescences that bear flowers on the larger ranges of size. However, intermediates between these Amazonian plants and the more compact populations from the Guianas region, the Venezuelan Coastal Range, and other parts of the Neotropics are commonly found, making it difficult to draw clear lines of discontinuity between the several forms. A revision of the complex is needed. Pleurothallis lappiformis A.H. Heller & L.O. Williams, Fieldiana, Bot. 31: 42. 1965. —Myoxanthus lappiformis (A.H. Heller & L.O. Williams) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 44: 38. 1986. —Echinella lappiformis (A.H. Heller & L.O. Williams) Pridgeon & M. W. Chase, Lindleyana 16: 253. 2001. —Echinosepala lappiformis (A.H. Heller & L.O. Williams) Pridgeon & M. W. Chase, Lindleyana 17: 101. 2002. Lithophyte; leaves ca. 19 × 5 cm, longer than stem; inflorescences 1–3-flowered; flowers deep red-purple, borne at soil level, almost always producing large, red, hairy capsules. Shady positions in shrublands, 700– 1700 m; Bolívar (Cerro Guaiquinima, Cerro Sarisariñama, Luepa). Nicaragua, Panama, Ecuador. ◆Fig. 466. Pleurothallis longisepala Barb. Rodr., Vellosia ed. 2, 1: 115. 1891. Epiphyte, creeping, shade-loving; leaves 2.5–3.5 mm long; flowers resupinate, yellowish with purple veins; dorsal sepal 5–10 mm long. Rain forests, 100–200 m; Amazonas (Río Atabapo). Amazonian Brazil. Pleurothallis longisepala might be only a variety of the more common P. miqueliana. Pleurothallis loranthophylla Rchb. f., Bot. Zeitung (Berlin) 10: 674. 1852. —Rhynchopera punctata H. Karst., Auswahl Gew. Venez. 21, t. 7. 1848 [1849], "Rhynchophora," non Pleurothallis punctata Lindl. 1835.

518

O RCHIDACEAE

Epiphyte, cespitose, shade-loving; leaves 6–12 cm long; flowers resupinate, purplish or greenish with purple spots; dorsal sepal 6.5– 12 mm long. Cloud forests, 1000–1200 m; Bolívar (La Escalera to Cerro Venamo region). Aragua, Carabobo, Distrito Federal, Falcón, Mérida, Miranda, Táchira, Zulia; Colombia, Ecuador, Peru, Bolivia. Pleurothallis maguirei Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 76: 113. 1999. —Anathallis maguirei, Pridgeon & M.W. Chase, Lindleyana 16: 249. 2001. Epiphyte, cespitose; leaves 5–9 × 1.5–1.8 cm; flowers resupinate, campanulate; sepals connate in the lower third, light golden yellow, petals and lip pale yellow; dorsal sepal 6.5–8.5 mm long. Dwarf cloud forests, 1400– 1500 m; Bolívar (La Escalera–Cerro Venamo area). Guyana. This recently described taxon is closely related to the variable and widespread Pleurothallis sclerophylla but in P. maguirei the sepals are obtuse, oblong, the lip has well-defined lateral lobes, and the column lacks the beak found in P. sclerophylla. Pleurothallis mentosa Cogn. in Mart., Fl. Bras. 3(4): 400. 1896. —Lepanthes yauaperyensis Barb. Rodr., Vellosia 2, 1: 117. 1891, non Pleurothallis yauaperyensis Barb. Rodr. 1891. —Anthereon mentosus, Pridgeon & M.W. Chase, Lindleyana 16: 252. 2001. Epiphyte, cespitose, erect; leaves 2–6.5 cm long; flowers resupinate, yellow-green to pale brown, often with brown or maroon spots; dorsal sepal 8–9 mm long. Rain forests, 100–500 m; Bolívar (quebrada Los Brasileros near Icabarú, Río Caura, Río Icabarú, Río Paragua), Amazonas (Raudal Monserrat, Río Siapa, Tamatama). Lara, Táchira; Amazonian parts of Ecuador, Peru, Brazil, and Bolivia. Pleurothallis minima C. Schweinf., Bot. Mus. Leafl. 3: 82. 1935. Epiphyte, minute, cespitose, usually growing on thin branches in shady or sunny spots; leaves 5–8 mm long; flowers resupinate, campanulate, sepals and petals brownish purple or maroon with a dark purple lip; dorsal sepal 2.7–3 mm long. Rain forests, 600–900 m; Bolívar (El Paují, Río Icabarú),

Amazonas (Ugueto). Distrito Federal, Miranda; Guyana, Amazonian Brazil. Pleurothallis minima has often been placed in the synonymy of P. barbulata, the judgements having been made based mostly on herbarium specimens. The differences (indicated in the key) are enough for the recognition of both species as distinct. Frequently both are found in the same tree and no intermediates occur. Pleurothallis miqueliana (H. Focke) Lindl., Fol. Orchid., Pleurothallis 17: 1859. —Specklinia miqueliana H. Focke, Tijdschr. Wis- Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 2: 199. 1849. —Acianthera miqueliana, Pridgeon & M.W. Chase, Lindleyana 16: 244. 2001. Pleurothallis fimbriata Lindl., Fol. Orchid., Pleurothallis 17. 1859. Epiphyte, creeping, usually growing low on tree trunks, shade-loving; leaves 2.5–4.5 mm long; flowers resupinate, sepals pale reddish brown, ochre-yellow toward apices, lip dark maroon; dorsal sepal 6–7.5 mm long. Rain forests, near sea level to 300(–600) m; Delta Amacuro (Río Arature), Bolívar (Altiplanicie de Nuria), Amazonas (widespread). Aragua, Distrito Federal; Guyana, Amazonian Ecuador, Peru, and Brazil. ◆Fig. 463. The Venezuelan Coastal Range collections of this typically Amazonian species show certain differences and might represent a distinct species. Pleurothallis morilloi Carnevali & I. Ramírez, Novon 3: 119. 1993. Epiphyte, creeping, shade-loving; leaves 2.3–3.2 cm long; flowers resupinate, not opening completely, sometimes cleistogamous; greenish yellow; lip greenish white; dorsal sepal 4 mm long. Rain forests, 600– 900 m; Bolívar (Río Akaruai, Río Aonda on Auyán-tepui). Táchira. ◆Fig. 472. Pleurothallis moritzii Rchb. f., Linnaea 22: 824. 1849. —Stelis moritzii, Pridgeon & M.W. Chase, Lindleyana 16: 264. 2001. Pleurothallis elegans auct. non (H.B.K.) Lindl. 1842: sensu Foldats in Lasser, Fl. Venez. 15(2): 262. 1970.

Pleurothallis 519

Pleurothallis tepuiensis auct. non Carnevali & I. Ramírez 1993: sensu G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 858. 2000. Epiphyte or lithophyte, cespitose or shortly creeping, usually sun-loving; leaves 5–10 cm long; flowers campanulate, resupinate, yellow; dorsal sepal 6–8 mm long. Cloud forests or open associations, (1300–) 1900–2800 m; Bolívar (Cerro Jaua, La Escalera to Cerro Venamo region, Macizo del Chimantá, Ptari-tepui), Amazonas (Cerro Coro Coro, Cerro Marahuaka). Aragua, Distrito Federal; Ecuador (?). ◆Fig. 459.

gion, Río Paragua, Waramasén). Peru, Amazonian Brazil, Bolivia.

Pleurothallis nanifolia Foldats, Bol. Soc. Venez. Ci. Nat. 22: 258. 1961. Epiphyte, minute, creeping; leaves 5–8 mm long; flowers resupinate, sepals pale maroon or dark maroon-red, petals pale pink with very dark maroon midvein, lip dark maroon-purple with white basal lobes; dorsal sepal 3.5–4 mm long. Rain forests, 100–300 m; Bolívar (Río Paragua), Amazonas (near Puerto Ayacucho, Río Atabapo). Brazil (Mato Grosso), Bolivia.

Pleurothallis phoenicoptera Carnevali & G.A. Romero, Novon 4: 88. 1994. Epiphyte, creeping, pendent or horizontally spreading, usually in well-illuminated positions, purple-tinged; leaves 3–4 cm long; flowers cleistogamous, sepals white or greenish, suffused purple within, petals translucent white, lip white lightly marked with purple on the margins; persistent perianth on the fruit recurved and resembling a bird’s head, fleshy; dorsal sepal 5–8.1 mm long. Sclerophyllous forests, 1200–1500 m; Bolívar (Cerro de la Piedra del Canaima, Salto Torón), Amazonas (Cerro Duida). Endemic.

Pleurothallis omissa Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 76: 175. 1999. —Acianthera omissa, Pridgeon & M.W. Chase, Lindleyana 16: 245. 2001. Epiphyte, short-creeping; leaves 5–7 × 1– 1.3 cm; inflorescence pendulous; flowers resupinate, campanulate, sepals light brown, the dorsal with 3 purple veins, petals translucent brown with a purple midvein, lip dark red-purple; dorsal sepal 1.75–2 mm wide. Rain forests, 200–300 m; Amazonas (Río Casiquiare). Endemic. This rare taxon is closely related to Pleurothallis fockei but the dorsal sepal is much longer (as long as the synsepal) and the lip is broadly rounded at apex (acute in P. fockei). Pleurothallis parvifolia Lindl., Companion Bot. Mag. 2: 355. 1836. —Specklinia parvifolia, Pridgeon & M.W. Chase, Lindleyana 16: 258. 2001. Epiphyte, cespitose, erect, shade-loving; leaves 6–10 mm long; flowers resupinate, yellow with red or purplish veins; dorsal sepal 6–8 mm long. Rain forests, 300–800 m; Bolívar (La Escalera to Cerro Venamo re-

Pleurothallis pemonum Carnevali & I. Ramírez, Ann. Missouri Bot. Gard. 77: 555. 1990. Epiphyte, minute, creeping; leaves 5–7 (–9) mm long; flowers deep wine red, the sepals semitranslucent, lip dark red with a central yellow-cream longitudinal concavity; dorsal sepal 3.8–4 mm long. Rain forests, 100–800 m; Bolívar (Cerro Guaiquinima, Río Akaruai, Río Carrao, Río Icabarú), Amazonas (Río Gavilán). Endemic. ◆Fig. 455.

Pleurothallis picta Lindl., Edwards’s Bot. Reg. 21: sub t. 1797. 1836. —Specklinia picta, Pridgeon & M.W. Chase, Lindleyana 16: 259. 2001. Epiphyte, cespitose; leaves 2.5–6 cm long; flowers resupinate, campanulate, cream or yellowish; dorsal sepal 6–9 mm long. Rain forests, 100–1000 m; Bolívar (quebrada Los Brasileros near Icabarú, upper Río Caura, Río Erebato), Amazonas (Boca Padamo, Cerro Huachamacari, Río Siapa, Sierra de la Neblina). Táchira, Trujillo; Guyana, Suriname, French Guiana, Ecuador, Brazil. Pleurothallis picta has been treated as a synonym of P. grobyi Bateman, but the narrower leaves and acuminate sepals are distinctive. There are several forms of this taxon in different areas of the Neotropics and some might represent distinct species. Pleurothallis prolifera Herb. ex Lindl., Edwards’s Bot. Reg. 15: t. 1298. 1830. —Acianthera prolifera, Pridgeon & M.W. Chase, Lindleyana 16: 245. 2001.

520

O RCHIDACEAE

Epiphyte, creeping, erect; leaves 3.2–4 cm long; flowers resupinate, purplish or maroon; dorsal sepal 5–9 mm long. Cloud forests, ca. 1500 m; Bolívar (Auyán-tepui). Southeastern Guyana, Brazil. Pleurothallis pruinosa Lindl., Edwards’s Bot. Reg. 28: misc. 75. 1842. Epiphyte, cespitose, usually sun-loving; leaves 2–5 cm long; flowers resupinate, pale yellow, creamy white, or greenish; dorsal sepal 2.5–4 mm long. Rain forests, 50–600 m; Delta Amacuro (Caño Capurito, Río Acure, Río Cuyubini), Bolívar (Altiplanicie de Nuria, Río Toro), Amazonas (Caño Yapacana, Salto Yureba). Anzoátegui, Aragua, Carabobo, Distrito Federal, Mérida, Miranda, Sucre, Zulia; widespread in Neotropics. Pleurothallis revoluta (Ruiz & Pav.) Garay, Caldasia 8: 520. 1962. —Humboldtia revoluta Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 1: 235. 1798. Epiphyte, cespitose; leaves 8–27 cm long; flowers nonresupinate, translucent, greenish or yellowish, often with red or purplish dots; dorsal sepal 1.1–2.5 mm long. Cloud forests, 1800–2000 m; Bolívar (Macizo del Chimantá, Ptari-tepui, Roraima-tepui), Amazonas (Sierra de la Neblina). Anzoátegui, Distrito Federal, Mérida, Monagas, Nueva Esparta, Portuguesa, Sucre; Trinidad-Tobago, Colombia, Ecuador, Peru, Bolivia.

greenish, yellowish, or hyaline; dorsal sepal 5–7 mm long. Rain or cloud forests, 600– 1500 m; Bolívar (Altiplanicie de Nuria, Auyán-tepui, La Escalera to Cerro Venamo region, Macizo del Chimantá, Ptari-tepui), Amazonas (Cerro Marahuaka). Anzoátegui, Miranda, Nueva Esparta, Sucre, Táchira; widespread in the Neotropics. Pleurothallis samacensis Ames, Sched. Orchid. 2: 22. 1923. —Specklinia samacensis, Pridgeon & M.W. Chase, Lindleyana 16: 259. 2001. Epiphyte, minute, cespitose, shade-loving, usually growing among bryophytes; leaves 1–2 cm long; flowers resupinate, often cleistogamous, yellowish brown with purple veins or totally purplish; dorsal sepal 5–6 mm long. Cloud forests 900–1800 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region). Aragua, Táchira; Guatemala, Costa Rica, Ecuador. ◆Fig. 464.

Pleurothallis rhombipetala Rolfe, Bull. Misc. Inform. Kew 1893: 4. 1893. Epiphyte, cespitose; leaves 9–10 cm long; flowers pale purple with dark purple dots, lip green with marginal spots of dark purple; dorsal sepal 9–10 mm long. Rain forests, ca. 2700 m; Bolívar (Roraima-tepui). Endemic. Pleurothallis rhombipetala has not been collected again since the original collection. It is not known whether it was collected in the Guyanan or Venezuelan part of Roraima.

Pleurothallis sandaliorum G.A. Romero & Carnevali, Harvard Pap. Bot. 5: 182. 2000. Pleurothallis coffeicola auct. non Schltr. 1929: sensu Foldats in Lasser, Fl. Venez. 15(2): 245. 1970, pro parte. Epiphyte, low on tree, short-creeping; leaves 4.5 cm long and wide, suborbicular; flowers with a green or yellowish dorsal sepal, petals green, purple-spotted, synsepal and lip purple; dorsal sepal 7 mm long, 2 mm wide. Rain forests, 100–200 m; Amazonas (upper Rio Yatúa, Yavita-Maroa road). Endemic. Pleurothallis sandaliorum is very closely related to Pleurothallis discophylla Luer & Carnevali, but in that species, the perianth segments are yellow-orange, brownish at the apex of the synsepal, and the lip is purplebrown with a brownish apex and elliptic in outline instead of obovate or obovatespathulate.

Pleurothallis ruscifolia (Jacq.) R. Br. in W.T. Aiton, Hortus Kew ed. 2, 5: 211. 1813. —Epidendrum ruscifolium Jacq., Enum. Syst. Pl. 29: 1760. Pleurothallis wendlandiana auct. non Rchb. f. 1888: sensu Dunst. & Garay, Venez. Orchid. Ill. 1: 357. 1959. Epiphyte, cespitose, shade-loving; leaves 8–15 cm long; flowers resupinate, white,

Pleurothallis sarcosepala Carnevali & I. Ramírez in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1144. 2000. Pleurothallis vitatta auct. non Lindl. 1838: sensu Foldats in Lasser, Fl. Venez. 15(2): 443. 1970, pro parte, as for the Guayana specimens cited. Epiphytic herbs 10–20 cm tall; leaves el-

Pleurothallis

liptic, 5–5.5 cm long, 2.5–2.7 cm wide; flowers not opening fully, greenish yellow with purple zones in petals, lip, and column, dorsal sepal 6 mm long, 2–2.5 mm wide. Cloud forests, 1200–1800 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region). Táchira. Plants of Pleurothallis sarcosepala are identical to those of P. rubroviridis Lindl., but the flowers are strikingly different. Pleurothallis rubroviridis is known from the West Indies, the Venezuelan Coastal Range, and the Andean slopes of Ecuador and Peru. Pleurothallis sclerophylla Lindl., Edwards’s Bot. Reg. 21: sub t. 1797. 1836. —Anathallis sclerophylla, Pridgeon & M.W. Chase, Lindleyana 16: 250. 2001. Pleurothallis stenopetala Lodd. ex Lindl., Edwards’s Bot. Reg. 21: misc. 95. 1836. Epiphyte, cespitose, erect, frequently sunloving; leaves 3.5–10 cm long; flowers resupinate, variable in shape and size of the sepals, yellow, pale orange, cream, or white; dorsal sepal 7.5–21 mm long. Rain or cloud forests, 800–1800 m; Bolívar (Auyán-tepui, Cerro Sarisariñama, Kavanayén, La Escalera to Cerro Venamo region, Macizo del Chimantá [Abacapá-tepui], Río Canaracuni, Roraimatepui), Amazonas (Cerro Aratitiyope, Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Yapacana, Cerro Yaví). Aragua, Distrito Federal, Lara, Mérida, Nueva Esparta, Táchira, Trujillo, Yaracuy; tropical South America. ◆Fig. 473. Pleurothallis sclerophylla is a very variable species regarding the length and pubescence of the sepals, number and disposition of flowers in the inflorescence, and vegetative size. Some of the extreme forms could represent distinct species, but a careful revision of the group is required across its entire distribution. Pleurothallis seriata Lindl., Edwards’s Bot. Reg. 26: misc. 75. 1840. —Specklinia seriata, Pridgeon & M.W. Chase, Lindleyana 16: 259. 2001. Pleurothallis diffusiflora C. Schweinf., Bot. Mus. Leafl. 3: 80. 1935. Epiphyte, cespitose; leaves 2.5–4.5 mm long; flowers nonresupinate, sepals yellowish orange with dark purple spots, petals orange with very dark purple at the margins

521

and apex; lip yellow with dark spots basally, the globose apex dark purple; dorsal sepal 5– 7 mm long. Rain forests, 400–600 m; Bolívar (Altiplanicie de Nuria). Guyana, Brazil. Pleurothallis spiculifera Lindl., Fol. Orchid., Pleurothallis 43. 1859. —Specklinia spiculifera, Pridgeon & M.W. Chase, Lindleyana 16: 259. 2001. Pleurothallis acutissima Lindl., Fol. Orchid., Pleurothallis 43. 1859. Epiphyte, usually growing low on trees among bryophytes, minute, creeping, shadeloving; leaves 1–2 cm long, often purpletinged; flowers resupinate, pale purple with a darker purple lip; dorsal sepal 4–6 mm long. Rain forests, 100–200 m; Amazonas (Río Mawarinuma, near San Carlos de Río Negro). Amazonian Brazil. ◆Fig. 468. Pleurothallis steinbuchiae Carnevali & G.A. Romero, Novon 4: 90. 1994. Epiphyte, minute, shortly creeping; leaves 5–8 mm long; flowers resupinate, sepals and petals translucent maroon-pink; lip dark maroon; dorsal sepal 3.3–3.5 mm long. Rain forests, 500–600 m; Bolívar (Cerro Guaiquinima, Río Churún near Guarimba). Endemic. Pleurothallis stenocardium Schltr., Notizbl. Königl. Bot. Gart. Berlin 6: 123. 1914. Epiphyte, cespitose; leaves 7–10 cm long; flowers purplish; dorsal sepal 4–5.5 mm long. Cloud forests, ca. 2000 m; Bolívar (Roraima-tepui). Guyana. Pleurothallis suspensa Luer, Selbyana 7: 125. 1982. Pleurothallis penduliflora auct. non Kraenzl. 1905: sensu Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 4: 231. 1966. Pleurothallis revoluta auct. non (Ruiz & Pav.) Garay 1962: sensu Foldats in Lasser, Fl. Venez. 15(2): 377. 1970, pro parte, as for Guayanan material. Epiphyte, cespitose; leaves 6–8.5 cm long; flowers nonresupinate, purple; dorsal sepal ca. 8 mm long. Rain or cloud forests, 500– 1200 m; Bolívar (La Escalera to Cerro Venamo region). Suriname. Collections of Pleurothallis suspensa have been previously identified in the literature

522

O RCHIDACEAE

as P. penduliflora Kraenzl., an Andean species from Ecuador and Peru, or as P. revoluta (Ruiz & Pav.) Garay, a widespread, related but clearly distinct species. Pleurothallis tepuiensis Carnevali & I. Ramírez, Novon 3: 121. 1993. Pleurothallis pulchella auct. non (H.B.K.) Lindl. 1825: sensu Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 65: 46. 1998. Epiphyte, cespitose; leaves 6–12(–14) cm long; flowers campanulate, hyaline or greenish with purplish veins; dorsal sepal 3.5–4 mm long. Cloud forests on tepui summits or slopes, 700–2200 m; Bolívar (Auyán-tepui, Kamarkawarai-tepui, Jaua-Sarisariñama massif, Kamarkawarai-tepui, Sierra de Maigualida). Endemic. ◆Fig. 461. Pleurothallis tepuiensis is very closely related to the Andean P. pulchella but is easily distinguished by its glabrous sepals, narrower petals (3.5 times longer than wide as opposed to 2.5–2.8 times longer than wide in P. pulchella), and rugose apical portion of the lip (smooth in P. pulchella). Pleurothallis pulchella, although known from the Eastern Cordillera southward to Peru and in Panama and Costa Rica, is not known from the Cordillera de Mérida in Venezuela. Pleurothallis tridentata Klotzsch, Allg. Gartenzeitung 8: 289. 1840. Epiphyte, cespitose; leaves 2.5–6.7 cm long; flowers resupinate, yellow or green-yellow; dorsal sepal 3.5–6 mm long. Rain forests, 500–600 m; Bolívar (La Escalera to Cerro Venamo region). Aragua, Carabobo, Distrito Federal, Falcón, Lara, Mérida, Miranda, Yaracuy; Guyana, Colombia, Peru. Pleurothallis trinitensis (Griseb.) Carnevali & G.A. Romero in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1145. 2000. —Pleurothallis sertularioides var. trinitensis Griseb., Fl. Brit. W. I. 609. 1864. Pleurothallis acutissima auct. non Lindl. 1856: sensu Foldats in Lasser, Fl. Venez. 15(2): 200. 1970. Pleurothallis spiculifera auct. non Lindl. 1859: sensu Dunst. & Garay, Venez. Orchid. Ill. 2: 303. 1961. Epiphyte, creeping; leaves 1–3.5 cm long; flowers rose-orchid, purple, or pale lavender;

lip maroon-purple; dorsal sepal 3–4.5 mm long. Rain forests, usually growing low on trees among bryophytes, 100–500 m; Bolívar (La Escalera to Cerro Venamo region, Macizo del Chimantá, Río Canaracuni, Salto Angel, Sierra de Lema), Amazonas (Río Cataniapo). Miranda; Trinidad-Tobago. Guyana, Suriname, French Guiana. ◆Fig. 467. Pleurothallis velaticaulis Rchb. f., Linnaea 22: 824. 1850. —Stelis velaticaulis, Pridgeon & M.W. Chase, Lindleyana 16: 267. 2001. Pleurothallis velatipes Rchb. f., Linnaea 22: 828. 1850. Epiphyte or subterrestrial, cespitose or shortly creeping; leaves 12–20 cm long; flowers resupinate, yellow or greenish; dorsal sepal 5–6 mm long. Cloud forests, 1900–2000 m; Amazonas (Sierra de la Neblina). Mérida, Táchira, Trujillo, Zulia; Colombia, Ecuador, Peru. Pleurothallis vittariifolia Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19: 26. 1923. —Specklinia vittariifolia, Pridgeon & M.W. Chase, Lindleyana 16: 259. 2001. Pleurothallis pertenuis C. Schweinf., Bot. Mus. Leafl. 3: 83. 1935. Epiphyte, minute, cespitose, shade-loving; leaves 1.5–3 cm long; flowers resupinate, campanulate, orange; dorsal sepal 4–5 mm long. Rain forests, 100–200 m; Delta Amacuro (Río Amacuro), Bolívar (La Escalera to Cerro Venamo region). Costa Rica, Panama, Guyana. ◆Fig. 465. Pleurothallis wilsonii Lindl., Ann. Mag. Nat. Hist. ser. 3, 1: 326. 1858. —Acianthera wilsonii, Pridgeon & M.W. Chase, Lindleyana 16: 247. 2001. Epiphyte or lithophyte, minute, creeping, shade-loving; leaves 1.5–5 cm long, often purple below and bronzy green-purple above; flowers resupinate, often cleistogamous, greenish yellow with purple streaks; dorsal sepal 3–4 mm long. Rain forests, 500–600 m; Bolívar (Altiplanicie de Nuria). Aragua, Distrito Federal; West Indies, Ecuador. Pleurothallis yauaperyensis Barb. Rodr., Vellosia ed. 2, 1: 116. 1891. —Acianthera yauaperyensis, Pridgeon & M.W. Chase, Lindleyana 16: 247. 2001.

Pleurothallis 523

Pleurothallis consimilis Ames, Orchidaceae 7: 116. 1922. Epiphyte or lithophyte, creeping, shadeloving; leaves 2–5 cm long, often purple abaxially; flowers resupinate, campanulate; sepals hyaline with purple spots, petals hyaline with purple veins, lip dull red with dark red base and callus; dorsal sepal 5.3–7 mm long. Rain forests, 900–1000 m; Bolívar (La Escalera to Cerro Venamo region), Amazonas (road between Yavita and Pimichín). Trinidad-Tobago, Amazonian. Brazil. Pleurothallis zephyrina Rchb. f., Bonplandia (Hanover) 3: 71. 1855. —Humboldtia zephyrina (Rchb. f.) O. Kuntze, Revis. Gen. Pl. 2: 668. 1891. —Trichosalpinx zephyrina (Rchb. f.) Fig. 453. Pleurothallis kerrii

Fig. 455. Pleurothallis pemonum

Luer, Phytologia 54: 398. 1983. Pleurothallis setigera Lindl., Fol. Orchid., Pleurothallis 38. 1859. Epiphyte, often growing among bryophytes, minute, cespitose; leaves 1–2.5 cm long; flowers resupinate, pale green or yellowish; dorsal sepal 7–8.5 mm long. Cloud forests, 1600–1800 m; Bolívar (Auyán-tepui). Lara, Mérida, Táchira, Trujillo; Colombia, Ecuador, Peru. Pleurothallis sp. A Minute, shortly creeping epiphyte, 2.5-3.5 cm tall counting the inflorescences; flowers hyaline with pinkish veins; dorsal sepal 2.5 mm long, 1.5 mm wide. Rare in rain forests, 120–150 m, Bolívar (vic. La Escalera to Cerro Venamo region). Endemic. Fig. 454. Pleurothallis hexandra

Fig. 456. Pleurothallis deborana

524

O RCHIDACEAE

Fig. 457. Pleurothallis holstii

1 cm

Fig. 458. Pleurothallis arenicola

Fig. 459. Pleurothallis moritzii

Pleurothallis 525

Fig. 460. Pleurothallis corniculata

Fig. 461. Pleurothallis tepuiensis

Fig. 462. Pleurothallis lanceana

526

O RCHIDACEAE

Fig. 463. Pleurothallis miqueliana

Fig. 464. Pleurothallis samacensis 1 cm

Fig. 465. Pleurothallis vittariifolia

Fig. 466. Pleurothallis lappiformis

Pleurothallis 527

Fig. 467. Pleurothallis trinitensis

Fig. 468. Pleurothallis spiculifera

Fig. 469. Pleurothallis grobyi

Fig. 470. Pleurothallis elvirana

528

O RCHIDACEAE

Fig. 471. Pleurothallis granitica

Fig. 472. Pleurothallis morilloi

Fig. 473. Pleurothallis sclerophylla

Polystachia 529

112. POLYCYCNIS Rchb. f., Bonplandia (Hanover) 3: 218. 1855. [Subtribe Stanhopeinae]. by Gustavo A. Romero-González Cespitose, robust epiphytes, or rarely terrestrial herbs. Stem modified into ovoid to subcylindric pseudobulbs of 1 internode, when young entirely concealed by distichous, scarious sheaths, eventually naked. Leaves 1–3, lanceolate, acute, plicate, articulate, short-petiolate, the petiole sometimes costate. Inflorescences 1– several, lateral, arising from the base of the pseudobulbs, erect to pendent, racemose, many-flowered; peduncle densely pubescent. Flowers resupinate, showy; floral bracts concave, narrowly ovate, much shorter than the densely pubescent, pedicellate ovary. Sepals and petals membranous, spreading to reflexed; sepals subequal, narrowly ovate, the lateral sepals sometimes slightly oblique; petals linear and acuminate to very narrowly obovate. Lip subsessile to conspicuously clawed, divided into a hypochile and an epichile; hypochile narrowly to broadly winged, pubescent or not, often with a central, axial callus, sometimes with a basal, bifurcate callus; epichile adnate to the abaxial surface of the hypochile, linear, linearobtriangular, or subtriangular, sometimes basally pilose. Column elongate, slender, terete, arched, apically clavate and abruptly winged or auriculate, footless; anther terminal, operculate, incumbent, 1-locular; pollinarium with 2 yellow pollinia, narrowly obovate, cleft, cartilaginous, and dorsoventrally flattened, the tegula long, linear-obtriangular, the viscidium subtriangular. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 9 species, 2 in Venezuela, 1 of these in the flora area. Polycycnis surinamensis C. Schweinf., Bull. Torrey Bot. Club 75: 224. 1948. Epiphyte; inflorescence pendent, to 12flowered; flowers yellow with maroon spots;

pedicel pilose. Wet forests, 500–1500 m; Bolívar (Altiplanicie de Nuria, summit of La Escalera), Amazonas (Cerro Duida). Guyana, Suriname, French Guiana. ◆Fig. 474.

Fig. 474. Polycycnis surinamensis

530

O RCHIDACEAE

113. POLYOTIDIUM Garay, Bot. Mus. Leafl. 18: 105. 1958. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes or twig epiphytes, erect or pendulous, mostly sun-loving. Roots many, elongate, wiry. Rhizomes short, creeping, appressed to substrate, or ± ascendent; pseudobulbs spindle-shaped to cylindric, apically 1-foliate, enveloped when young by several imbricate, tubular, scarious sheaths that are eventually deciduous; leaves terete or subterete, articulate, erect, linear-subulate, fleshy-coriaceous. Inflorescences originating from pseudobulb base, racemose, lax, successively flowered, occasionally producing 1 or 2 branches at the base of the rachis; peduncle and rachis terete, peduncle remotely few-bracteate, rachis straight; floral bracts much shorter than the pedicellate ovary. Flowers resupinate, spreading, short-lived, showy due to bright orange color; pedicellate ovary elongate, terete. Perianth segments not widely spreading, subfleshy; dorsal sepal free, concave, elliptic or elliptic-ovate, forming a hood over the column and lip; lateral sepals connate about or above middle into a concave synsepal with divergent apices, somewhat saccate at base; petals subsimilar to dorsal sepal, at first parallel to column and enclosing it, later with spreading apices. Lip erect, sessile, rigidly attached to base of column, divided into hypochile (basal area of a complex lip) and epichile (distal area of a complex lip); hypochile saccate, basally with a pair of fleshy swellings, the opening completely covered by the column; epichile flat, cuneate to fan-shaped, subpapillate, basally with a retrorse callus. Column erect, footless, arched, apically dilated, with 4 wings; basal wings originating from the sides of the column with their parallel sides enclosing the stigma, the lower margins completely connate to form a shallow, cup-like cavity immediately beneath the stigmatic surface; the apical wings originating from the column apex and enclosing the basal part of the anther; anther dorsal, operculate, 1locular, elongate, rostrate, inserted in middle of column; pollinia 2, subglobose-obovoid, waxy, white, tegula oblong, ca. 3 times longer than pollinia, ca. 4 times longer than wide, viscidium small, strap-like; clinandrium conspicuous, with entire margins; rostellum conspicuous, transverse, highly bifid; stigma ventral, elliptic, very concave. Colombia, Venezuela, Brazil; 1 species. Polyotidium huebneri (Mansf.) Garay, Bot. Mus. Leafl. 18: 105. 1958. —Hybochilus huebneri Mansf., Repert. Spec. Nov. Regni Veg. 36: 61. 1934. Epiphyte or twig epiphyte; pseudobulbs 1–2 cm long; leaves 5–20 cm long, pale green, sometimes freckled with darker green; inflorescence ± equal to somewhat longer than the subtending leaf, the peduncle at first erect and later bending downward to become almost horizontal; rachis, floral bracts, pedicellate ovary, and flowers bright orange; flowers variable in size, 4–11 mm diameter, campanulate. Savannas, open places in rain forests, 100–200 m; Amazonas (Río Autana, San Carlos de Río Negro-Solano). Amazonian Colombia and Brazil. ◆Fig. 475.

Fig. 475. Polyotidium huebneri

Polystachia 531

114. POLYSTACHYA Hook., Exot. Fl. 2: t. 103. [1824] 1825. [Subtribe Polystachynae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes or rarely lithophytes, erect, sun- to shade-loving. Rhizome abbreviate, thus forming cespitose plants. Stems generally abbreviate, rarely elongate, basally thickened into homoblastic pseudobulbs enveloped by the leaf sheaths, which eventually remain naked after the fall of the leaf-sheaths. Leaves conduplicate, articulate with its sheaths, generally 2–5 borne distichously over the pseudobulb, the sheaths usually connivent into a ± well-developed pseudostem; the blades linear-oblong or linear-elliptic, less commonly elliptic or obovate, basally attenuated into a ± well-developed pseudopetiole. Inflorescences terminal, solitary, a raceme or a panicle of congested, erect or ascending racemes, longer or shorter than leaves; peduncle clothed by tubular sheaths, compressed, erect or arched; rachis abbreviate or subelongate, bearing an apically congested raceme of successive flowers, pubescent or glabrous; floral bracts usually small, subfleshy; pedicellate ovary relatively short. Flowers fleshy, minute to medium-sized, nonresupinate, frequently globose or saccate, fragrant or inodorous. Sepals wider than petals, usually broadly elliptic or ovate, acute to rounded, apiculate. Lateral sepals oblique and adnate to column foot, frequently forming a prominent mentum, commonly much larger than the dorsal; petals usually oblong or narrowly obovate, generally much narrower than sepals. Lip articulate with column foot, usually 3-lobed; disk with a conspicuous callus, frequently covered with multiseptate trichomes (mealy hairs or pseudopollen) or wax, which could extend to the lobes or wholly cover the lip surface. Column short, basally produced into a variously developed column foot, exauriculate, anther terminal incumbent, 1-locular or imperfectly 2-locular; pollinia 4, waxy, with a small but definite tegula and a viscidium; stigma ventral. Capsule oblong-ellipsoid. Tropical Africa and Asia, U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, northern Argentina; 100–200 species, 4 in Venezuela, all in the flora area. Key to the Species of Polystachya 1. 1. 2(1). 2.

3(2). 3.

Columnar foot about 1/2 the length of the lip; mentum of lateral sepals very prominent ........................................................................... P. concreta Columnar foot < 1/3 the length of the lip; mentum of the lateral sepals not prominent ............................................................................................... 2 Central lobe of the lip transversely oblong-elliptic, wider than its length, overlapping with the lateral lobes ...................................... P. cavanayensis Central lobe of the lip not transversely oblong-elliptic and as long as or longer than its width, not overlapping with the lateral lobes ................................................................................................................ 3 Leaves 15–30 times longer than wide; lip as long as wide or wider than long, lateral lobes in the basal 1/2 ........................................... P. amazonica Leaves 4–10 times longer than wide; lip longer than wide (1.5–2 times), lateral lobes in the basal 1/3 ........................................................... P. foliosa

532

O RCHIDACEAE

Polystachya amazonica Schltr., Beih. Bot. Centralbl. 42(2): 77. 1925. Polystachya stenophylla Schltr., Beih. Bot. Centralbl. 42(2): 77. 1925. Epiphyte, small-sized; leaves narrow; perianth segments greenish or yellowish; lip white. Humid forests, 50–100 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Colombia, Guyana, Ecuador, Brazil.

Epiphyte; inflorescences racemes; flowers greenish or yellowish. Forests, shrublands, near sea level to 700(–1000) m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Paraguay.

Polystachya cavanayensis Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 6: 378. 1976. Epiphyte, small; flowers greenish; lateral lobes of lip overlapping the central one. Rain forests, 900–1300 m; Bolívar (Gran Sabana). Apure, Barinas, Portuguesa, Táchira. Polystachya concreta (Jacq.) Garay & Sweet, Orquideología 9: 206. 1974. —Epidendrum concretum Jacq., Enum. Syst. Pl. 30. 1760. Onychium flavescens Blume, Bijdr. 325. 1825. —Polystachya flavescens (Blume) J.J. Sm., Orch. Java. 284. 1905. Epiphyte; flowers greenish with a long columnar foot. Evergreen, seasonally dry to rain forests, 50–1200 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; U.S.A., Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Madagascar, Ceylon, India, Indonesia, Phillippines, Java. ◆Fig. 476. Polystachya foliosa (Hook.) Rchb. f. in Walp., Ann. Bot. Syst. 6: 640. 1863. —Stelis foliosa Lindl., Ann. Nat. Hist. 2: 330, t. 17. 1839.

Fig. 476. Polystachya concreta

115. PONTHIEVA R. Br. in W.T. Aiton, Hortus Kew ed. 2, 5: 199. 1813. [Subtribe Cranichidinae]. by Germán Carnevali, Ivón M. Ramírez-Morillo, and Carlos A. Vargas Terrestrial or muscicolous on soil, rocks, or branches, erect, cespitose, glabrous or variously pubescent or pilose, mostly shade-loving. Roots fasciculate, villous, fleshy or ± fibrous. Stems very short, cylindric. Leaves convolute, membranous, not articulate, mostly basal and disposed in rosette, sessile or variously petiolate, erect or spreading, ovate, elliptic to lanceolate, glabrous to pilose. Inflorescence terminal from central rosette, solitary, racemose; peduncle terete, erect, longer than leaves, remotely few-sheathed, pubescent or glandular-pilose; rachis lax- or ± dense-flowered, erect, pubescent as in peduncle; floral bracts conspicuous but always shorter

Prescottia 533

than pedicellate ovary; pedicellate ovary thin, elongate, terete, variously pubescent. Flowers nonresupinate, spiral on the rachis, suberect to spreading, mostly inconspicuous, with widely spreading perianth segments, short-lived; perianth segments membranous, mostly white but sometimes greenish or yellowish, often with purple or maroon spots, especially on lateral sepals; sepals pubescent, especially outside; dorsal sepal free, usually smaller and less showy than the laterals; lateral sepals free or variously connate; petals adnate much above the base to the column, asymmetrical, usually clawed, glabrous or more commonly pubescent outside or marginally, forming a lip-like structure. Lip smaller and fleshier than the other perianth segments, long clawed but the claw totally or partially adnate to the ventral face of column, so that lip appears to be inserted on the upper 1/2 of the column; free portion erect or ± spreading, simple or variously 3-lobed, flat, concave, or excavate, glabrous or pubescent; disk variously callose, rarely ecallose. Column erect, fleshy, cylindric, dilated in the upper 1/3; anther dorsal, erect, 2-locular; pollinia 4, clavate, brittle, with small terminal caudicles, with a small hamuli (pollinium stalk derived from the apex of the rostellum), viscidium terminal, small; clinandrium short; rostellum horizontal, subequal to anther, posteriorly concave; stigma terminal or subventral, entire. Capsules ovoid or ellipsoid. U.S.A. (Carolina, Florida, Louisiana, Virginia), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia, northern Argentina; ca. 30 species, 8 in Venezuela, 2 of these in the flora area. Key to the Species of Ponthieva 1.

1.

Lip at base conspicuously angled or lobed, with a basal and an apical callus; petals transversely oblong, oblique, with the apex rounded or truncate .........................................................................................P. diptera Lip simple with only a basal callus; petals obliquely obtriangular, with the apex acute ................................................................................ P. ovatilabia

Ponthieva diptera Linden & Rchb. f., Bonplandia (Hanover) 2: 278. 1854. Muscicolous or terrestrial, 40–100 cm tall in flower, shade-loving; foliar blades 5–14 × 2–5.8 cm; flowers yellowish with purple or carmine spots, lateral sepals 6.3–12 mm long. Cloud forests, 1300–1500 m; Bolívar (La Escalera to Cerro Venamo region). Aragua; Cuba, Jamaica, Colombia, Ecuador, Peru.

Ponthieva ovatilabia C. Schweinf., Bot. Mus Leafl. 19: 211. 1961. Muscicolous or terrestrial, 40–90 cm tall in flower, shade-loving; foliar blades 7–12 × 2.2–3.8 cm; flowers with sepals and petals greenish white; lip white, lateral sepals 5.5– 8 mm long. Cloud forests, 700–1300 m; Bolívar (La Escalera to Cerro Venamo region), Amazonas (Sierra de la Neblina). Guyana. ◆Fig. 477.

116. PRESCOTTIA Lindl. in Hook., Exot. Fl. 2: pl. 115. 1824. [Subtribe Prescottiinae]. Decaisnea Brongn. in Duperrey, Voy. Monde (Phan.) 192, t. 39. 1829. —Decaisnea Lindl. in Wall. & Benth., Numer. List no. 7388. 1831. Galeoglossum A. Rich. & Galeotti, Ann. Sci. Nat. Bot. sér. 3, 3: 31. 1845. by Germán Carnevali, Ivón M. Ramírez-Morillo, and Carlos A. Vargas Terrestrial or humicolous herbs, sublithophytes, rarely epiphytes, cespitose, acaulescent or very shortly caulescent, glabrous or rarely pubescent throughout.

534

O RCHIDACEAE

Fig. 477. Ponthieva ovatilabia

Fig. 478. Prescottia oligantha

Roots fleshy, often tuberous, fasciculate, usually villous. Leaves membranous to subfleshy, convolute, rosulate or rarely cauline, sometimes only 1 present, usually erect, rarely appressed to the substrate, variously petiolate or sometimes subsessile; petiole channeled. Inflorescences terminal, erect, always longer than leaves, simple, spike-like; peduncle straight, often thick, variously covered with bract-like sheaths; rachis straight, densely many-flowered; floral bracts ± equal to the pedicellate ovary; pedicellate ovary thick, fusiform to clavate, often longer than the sepals. Flowers inconspicuous, nonresupinate, opening successively on the spike, erect, greenish to reddish, short-lived; sepals basally connate to form a short cup, spreading or reflexed, thin; petals narrower than the sepals, basally adnate to sepal cup, often circinate. Lip fleshy, slipper-like, galeate, or cochleate, often enclosing the column, at base with 2 retrorse auricles, clawed; claw adnate to sepal cup; disk ecallose. Column erect, short, fleshy, apically winged, basally produced into a short foot which is confluent with the claw of the lip; anther erect, dorsal; pollinia 4, granular, soft and mealy; without tegula or caudicles, viscidium small; clinandrium erect, acuminate, its margins connate with the rostellum; rostellum laminar, soft, emarginate, parallel to the anther and ± of the same length; stigma entire, ventral.

Prescottia 535

U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina, Uruguay; ca. 25 species, 5 in Venezuela, all in the flora area. Key to the Species of Prescottia 1. 1. 2(1). 2. 3(2).

3.

4(3). 4.

Leaves linear, not petiolate ................................................ P. auyantepuiensis Leaves elliptic, ovate, or ovate-elliptic, never linear; petioles always present, variously developed ................................................................. 2 Petioles inconspicuous, to 2.5 cm long; blades < 5 cm; lip 1–2 mm long ................................................................................................... P. oligantha Petioles (3–)5–25(–35) cm long; blades (4.5–)6–16 cm long; lip 2–7 mm long ......................................................................................................... 3 Upper 1/2 of lip longitudinally carinate; leaves fleshy when fresh, thinly chartaceous when dry; petiole, peduncle, rachis, and bracts dull red, brownish red, or pink ................................................................... P. carnosa Upper 1/2 of lip not longitudinally carinate; leaves thinly fleshy or membranous when fresh, membranous when dry; petiole, peduncle, rachis, and bracts green or pink ........................................................................ 4 Lip ca. 4 mm long ........................................................................ P. stachyodes Lip ca. 2 mm long ........................................................................ P. tepuyensis

Prescottia auyantepuiensis Carnevali & G.A. Romero in G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1145. 2000. Prescottia orchioides auct. non Lindl. 1845: sensu Foldats in Lasser, Fl. Venez. 15: 377, fig. 139. 1969; Dunst. & Garay, Venez. Orchid. Ill. 3: 272. 1965; Dunst. & Garay, Orchids Venez. Ill. Field Guide 871. 1979. Muscicolous herb ca. 35 cm tall, sun-loving; petiole, peduncle, rachis, fruits, and bracts green; lip ca. 2 mm long. Open places on tepui summits, 1800–1900 m; Bolívar (Auyán-tepui). Endemic. Prescottia carnosa C. Schweinf., Fieldiana, Bot. 28: 173, fig. 28. 1951. Muscicolous herb 20–50 cm tall, sun- or shade-loving; petiole, peduncle, rachis, fruits, and bracts dull red, brownish red, or pink. Forests, shrublands, tepui summits, (1400–)2200–2800 m; Bolívar (Auyán-tepui, Ilú-tepui, Kamarkawarai-tepui, Kukenántepui, Macizo del Chimantá, Murisipántepui, Roraima-tepui), Amazonas (Cerro Sipapo). Guyana (Mount Roraima).

Prescottia oligantha (Sw.) Lindl., Gen. Sp. Orchid Pl. 454. 1840. —Cranichis oligantha Sw., Prodr. 120. 1788. Muscicolous herb 8–40 cm tall, sun-loving, petiole, peduncle, rachis, fruits, and bracts green. Savannas, open rocky places, 900–1600 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Gran Sabana, Río Abapó, Sororopán-tepui). Anzoátegui, Aragua, Portuguesa; U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Guyana, Ecuador, Peru, Brazil, northern Argentina. ◆Fig. 478. Prescottia oligantha has wide floral and vegetative variation throughout its distribution, and is probably a complex including more than one species. Prescottia stachyodes (Sw.) Lindl., Edwards’s Bot. Reg. 22: sub t. 1916. 1836. —Cranichis stachyodes Sw., Prodr. 120. 1788. Muscicolous herb (30–)50–100 cm tall, shade-loving; petiole, peduncle, rachis, fruits, and bracts green, dull red, brownish red, or pink. Rain or cloud forests, (200–) 1200–2500 m; Bolívar (Auyán-tepui, Macizo

536

O RCHIDACEAE

del Chimantá, Ptari-tepui, Río Caura, near Santa Elena de Uairén), Amazonas (Cerro Aracamuni, Cerro Aratitiyope, Cerro Marahuaka). Anzoátegui, Aragua, Carabobo, Distrito Federal, Mérida, Miranda, Monagas, Yaracuy; southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, southern Brazil, northern Argentina. Prescottia stachyodes has wide floral and

vegetative variation throughout its distribution, and is probably a complex including more than one species. Prescottia tepuyensis Carnevali & C.A. Vargas, Lindleyana 11: 236. 1996. Herb 30 cm tall; petiole, peduncle, rachis, fruits, and bracts green. Open places, 1200– 1400 m; Amazonas (Cerro Aracamuni, Cerro Marahuaka). Endemic.

117. PROSTHECHEA Knowles & Westc., Fl. Cab. 2: 111. 1838. [Subtribe Laeliinae]. Epidendrum L., Gen. Pl. 272. 1737, pro parte. Anacheilium Hoffmanns., Verz. Orchid. 21. 1824. Encyclia Hook., Bot. Mag. 55: t. 2831. 1828, pro parte. Hormidium Lindl. ex Heynh., Nom. Bot. Hort. 880. 1841. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, lithophytes, or sometimes subterrestrial herbs, usually erect, rarely pendulous, mainly sun-loving but often growing in shade. Rhizome short, creeping to subscandent, terete, rooting at internodes. Pseudobulbs present (excepting in one Central American species), heteroblastic (one internode), pear-shaped, fusiform, or sometimes cylindric to subspheric, laterally compressed or not, apically or subapically 1–3(–6)-foliate, basally enveloped by tubular, applicate, eventually deciduous sheaths. Leaves subcoriaceous to rigid-fleshy, conduplicate, articulate (with the exception of some West Indian species), usually flat but sometimes concave, basally sheathing, not petiolate, linear, oblong to elliptic or broadly elliptic, midvein not prominent. Inflorescences terminal, from the mature pseudobulb or rarely from the developing pseudobulb, racemes or panicles, rarely 1-flowered, basally often subtended by 1 or 2 spathes; peduncle and rachis smooth to verruculose, peduncle few to many articulations; bracts usually inconspicuous; rachis straight to zigzag. Flowers resupinate or not, fragrant, very small to large and showy, long-lived, sometimes cleistogamous; floral bracts usually shorter than pedicellate ovary, often inconspicuous, fleshy to membranous; pedicellate ovary cylindric to trigonous, often winged, smooth or verruculose, longer than column. Perianth segments fleshy or fleshymembranous, variously colored, widely spreading to subcampanulate; sepals free, similar or the laterals somewhat oblique, often dorsally carinate; petals usually similar to sepals. Lip free to base from column or ventrally connate in its basal 1/3–1/2, fleshier than the other perianth segments; blade unlobed and usually concave or convex, or 3-lobed, flat or somewhat convex, lateral lobes narrower than central lobe; disk glabrous or pubescent, almost ecallose to conspicuously callose; calli usually of several raised, longitudinal keels, or plate-like. Column erect, footless, straight or somewhat recurved, parallel to lip; anther operculate, incumbent, 2-locular; clinandrium 3-dentate, teeth conspicuous or not; pollinia 4, subreniform, with caudicles, rostellum transversely perpendicular to column; stigma ventral, semiorbicular, obcordate or transversely fiddle-shaped. Capsules 3-winged. Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, southern Brazil, Bolivia; 90–95 species, 18–19 in Venezuela, 7 of these in the flora area. Prosthechea as here circumscribed includes a number of taxa formerly included

Prosthechea 537

in a broad, polyphyletic concept of Encyclia. However, it differs from Encyclia in its prominent inflorescence spathe, its fleshy, knob-like, obtuse or truncate midtooth appendage, which is not appressed to the anther, the 3-winged or -angled fruit, and the capsule suture covered by a strap of tissue. “Vegetative and floral material of Prosthechea species precipitate lumps of glycosides whithin the tissues when preserved in alcohol, further differentiating it from Encyclia material ...” (see Higgins, Phytologia 82: 370–383. 1997). Key to the Species of Prosthechea 1.

1.

2(1).

2.

3(2).

3.

4(3).

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

6.

Plants distinctly creeping with a branching rhizome, the distance between the pseudobulbs on the rhizome about as long or longer than the pseudobulbs themselves; lip 3-lobed; petals < 2.5 mm wide .... P. pygmaea Plants cespitose or shortly creeping, rhizome rarely branching, the distance between pseudobulbs on the rhizome usually much shorter than the pseudobulbs themselves; lip simple or shallowly lobed; petals 4 mm wide ........................................................................................................ 2 Lip concave or, if flat, then with smooth margins; disk and blade of lip with 5–17 longitudinal purple or pink stripes; petals acute or acuminate; inflorescences 1–5(–7)-flowered; the rachis about as long or longer than the peduncle ....................................................................... 3 Lip flat or convex with smooth or erose margin; disk and blade of lip without longitudinal purple or pink stripes; petals obtuse to rounded; inflorescences 2–30-flowered, when < 4-flowered, the rachis shorter than peduncle ................................................................................................. 5 Pseudobulbs 2-leaved; leaves 7–10 mm wide; inflorescences 1 or 2(3)-flowered; lip flat or shallowly concave with the longitudinal stripes only in basal 1/2, apex acute or obtuse; lip blade ca. 1 cm long .......... P. calamaria Pseudobulbs 1-leaved; leaves at least 12 mm wide; (usually > 15 mm wide); inflorescences (1)2–6-flowered; lip concave with the longitudinal stripes reaching the apex or almost, apex acuminate; lip blade ca. 2 cm long ......................................................................................................... 4 Plants flowering from the developing pseudobulb; stigmatic surface pandurate, plants from lowland rain forests, at elevations from near sea level to 600 m ......................................................................... P. aemula Plants flowering from the mature pseudobulb; stigmatic surface obcordate; plants from tepui highlands at elevations above 1500 m ............... P. jauana Dorsal sepal 17–21(–25) mm long; callus restricted to basal 1/2 of lip; stigmatic surface obcordate .......................................................... P. tigrina Dorsal sepal 9–13 mm long; callus occupying most of the lip blade; stigmatic surface transversely pandurate .................................................. 6 Pseudobulbs narrowly fusiform or cylindric, slightly compressed if at all, apically 2–4-leaved; leaves subdistichous due to the elongation of apical pseudobulb internodes; sepals usually verruculose without; plants from cloud forests at elevations usually above 1200 m ........ P. crassilabia Pseudobulbs ovoid-fusiform, strongly compressed, apically 1- or 2(3)leaved; leaves almost opposite due to telescoping of apical pseudobulb internodes; sepals always smooth without; plants from rain forests at elevations usually below 1000 m .................................................... P. vespa

538

O RCHIDACEAE

Prosthechea aemula (Lindl.) W.E. Higgins, Phytologia 82: 376. 1997. —Epidendrum aemulum Lindl., Edwards’s Bot. Reg. 22: t. 1898. 1836. —Epidendrum fragans var. aemulum (Lindl.) Barb. Rodr., Gen. Spec. Orchid. 2: 136. 1882. —Encyclia fragans subsp. aemula (Lindl.) Dressler & Pollard, Phytologia 21: 440. 1971. —Encyclia aemula (Lindl.) Carnevali & I. Ramírez, Monogr. Syst. Bot. Missouri Bot. Gard. 45: 1257. 1993. Epidendrum fragans auct. non Sw. 1788: sensu Foldats in Lasser, Fl. Venez. 15(3): 252. 1970; Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 4: 87. 1966. Epiphyte or rarely a lithophyte; leaves 7– 30 cm long; flowers non resupinate, white or cream with longitudinal purple stripes on the lip; dorsal sepal 20–30 mm long. Moist, semideciduous forests, granitic outcrops, open shrublands on sandy soil, near sea level to 300(–600) m; widespread in the flora area. Widespread in northern Venezuela; Panama, Trinidad-Tobago, eastern South America. ◆Fig. 479. Populations from some areas of Brazil, currently attributed to Prosthechea aemula, could belong to an as yet unnamed subspecies. The closest relative of this species, P. fragrans (Sw.) W.E Higgins, flowers from the mature pseudobulb and usually produces more flowers per inflorescence. The pseudobulbs of P. fragrans are also longer and narrowly ellipsoid. Prosthechea fragrans is known from Mexico to Panama and the West Indies. It enters South America at two points, the first the Pacific coast of Colombia and Ecuador, presumably as an extension range of the Panamanian populations; the second at Guyana, Suriname, French Guiana, and Brazil (Ceará), probably as an extension range of the West Indian populations of the species. Prosthechea calamaria (Lindl.) W.E. Higgings, Phytologia 82: 377. 1997. —Epidendrum calamarium Lindl., Edwards’s Bot. Reg. 24: misc. 88, no. 163. 1838. —Encyclia calamaria (Lindl.) Pabst, Orquídea (Niteroi) 29: 276. 1967 [1972]. —Anacheilium calamarium (Lindl.) Pabst, Moutinho & Pinto, Bradea 3: 183. 1981.

Epiphyte to 15 cm tall; sepals and petals greenish yellow or white with 3 purple streaks in lower half; lip white with 5–7 purple streaks on disk; dorsal sepal 10.5– 11.5 mm long. Rain forests, 500–600 m; southern Amazonas (upper Río Siapa). Suriname, French Guiana, southeastern and Amazonian Brazil. The Venezuelan material here treated as Prosthechea calamaria differs in several ways from the Brazilian populations and may represent a different variety or species. Prosthechea crassilabia (Poepp. & Endl.) Carnevali & I. Ramírez, comb. nov. —Epidendrum crassilabium Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 1, t. 102. 1838. —Encyclia crassilabia (Poepp. & Endl.) Dressler, Brittonia 13: 264. 1961. Epiphyte, rarely terrestrial, 25–35 cm tall; leaves 12–15 cm long, 2.5–5 cm wide; inflorescences 5–15-flowered; flowers buff yellow with maroon spots within; lip white or pale yellow; dorsal sepal 12–14 mm long. Cloud forests, 1200–1900 m; Bolívar (La Escalera to Cerro Venamo region, Kavanayén), Amazonas (Sierra Parima). Aragua, Distrito Federal, Falcón, Lara, Mérida, Yaracuy; Costa Rica, Panama, probably widespread in tropical South America. Prosthechea crassilabia is the name based on the oldest binomial in a complex of species around P. vespa characterized by slender, cylindrical, nonlaterally compressed pseudobulbs. These plants range from Costa Rica southward to Brazil and tend to grow in cloud forests at intermediate elevations. There are probably several taxa included in the specific circumscription adopted here and there are several names available (Epidendrum longipes Klotzsch, E. saccharatum Kraenzl., E. baculibulbon Schltr., etc.). None of these names have combinations in Prosthechea. The species apparently differ by combinations of plant and flower size and color. A revision of this complex is required to determine the number of taxa involved and their distributions and circumscriptions. The name Prosthechea vespa is here restricted to a Guayanan and Amazonian species with elliptic or ovate, strongly flattened pseudobulbs that usually grows at elevations below 1000 m.

Prosthechea 539

Fig. 479. Prosthechea aemula

Prosthechea jauana (Carnevali & I. Ramírez) W.E. Higgins, Phytologia 82: 378. 1997. —Encyclia jauana Carnevali & I. Ramírez, Lindleyana 9: 67. 1994. Lithophyte or low epiphyte; flowers whitegreen or creamy-white, lip white-green with 17–19 dark wine purple longitudinal lines; dorsal sepal 19–20 mm long. Dwarf, open tepui forests, 1500–2100 m; Bolívar (JáuaSarisariñama), Amazonas (Cerro Yaví). Endemic. ◆Fig. 480. Prosthechea jauana is similar to P. fragrans, but in P. jauana the pseudobulbs are shorter and elliptic, flowers are fewer per inflorescence, the lip is proportionally broader, the stigmatic surface is ovate-cordate, the callus is depressed, and the plants grow at higher elevations in rocky tepui associations. The populations of the P. fragrans–P. aemula complex from Guyana, Suriname, French Guiana, and Brazil (Ceará) that are discussed under P. aemula may belong here. Prosthechea pygmaea (Hook.) W.E. Higgins, Phytologia 82: 380. 1997. —Epidendrum pygmaeum Hook., Bot. Mag. 60: 5. 3233. 1833. —Epidendrum pygmaeum Hook., J. Bot. (Hooker) 1: 49.

Fig. 480. Prosthechea jauana

1834, nom. superfl. —Encyclia pygmaea (Hook.) Dressler, Brittonia 13: 264. 1961. Epiphyte or rarely lithophyte to 12 cm tall; flowers often cleistogamous, nonresupinate, perianth segments green or greenish yellow; lip white. Rain forests or cloud forests on tepui slopes, 400–1200 m; Bolívar (Altiplanicie de Nuria, Uei-tepui); Anzoátegui, Aragua, Miranda, Nueva Esparta, Táchira; widespread in the Neotropics. Prosthechea tigrina (Linden ex Lindl.) W.E. Higgins, Phytologia 82: 381. 1997. —Epidendrum tigrinum Linden ex Lindl., Orchid. Linden. 9. 1846. —Encyclia tigrina (Linden ex Lindl.) Carnevali & I. Ramírez, Ernstia 36: 9. 1986.

540

O RCHIDACEAE

Epidendrum pamplonense Rchb. f., Linnaea 22: 837. 1850. —Encyclia pamplonense (Rchb. f.) Carnevali, Phytologia 55(5): 228. 1984. —Prosthechea pamplonensis (Rchb. f.) W.E. Higgins, Phytologia 82: 379. 1997. Usually an epiphyte 20–100 cm tall, including the inflorescence; flowers frequently subcampanulate, fleshy, often cleistogamous; petals and sepals greenish or yellowish with purple, red, or brown spots; column white, lip white basally, the apical 3/4 rose or dull magenta; dorsal sepal 17–21 mm long. Locally common in cloud forests on tepui slopes or summits, 900–2200 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region, Macizo del Chimantá, Ptaritepui, Uaipán-tepui). Mérida, Táchira; Colombia. Occasional albino forms of Prosthechea tigrina have been found with entirely yellow, unspotted petals and a yellow lip with a magenta margin. Further study could demonstrate that P. pamplonense is a distinct spe-

cies, but a thorough study of the complex across its entire range is required. Plants reported from southeastern Brazil as P. tigrina probably belong in other taxa. Prosthechea vespa (Vell.) W.E. Higgings, Phytologia 82: 381. 1997. —Epidendrum vespa Vell., Fl. Flumin. Icon. 9: pl. 27. 1827 [1831]. —Encyclia vespa (Vell.) Dressler, Phytologia 21: 441. 1971. Epidendrum variegatum Hook., Bot. Mag. 59: t. 3151. 1832, non Sw. 1788, nec Koen. 1791. Epiphyte or lithophyte, 20–80 cm tall, very variable; pseudobulbs cylindric to suborbicular; inflorescences 5–30-flowered; sepals and petals green or yellowish with brown or maroon blotches or spots; lip white or pale green, often with purple stripes. Cloud or rain forests, near sea level to 1000 (–1200) m; widespread in Delta Amacuro, Bolívar, and Amazonas. Apure; Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia.

118. PSILOCHILUS Barb. Rodr., Gen. Spec. Orchid. 2: 273. 1882. [Tribe Triphoreae]. Pogonia Juss., Gen. Pl. 65. 1789, pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial or humicolous herbs, shade-loving. Rhizome terete, basally creeping and leafless, apically erect and leafy, rooting at internodes, usually branching below middle after flowering. Roots few, fleshy, pubescent. Leaves 2–5 per stem, distichous, distant and leaving portions of the stem naked, not articulated, membranous, plicate, sometimes silvery-variegated; petioles basally dilated and sheathing the stem. Inflorescences terminal, racemose or rarely branching, few–20-flowered, mostly shorter than the leaves; bracts usually conspicuous, boat-shaped, sometimes subfoliaceous. Flowers resupinate, subcampanulate, fugacious, successive or rarely several in simultaneous anthesis. Pedicellate ovary terete or subtrigonous, arched. Perianth segments membranous to thin-fleshy, greenish or whitish, often with purple lines or tinges, usually narrow-lanceolate to narrow-obovate; sepals free, subequal or the laterals oblique, dorsally carinate; petals usually narrower than sepals, often falcate. Lip completely free from column, as long as the petals, usually whitish with purple lines or zones; blade narrower toward base, broader and distinctly 3-lobed in the apical 1/2, rarely lobed below the middle; disk with several keels. Column elongate, terete, tapering toward base, usually arched; anther erect, terminal, articulated to column apex, 2-locular; pollinia 4, granulose. Stigma ventral. Capsules ellipsoid to obovoid, often ± trigonous. Central America, West Indies, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia; ca. 7 species, 3 in Venezuela, 2 of these in the flora area.

Psilochilus 541

Plants of Psilochilus resemble members of the Commelinaceae, and unlike most orchids, they show gregarious flowering. Key to the Species of Psilochilus 1. 1.

Dorsal sepal ca. 30 mm long; lip 3-lobed near or below the middle, midlobe broadly clawed ....................................................................... P. dusenianus Dorsal sepal 15–20 mm long; lip 3-lobed in the apical 1/3, midlobe not clawed ................................................................................. P. macrophyllus

Psilochilus dusenianus Kraenzl. ex Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 3: 274. 1965. Herb to 30 cm tall; flowers yellowish with some red in the lip. Cloud forests, 900–1600 m; Bolívar (La Escalera to Cerro Venamo region, Uaipán-tepui), Amazonas (Cerro Aracamuni). Amazonian Brazil. ◆Fig. 481. Psilochilus macrophyllus (Lindl.) Ames, Orchidaceae 7: 45. 1922. —Pogonia macrophylla Lindl., Ann. Mag. Nat. Hist. ser. 3, 1: 355. 1858. Pogonia physuraefolia Rchb. f., Ned. Kruidk. Arch. 4: 324. 1859. Herb to 30 cm; sepals and petals greenish white, lip greenish yellow or white with purple midlobe. Found in dense localized colonies in cloud forests, 800–1700 m; Bolívar (La Escalera to Cerro Venamo region, Macizo del Chimantá [Abacapá-tepui], Uei-tepui), Amazonas (Cerro Duida, Sierra de la Neblina). Aragua, Mérida, Sucre; Mexico, Guatemala, Costa Rica, West Indies, Guyana, Peru.

Fig. 481. Psilochilus dusenianus

542

O RCHIDACEAE

119. PSYCHOPSIS Raf., Fl. Tellur. 4: 40. 1836 [1838]. —Oncidium sect. Glanduligera Lindl., Fol. Orchid., Oncidium 2. 1855. [Subtribe Oncidiinae]. Papiliopsis E. Morren ex Cogn. & Marchal, Pl. Feuill. Omem. 2: sub t. 55. 1874; E. Morren, Belg. Nov. 1: 95. 1877. Psychopsiella Lückel & Braem, Orchidee (Hamburg) 33: 7. 1982. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, cespitose or shortly creeping, erect. Rhizome abbreviate, relatively thin; stems pseudobulbose-thickened; pseudobulbs heteroblastic (one internode), densely aggregated or closely creeping, orbicular, ancipitous (flattened with sharp edges), basally enveloped by several small, imbricate, scarious sheaths none of which develop foliar blades, apically 1-leaved. Leaves conduplicate, articulate, thickly coriaceous, basally shortly conduplicate pseudopetiolate; blade frequently red- or brown-spotted, surfaces with the texture of fine sand-paper. Inflorescences lateral, originating from pseudobulb base or from the axils of the sheaths, often with the same color pattern as the leaves, racemose or rarely with 1–2 branches, 3–10 times longer than leaves, erect; peduncle straight or flexuous, terete or laterally flattened in the upper 1/4; with several tubular sheaths; rachis terete or laterally flattened, much shorter than the peduncle, densely flowered; floral bracts inconspicuous, coriaceous, concave, broadly triangular; pedicellate ovary terete, longer than bracts. Flowers resupinate, large and showy or medium-sized, spreading, with widely spreading perianth segments, lasting 4–7 days, successive on the rachis, only 1 open at a time; perianth segments membranaceous, yellowish with dull orange or brownish zones, blotches, or mottles, rarely the whole flower yellowish, margins ± undulate; sepals free, dissimilar or similar in 1 species; dorsal sepal narrow-linear or linear-oblanceolate, rarely elliptic, about 2 times as long as the laterals or rarely ± equal to them; lateral sepals mostly showy, oblong-lanceolate to narrowly ovate-lanceolate, falcate; petals often similar to dorsal sepal and somewhat diverging from it in natural position. Lip spreading, showy, sessile and rigidly attached to column base, pandurate-obovate, with a deep constriction, which may sometimes be prolonged into a short, broad isthmus, deeply 3-lobed in the basal 1/3 due to constriction; lateral lobes semiquadrate, semiobovate, or semirounded; central lobe much larger than the laterals, transversely subquadrate, transversely elliptic, to suborbicular, apically rounded to emarginate; disk with a prominent, fleshy, tuberculate or crested, glabrous or minutely papillose callosity. Column relatively short, erect, ± geniculate, footless, apically with 2 well-developed, subquadrate or triangular-oblong wings with subentire or dentate to laciniate margins, each wing usually with 1 linear-subulate, elongate process above, capitate (thickened at the apex into small, blackish semispheric glands); anther terminal, operculate, incumbent, very convex, 1-locular; pollinia 2, waxy, obpear-shaped, longer to ± equal to the tegula, tegula short, broad, concave, viscidium broad, horseshoe-shaped; clinandrium short, truncate; rostellum transverse; stigma ventral, broad, suborbicular or subquadrate. Capsules oblong, attenuated toward base and apex, inconspicuously 3-edged. Costa Rica, Panama, Colombia, Venezuela, Trinidad-Tobago, Suriname, French Guiana, Ecuador, Peru, Brazil; 4 species, 1 or probably 2 in Venezuela, 1 of these in the flora area.

Pterichis 543

Fig. 482. Psychopsis papilio

Psychopsis papilio (Lindl.) H.G. Jones, J. Barbados Mus. Hist. Soc. 35(1): 32. 1975. —Oncidium papilio Lindl., Bot. Reg. 11: t. 910. 1825. —Mariposa. Psychopsis picta Raf., Fl. Tellur. 4: 40. 1836 [1838]. Epiphyte, sun- or shade-loving; pseudobulbs reddish; leaves 12–22 cm long; inflo-

rescence 60–120 cm long; dorsal sepal 7.5–13 cm long. Rain or low-elevation cloud forests, 800–1000 m; Bolívar (La Escalera to Cerro Venamo region). Aragua, Carabobo, Distrito Federal, Lara, Mérida, Miranda, Monagas, Trujillo; Panama, Colombia, Trinidad-Tobago, Suriname, Ecuador, Peru, northern Brazil. ◆Fig. 482.

120. PTERICHIS Lindl., Gen. Sp. Orchid. Pl. 444. 1840. [Subtribe Cranichidinae]. Acraea Lindl. in Benth., Pl. Hartw. 155. 1839 [1845], pro parte. by Germán Carnevali, Ivón M. Ramírez-Morillo, and Carlos A. Vargas Terrestrial, sun-loving, growing at high elevations. Rhizome short or lacking; roots few, fasciculate, tuberous. Leaves in a basal rosette, 1 or 2(3), basally attenuated into a channeled pseudopetiole, or absent during flowering (hysteranthous plants). Inflorescence terminal from center of the rosette, pubescent; peduncle erect, stiff, simple, pubescent, with several sheaths; sheaths tubular, distant, leaving most of the peduncle naked, lax- or dense-spiciform, bearing flowers in apical 1/4; floral bracts conspicuous, boat-shaped. Pedicellate ovary thick, pubescent, subequal to conspicuously longer than its bract. Flowers nonresupinate, lasting 1 or 2 days, successive or 2–4 in simultaneous anthesis. Perianth segments subfleshy, pubescent outside, usually reddish, brownish, or yellowish with red or brown streaks; sepals free or the laterals somewhat connate basally, subequal or the laterals broader, usually elliptic or lanceolate; petals often narrower than the sepals, straight or falcate. Lip free to base, fleshier than other perianth segments, concave in natural position, simple or apically lobed, often with verrucose-glandular structures at its margin,

544

O RCHIDACEAE

apical lobe usually recurved, lateral lobes partially embracing the column; disk pubescent or glandular. Column short and broad, footless; anther dorsal, erect; clinandrium small; rostellum horizontal; pollinia 4, clavate, brittle, with small terminal caudicles, with a small hamulus (pollinium stalk derived from the apex of the rostellum), viscidium terminal, small; stigma ventral, entire. Capsules oblong or ellipsoid. Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia; ca. 20 species, 5 in Venezuela, 1 of these in the flora area. This is basically a high-elevation, Andean genus whose occurrence in the Venezuelan Guayana is unusual. The species are poorly delimited and are in need of revision. Pterichis galeata Lindl., Gen. Sp. Orchid. Pl. 445. 1840. Herb to 40 cm; flowers brown-maroon or brownish purple. Open, boggy associations on tepui summits, 2500–2700 m; Amazonas (Cerro Marahuaka). Mérida, Táchira, Trujillo; Colombia, Ecuador, Peru, Bolivia. ◆Fig. 483. Some individuals of Pterichis galeata from the flora area have lobed petals, a character unknown elsewhere.

Fig. 483. Pterichis galeata

121. QUEKETTIA Lindl., Edwards’s Bot. Reg. 25: misc. 3. 1839. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Twig epiphyte, cespitose, erect to pendulous, sun-loving. Roots relatively long and wiry. Pseudobulbs very small, 1-foliate, laterally compressed, clothed by imbricating sheaths of which the innermost 1 or 2 may have foliar blades. Leaves terete to subterete, very rarely flat. Inflorescences axillary, shorter than to longer than leaves, racemose to paniculate; peduncle subterete, remotely few-bracteate, erect to arched; floral bracts concave. Flowers minute, resupinate or not. Perianth segments membranaceous, subparallel to apically spreading. Lateral sepals basally variously

Reichenbachanthus 545

connate; dorsal sepal free; petals subequal to dorsal sepal or broader. Lip adnate to column base, obovate, broadly rhombic to narrowly rhombic, ecallose or with a transverse keeled callus below the middle. Column short to relatively elongate, footless, apically with conspicuous wings; anther terminal, incumbent, operculate, 1locular; pollinia 2, cartilaginous, ovoid to ellipsoid, attached to a short, narrow tegula and to a small viscidium. Venezuela, Guyana, Suriname, French Guiana, Ecuador, Brazil; 1 species. Quekettia microscopica Lindl., Edwards’s Bot. Reg. 25: misc. 3. 1839. Leaves subterete; inflorescences shorter than leaves, racemose or with 1 or 2 lateral branches; flowers minute, bright yellow green; lip bright golden yellow with 2 small maroon marks near apex. Rain forests, 50– 500 m; Bolívar (Río Caura, Río Caroní, Río Cuyuní), Amazonas (Río Padamo). Monagas; Guyana, Suriname, Brazil. ◆Fig. 484. Fig. 484. Quekettia microscopica

122. REICHENBACHANTHUS Barb. Rodr., Gen. Spec. Orchid. 2: 164. 1882. [Subtribe Laeliinae]. Fractiunguis Schltr., Anexos Mem. Inst. Butantan, Secc. Bot. 1(4): 56. 1922. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes or rarely muscicolous on rocks, first erect, then arching, later pendulous, cespitose. Roots filiform. Rhizome very abbreviate, appressed to substrate; stems cylindric, basally attenuate and substipitate, apically 1-leaved, enveloped by several scarious, tubular, imbricate sheaths, proliferating at apex and forming branched, elongate, pendulous plants to 1 m long, ± fruticose when old. Leaves semiterete to subterete, articulate, narrowly linear-subulate, acute, straight or somewhat falcate. Inflorescences 1–3, flowering 1 at a time, terminal, much shorter than the leaves, fasciculate, 1–few-flowered; peduncle very abbreviate, enveloped by several imbricate, scarious sheaths; pedicellate ovary thin, ± arched; floral bracts inconspicuous. Flowers resupinate, inconspicuous, short-lived, often cleistogamous; perianth segments reflexed, membranous; sepals free, subequal or the laterals somewhat larger, ovate-elliptic or lanceolate, acute or acuminate; petals shorter and narrower than sepals. Lip fleshier than other perianth segments, rigid, basal margins connate with the basal portion of column forming a prominent cup or sac that projects into the ovary and is noticeable from the outside; free portion dilated into an articulate, very convex, twice-geniculate blade; blade 3-lobed, lateral lobes rounded or subtriangular, central lobe variously emarginate to bifid making the lip appear 4lobed; disk with 2- or 4-keeled central veins. Column erect, straight, ± elongate, with a prominent foot, apically ± dilated; anther terminal, operculate, incumbent, semiglobose, 2-locular; pollinia 4, waxy, laterally compressed, ovoid, with yellow caudicles; clinandrium with unequally 3-lobed margin; rostellum transverse; stigma ventral, pandurate. Capsules subspheric or ellipsoid.

546

O RCHIDACEAE

Costa Rica, Panama, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 species, 1 in Venezuela. Some authors consider Reichenbachanthus a synonym of Scaphyglottis Poepp. & Endl. Reichenbachanthus reflexus (Lindl.) Brade, Rodriguésia 1: 55. 1935. —Scaphyglottis reflexa Lindl., Edwards’s Bot. Reg. 25: misc. 20. 1839. —Hexisea reflexa (Lindl.) Rchb. f. ex Griseb., Fl. Brit. W. I. 623. 1864, non Rchb. f. 1877. Reichenbachanthus modestus Barb. Rodr., Gen. Spec. Orchid. 2: 165. 1882. Epiphyte 25–100 cm long; stems 3–17 cm long; leaves 10–30 cm long; perianth segments 4–6 mm long, sepals and petals pale green, lip white with purple or pink on the callus, column pale green with a dark purple anther. Rain or cloud forests, (50–)400–1400 m; Delta Amacuro (Río Cuyubini), Bolívar (widespread but most common in the basins of Río Caroní and Río Cuyuní), Amazonas (Cerro Aratitiyope, Cerro Duida, Cerro Huachamacari, Sierra Parima, Sierra de la Neblina). Aragua, Distrito Federal, Miranda, Sucre, Táchira; West Indies, Costa Rica, Panama, Guyana, Suriname, French Guiana, Ecuador, Peru, northern Brazil. ◆Fig. 485.

Fig. 485. Reichenbachanthus reflexus

123. RESTREPIOPSIS Luer, Selbyana 2: 199. 1978. [Subtribe Pleurothallidinae]. Restrepiella Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 4: 266. 1966, pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, cespitose, erect, usually inconspicuous. Rhizome abbreviate. Ramicauls 1-leaved apically, erect, terete, enveloped by several scarious, tubular sheaths that are glabrous (pubescent in one species). Leaves conduplicate, articulate, coriaceous to fleshy-coriaceous or subterete, linear-elliptic to suborbicular, sometimes attenuated into a pseudopetiole. Inflorescences originating from near the apex of the ramicaul without an annulus, 1-flowered, solitary or in fascicles; peduncle short, terete; floral bracts tubular; pedicel short or very elongate, always longer than the floral bract, articulate with the ovary; ovary terete, often ridged. Flowers inconspicuous, resupinate, usually greenish, yellowish, or hyaline, often with the veins of a darker hue or reddish or purplish; perianth segments membranous, usually widely spreading; sepals subequal or the dorsal broader, free or more often the laterals connate in the basal 1/2; petals subequal to the sepals or more commonly, smaller, frequently subparallel to the column. Lip articulate to the column foot, erect, fleshier than the other perianth segments, subsimple or 3-lobed below the middle; lateral lobes much smaller than the central, usually ± falcate, porrect (extended horizontally) and erect, ± embracing the column; central lobe usually flat or ± concave, ovate-oblong, oblong, or obovate; disk with 2 or 4 longitudinal keels.

Rodriguezia 547

Column erect or ± curved, subterete, elongate; anther apical, incumbent, operculate, 2- or imperfectly 4-locular; pollinia 4, waxy, pear-shaped or ± clavate, viscidium very small; clinandrium shallow, mostly entire; rostellum transverse; stigma ventral, small. Central America, Colombia, Venezuela, Ecuador, Peru, Bolivia; ca. 18 species, 4 in Venezuela, 1 of these in the flora area. Restrepiopsis is very similar and probably closely related to Octomeria R. Br., differing only in the pollinia number (8 in Octomeria). Restrepiopsis tubulosa (Lindl.) Luer, Selbyana 2: 200. 1978. —Pleurothallis tubulosa Lindl., Fol. Orchid. Pleurothallis 19. 1859. Pleurothallis viridula Lindl., Fol. Orchid., Pleurothallis 19. 1859. —Restrepiella viridula (Lindl.) Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 4: 268. 1966. —Restrepiopsis viridula (Lindl.) Luer, Selbyana 2: 200. 1978. Epiphyte 6.5–21 cm tall, erect; flowers green-yellow, often with 3 purple stripes in the dorsal sepal; dorsal sepal 6–10 mm long, lateral sepals free or connate only at the very base; lip with a pair of small lateral lobes at the basal 1/4. Cloud forests, 2000–2600 m; widespread in Bolívar and Amazonas. Aragua, Distrito Federal, Mérida, Táchira, Trujillo. Colombia, Ecuador. ◆Fig. 486.

Fig. 486. Restrepiopsis tubulosa

124. RODRIGUEZIA Ruiz & Pav., Fl. Peruv. Prodr. 115, t. 25. 1794. [Subtribe Oncidiinae]. Burlingtonia Lindl., Edwards’s Bot. Reg. 23: t. 1927. 1837. by Germán Carnevali and Ivón M. Ramírez-Morillo Usually twig epiphytes, inconspicuous to showy, erect to pendulous, cespitose to long-rhizomatous and creeping to ascendent or subscandent, usually sun-loving. Rhizome abbreviate to long and wiry, bearing pseudobulbs at variable distances, rooting at nodes. Pseudobulbs heteroblastic (one internode), short, ovoid or ellipticoblong, ancipital (flattened and with sharp edges) to subquadrangular, apically 1- or (rarely) 2-leaved, almost totally clothed by sheaths the innermost 1–4 of which may have foliar blades. Leaves conduplicate, articulate, coriaceous, usually oblong to oblong-elliptic, apically acute, obtuse to obliquely emarginate, the apical leaves often with a short, conduplicate pseudopetiole. Inflorescences 1–5, from pseudobulb base or from the axils of the sheaths, racemose, erect, arching to pendulous, few- to manyflowered, short- to long-pedunculate; peduncle terete, remotely sheathed; floral bracts inconspicuous; pedicellate ovary terete, elongate. Flowers ± showy, resupinate, distichous on the rachis, sometimes secund (all turned to the same side), open-

548

O RCHIDACEAE

ing successively or ± simultaneously, usually white with yellow on lip to red, yellow, or cream, sometimes with yellow, pink, purple, or red spots, bands, or blotches. Perianth segments membranous to subfleshy, basally parallel to column, apical 1/2 usually spreading; dorsal sepal free, basally concave and ± hooded on the column; lateral sepals variously connate into a conduplicate synsepal, usually geniculate to gibbous, when partially free, the apices of lateral sepals ± divaricate; petals similar to dorsal sepal but usually wider, often subparallel to column with straight or spreading apices. Lip larger than other perianth segments, basally clawed and parallel to the column, often basally produced into a gibbous sac or spur decurrent with ovary, blade usually obovate, flat to ± convex; disk with several parallel keels. Column erect, relatively short to ± elongate, subcylindric, apically dilated, often variously auriculate at apex, basally footless or almost; anther terminal or ± dorsal, incumbent, operculate, 1-locular; pollinia 2, waxy, ovoid to subglobose with a well-developed, linear to ± spatulate tegula and a small, strip-like viscidium; clinandrium shallow, entire; rostellum transverse; stigma ventral. Capsules ellipsoid. Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 30 species, 6 in Venezuela, 4 of these in the flora area. The species with large white and yellow flowers are pollinated by large bees while the reddish-flowered species are pollinated by hummingbirds. Key to the Species of Rodriguezia 1. 1.

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

Flowers pink, rose, or purple, ± secund on the rachis; lip 1–2 cm long ................................................................................................. R. lanceolata Flowers white, sometimes with rose or purple tinges or dots at or near the base of perianth segments and lip, distichous on the inflorescence; lip 3–6 cm long ............................................................................................ 2 Column 9–12 mm long, shorter than pedicellate ovary ............. R. batemanii Column 17–25 mm long, longer than pedicellate ovary ........................... 3 Perianth segments obtuse; leaves obtuse, 4–5 times longer than wide; lip base cucullate, unspreadable ..................................................... R. candida Perianth segments acute; leaves acute, 7–8 times longer than wide; lip base cuneate, spreadable .............................................................. R. leeana

Rodriguezia batemanii Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 41. 1836. Twig epiphyte; inflorescences arching to pendulous, 3–10-flowered; flowers relatively showy with dorsal sepal 16–35 mm long, sepals and petals white with some pink or purple toward the base; lip white with yellow on the disk and callus. Rain forests, 50–200 m; widespread in southern Amazonas; Amazonian Ecuador, Peru and Brazil. Rodriguezia candida Bateman ex Lindl., Edwards’s Bot. Reg. 23: sub t. 1927. 1837.

Twig epiphyte; inflorescences spreading or arching, 1–7-flowered; flowers showy with dorsal sepal 25–31 mm long, sepals and petals white, with some clear yellow at callus and center of disk. Rain forests, 200–500 m; Bolívar (La Escalera to Cerro Venamo region, upper Río Caura). Guyana, Brazil. Rodriguezia lanceolata Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 1: 219. 1798. Rodriguezia secunda H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 367, t. 92. 1815 [1816]. Twig epiphyte; inflorescences spreading to ± nodding; flowers showy with dorsal se-

Rudolfiella 549

pal 9–14 mm long, perianth segments pink to deep coral red. Rain forests, open shrublands, riparian forests, 50–900 m; Delta Amacuro (widespread), Bolívar (widespread), Amazonas (widespread in northern part, scattered in southern part). Widespread in northern Venezuela; Panama, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 487. Rodriguezia leeana Rchb. f., Gard. Chron. n.s. 20: 38. 1883. Twig epiphyte; inflorescences arching to pendulous; flowers relatively large and showy with dorsal sepal 25–40 mm long; perianth segments white flushed with variable amount of pink toward base; lip white with yellow on disk. Rain forests, 50–200 m; widespread in southern Amazonas. Suriname, Ecuador, Peru, Amazonian Brazil.

Fig. 487. Rodriguezia lanceolata

125. RUDOLFIELLA Hoehne, Arq. Bot. Estado São Paulo 2: 13. 1944. —Schlechterella Hoehne, Arq. Bot. Estado São Paulo 2: 13. 1944, non K. Schum. 1899. [Subtribe Lycastinae]. Bifrenaria Lindl., Gen. Sp. Orchid. Pl. 152. 1833, pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, cespitose or shortly creeping, conspicuous, erect. Rhizome abbreviate; pseudobulbs heteroblastic (one internode), clumped on the rhizome and ± overlapping, broadly ellipsoid to orbicular, strongly laterally compressed, apically 1leaved, basally clothed by scarious, ± imbricate, not leaf-bearing sheaths that eventually disintegrate, surface of young pseudobulb usually dark maroon spotted or blotched. Leaves plicate, convolute, articulate, coriaceous, erect or spreading, conspicuously petiolate; blade elliptic or oblong-elliptic, often maroon spotted; petioles shorter than blades, subterete, sulcate. Inflorescences lateral racemes, from the base of the pseudobulb, ± equal to the leaves, often purple-tinged; peduncle terete, erect, remotely few-sheathed, sheaths tubular, acute, often maroon-spotted; rachis subequal or shorter than peduncle, ± zigzag, erect or more often ± arching; floral bracts inconspicuous; pedicellate ovary terete, elongate. Flowers resupinate, ± showy, long-lasting, spiralled on the rachis, anthesis ± simultaneous; perianth segments coriaceous, yellow or orange-yellow with maroon or chestnut brown spots or blotches on the lower 1/2; sepals free, subequal, elliptic or oblong-elliptic; dorsal sepal hooded over the column; lateral sepals widely spreading, attached to the margins of the column foot; petals narrower than the sepals, basally attenuated, parallel to the column. Lip articulate to the apex of the column foot, paler and fleshier than sepals, long-clawed, the claw as long as the blade, above the claw sharply 3-lobed; claw pubescent, parallel to column, flattened, thickened toward apex, as long as the blade; lateral lobes spatulate or dolabriform, erect and ± embracing the column, retrorse when flattened; central lobe spreading or ± reflexed, broader than the laterals, suborbicular to transversely elliptic or subquadrate; disk parallel to column, with a

550

O RCHIDACEAE

conspicuous, tuberculate callosity. Column elongate, semiterete, curved, minutely pubescent, basally with a prominent foot, wingless; anther terminal, operculate, incumbent, imperfectly 2-locular; pollinia waxy, 4 in 2 subequal pairs, dorsoventrally compressed, ± superposed, tegula (pollen stalk derived from the column) split in 2 arms, viscidium broad, ± horse shoe-shaped; clinandrium shallow; rostellum transverse; stigma ventral, entire. Panama, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, most diverse in the Amazon Basin; ca. 6 species, 1 in Venezuela. Rudolfiella aurantiaca (Lindl.) Hoehne, Arq. Bot. Estado São Paulo 2(1): 13. 1944. —Bifrenaria aurantiaca Lindl., Edwards’s Bot. Reg. 22: t. 1875. 1836. —Schlechterella aurantiaca (Lindl.) Hoehne, Arq. Bot. Estado São Paulo 2(1): 13. 1944. Epiphyte, mostly sun-loving; leaves 13.5– 37 cm long; flowers with sepals 11–16 mm long, lip 9–15 mm long. Locally common in rain forests, 50–800(–1100) m; Delta Amacuro (Sacupana), Bolívar (Auyán-tepui, Canaima, La Escalera to Cerro Venamo region, lower Río Caura, Río Paragua, near Santa Elena de Uairén, Uaiparú), Amazonas (Cerro Morrocoy, Cerro Yutajé, near Puerto Ayacucho, Río Ararí, Río Baría, Río Sipapo, Tamanaco). Monagas, Sucre; Amazonian Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil. ◆Fig. 488.

Fig. 488. Rudolfiella aurantiaca

126. SACOILA Raf., Fl. Tellur. 2: 86. 1836 [1837]. [Subtribe Spiranthinae]. Spiranthes Rich., De Orchid. Eur. 20, 28, 36. Aug.–Sept. 1817, nom. cons., pro parte. Stenorrhynchos, Rich. ex Spreng., Syst. Veg. 3: 677. 1826, pro parte. by Germán Carnevali, Ivón M. Ramírez-Morillo, and Carlos A. Vargas Terrestrial herbs. Roots fasciculate, fleshy, hairy, tuberous. Stem erect, vaginate or bracteolate. Leaves either basal or cauline, commonly absent at anthesis (in hysteranthous plants), cuneate, sessile. Inflorescence a terminal, ± dense, many-

Sacoila 551

flowered raceme, borne on a leafless plant. Flowers often showy, greenish to deep crimson, resupinate, large for the subtribe; ovary clavate to fusiform, with a short pedicel, somewhat twisted. Sepals free, subsimilar, connivent, usually lanceolate; dorsal sepal concave; lateral sepals decurrent on column foot, and together form a ± free-projecting, spur-like extension. Petals joined with dorsal sepal, basally decurrent on column foot. Lip sessile, from a cuneate base conduplicate, somewhat arched, margins on middle joined with sides of column, at base with linear thickenings. Column short, stout, basally extended in a long foot which is decurrent on sides of ovary, with the end protruding into a noticeable mentum; anther ovate, acute or acuminate, concave, much shorter than the rostellum; pollinia 4, soft, narrowly clavate, with a linear, sheath-like viscidium, covering the apex of the rostellum; rostellum rigid-linear-acicular, sharp-pointed; stigmas 2, terminal, confluent. Capsule elliptic. U.S.A. (Florida), southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina; 10 species, 1 in Venezuela. Most modern authors include this genus in a broadly defined Spiranthes or in its segregate, Stenorrhynchos. Sacoila lanceolata (Aubl.) Garay, Bot. Mus. Leafl. 28: 352. 1980. —Limodorum lanceolatum Aubl., Hist. Pl. Guiane 821. 1775. —Stenorrhynchos lanceolatum (Aubl.) Rich ex Spreng., Syst. Veg. 3: 710. 1826. —Spiranthes lanceolata (Aubl.) León, Contrib. Ocas. Mus. Hist. Nat. “De La Salle” 8: 358. 1946. Satyrium orchioides Sw., Prodr. 118. 1788. —Spiranthes orchioides (Sw.) A. Rich., Sagra, Hist. Phys. Cuba, Bot. Pl. Vasc. 252. 1850. Terrestrial, 30–90 cm tall at anthesis, sun-loving; leaves 2 or 3, 10–40 cm long, commonly developing after flowering; flowers variable in color, greenish white to brick red or deep crimson; dorsal sepal 13–22 mm long. Open places, 400–500 m; Bolívar (lower slopes of Auyán-tepui). Widespread and common in Venezuela and Neotropics. ◆Fig. 489.

Fig. 489. Sacoila lanceolata

552

O RCHIDACEAE

127. SARCOGLOTTIS C. Presl, Reliq. Haenk. 1: 95. 1827. [Subtribe Spiranthinae]. Spiranthes Rich., De Orchid. Eur. 20, 28, 36. Aug.–Sept. 1817, nom. cons., pro parte. Pelexia Poit. ex Lindl., Bot. Reg. 12: sub t. 985. Jun 1826, nom. cons, pro parte. Veyretia Szlach., Fragm. Florist. Geobot. Supp. 3: 115. 1996, pro parte. by Germán Carnevali, Ivón M. Ramírez-Morillo, and Carlos A. Vargas Usually terrestrial herbs, more rarely subepiphytes, very variable in habit, minute to rather large. Roots fasciculate, fleshy, tuberous. Leaves convolute, subfleshy, basal, rosulate, subsessile to almost petiolate, sometimes variegated or bicolorous. Inflorescences erect, slender to stout, variously bracteate, terminated by a few- to many-flowered racemes or spikes. Flowers fleshy, resupinate, mediumsized to relatively large in size, sometimes rather showy; ovary sessile ± fusiform, scarcely twisted. Sepals very unequal, subparallel with spreading, often pubescent. Dorsal sepal erect, concave; lateral sepals long-decurrent on ovarian wall without any observable line of adnation, the free apices ± falcate; petals joined with dorsal sepal, decurrent at base. Lip clawed, distinctly sagittate at base, margins near middle joined with column sides, often with a reflexed terminal lobe. Column rather short with long foot, which is embedded the full length internally in ovarian tissue and with it the ± connate lateral sepals form a prominent internal nectary or cuniculus, not discernible externally; anther ovate, cordate, obtuse; pollinia 4, soft, mealy, clavate with a large, thick, abaxial viscidium; rostellum laminar, soft ligulate, ± truncate; stigmas free to approximate, ± touching each other in middle. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, southern Brazil, Bolivia, Paraguay, northern Argentina; ca. 35 species, 5 in Venezuela, all in the flora area. Key to the Species of Sarcoglottis 1. 1. 2(l). 2.

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

4.

Plants without leaves during anthesis (hysteranthous), usually growing in open places ......................................................................................... 2 Plants leaf-bearing during anthesis (synanthous), usually growing in the understory of forests .............................................................................. 3 Sepals glabrous outside; dorsal sepal linear-oblong, obtuse; lip apically truncate and 3-lobed; disk glabrescent ...................................... S. aphylla Sepals pubescent or glabrescent outside; dorsal sepal basally ovate-elliptic, apically attenuate into a oblong appendix; lip apically contracted into a rhombic-ovate apical lobe; disk densely pubescent ......... S. simplex Plants < 15 cm tall; leaves < 10 cm long; flowers white; lip 1.5 cm long ................................................................................................... S. stergiosii Flowering plants at least 20 cm tall (usually higher); leaves 15 cm long; flowers greenish, deep green or cream; lip at least 2.5 cm long .......... 4 Leaves 4–7 times longer than wide, gradually attenuated toward base; floral bracts > 1/2 the length of the pedicellate ovary; dorsal sepal 2 cm long; apical lobe of lip as longer as wide or longer than wide .... S. acaulis Leaves 2–3 times longer than wide, abruptly attenuate toward base; floral bract < than 1/2 the length of the pedicellate ovary; dorsal sepal 1.8 cm long; apical lobe of lip wider than long ......................... S. metallica

Sarcoglottis 553

Sarcoglottis acaulis (Sm.) Schltr., Repert. Spec. Nov. Regni Veg. Beih. 6: 53. 1919. —Neottia acaulis Sm., Exot. Bot. 2: 91, t. 105. 1806. Sarcoglottis grandiflora auct. non Lindl. 1842: sensu Dunst. & Garay, Orchids Venez. Ill. Field Guide 904. 1979. Terrestrial to humicolous, 30–120 cm tall, erect, shade-loving; flowers large for the alliance, green or creamy. Rain forests, 100–500 m; Bolívar (lower Río Caroní, Río Caura). Aragua, Distrito Federal, Falcón, Miranda, Sucre. Ecuador. Sarcoglottis aphylla (Ridl. ex Thurn) Schltr., Beih. Bot. Centralbl. 37(2): 474. 1919. —Pelexia aphylla Ridl. ex Thurn, Timehri 5: 205. 1886. —Pelexia aphylla Ridl. ex Oliv., Trans. Linn. Soc. London, Bot. 2: 284, t. 48b. 1887, nom. superf., non Pelexia aphylla (Vell.) Schltr. 1920. —Spiranthes aphylla (Ridl. ex Thurn) Cogn. in Mart., Fl. Bras. 3(4): 215. 1895. —Veyretia aphylla (Ridl. ex Thurn) Szlach., Fragm. Florist. Geobot. Supp. 3: 116. 1995. Terrestrial, erect, sun-loving; flowers medium-large for the genus, white. Rain or cloud forests; Bolívar (Roraima-tepui). Brazil. There is only one collection of Sarcoglottis aphylla in the flora area. It is from Roraimatepui, but the exact locality was not given. Sarcoglottis metallica (Rolfe) Schltr., Beih. Bot. Centralbl. 37(2): 417. 1920. —Spiranthes metallica Rolfe, Bull. Misc. Inform. Kew 1896: 46. 1896. Sarcoglottis picta auct. non Klotzsch 1842: sensu Garay & Dunst. in Dunst. & Garay, Orchids Venez. Ill. Field Guide 903. 1979. Sarcoglottis acaulis auct. non (Sm.) Schltr. 1919: sensu Foldats in Lasser, Fl. Venez. 15(1): 302. 1969. Terrestrial or humicolous, to 20 cm tall, rosulate, erect, shade-loving; upper surface of leaves usually velvety or coppery brown, the lower surface silvery or black; flowers medium-sized for the genus, greenish. Evergreen lowland to lower montane forests, 100– 1000 m; Bolívar (La Escalera to Cerro Venamo region), Amazonas (Cerro Aratitiyope, Cerro Yutajé, Sierra de la Neblina). Guyana.

Sarcoglottis simplex (Griseb.) Schltr., Beih. Bot. Centralbl. 37(2): 421. 1920. —Spiranthes simplex Griseb., Fl. Brit. W. I. 641. 1864. —Veyretia simplex (Griseb.) Szlach., Fragm. Florist. Geobot. suppl. 3: 116. 1995. Terrestrial, medium-sized to small, erect, sun-loving; flowers moderately small for the genus, greenish or greenish white. Open Trachypogon savannas, 50–200 m; Bolívar (Río Caroní basin), Amazonas (Río Cataniapo basin). Brazil, Paraguay. Sarcoglottis stergiosii Carnevali & I. Ramírez, Novon 3: 124. 1993. Terrestrial to humicolous, small to minute, shade-loving; upper surface of leaves green or purple-brown, the lower surface silvery or green; flowers white. Rain forests, 50–1200 m; southern Amazonas (Río Negro basin). Endemic. ◆Fig. 490.

Fig. 490. Sarcoglottis stergiosii

554

O RCHIDACEAE

128. SCAPHOSEPALUM Pfitzer in Engl. & Prantl., Nat. Pflanzenfam. 2(6): 139. 1888. [Subtribe Pleurothallidinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, lithophytes, or subterrestrial herbs, erect to subpendulous, mainly cespitose or else creeping. Roots few to many, slender to coarse or fleshy; rhizome abbreviate to elongate between ramicauls, horizontal or ascending; ramicauls ascending to erect, slender to stout, terete or ventrally canaliculate, shorter than the leaf, 1-foliate apically, partially or completely enclosed by 2 or 3 imbricating sheaths. Leaves conduplicate, erect, thin to thick-coriaceous, smooth, green or dark green, rarely purplish, elliptic to narrow-elliptic or obovate to narrow-obovate, the apex acute to obtuse, shallowly notched with a mucro in the sinus, the base narrowly cuneate into a channeled pseudopetiole. Inflorescences emerging from an annulus, originating laterally close to the base of the ramicaul or subapically from near the apex of the ramicaul, a successively flowered, loose to dense, few- to many-flowered raceme, sometimes several flowers produced simultaneously, longer to shorter than the leaves, smooth to coarsely verrucose, sometimes branching and producing new shoots; peduncle erect to ascending or subpendulous, slender to stout, with 1–several bracts; rachis straight to strongly zigzag. Flowers nonresupinate, secund (with the flowers to one side) or distichous on the rachis, campanulate or suburceolate to widely open; floral bracts thin and tubular to broad, fleshy and conduplicate, longacuminate to obtuse, longer to shorter than the pedicel; pedicel thin to stout, sometimes verrucose; ovary smooth, verrucose to papillose. Sepals ± fleshy, variously colored, smooth, verrucose, spiculate, or pubescent, basally concave, apically contracted into vestigial to elongate tails; dorsal sepal basally connate to the lateral sepals or free, 3-costate; lateral sepals connate into a concave synsepal, with a pair of ± distinct, fleshy, flattened, elliptic to lunate or triangular calli (the cushion), occupying the distal and internal portion of the sepal blade within; tails slender to thickened and verrucose, erect, divergent to retrorse; petals much smaller than the sepals, about as long as the column, ± hyaline, ovate, often ventricose, sometimes apiculate. Lip hinged to apex of column foot, about the length of the column, usually reflexed near the middle, oblong, ovate, or often pandurate, sometimes with small lateral lobes near the middle; disk with a small pair of usually crested lamellae near the middle; hypochile ± quadrate and concave, the base truncate and often minutely bilobed; epichile acute, obtuse to rounded, smooth to denticulate and verrucose. Column semiterete, broadly winged above the middle, basally produced into a thick column foot; anther ventral, incumbent, operculate, 2-locular; clinandrium entire or toothed; pollinia 2, pear-shaped, with elastic caudicles; rostellum retrorse; stigmatic surface ventral. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia; 30 species, 2 in Venezuela, 1 of these in the flora area. Scaphosepalum breve (Rchb. f.) Rolfe, J. Bot. 28: 136. 1890. —Masdevallia brevis Rchb. f., Gard. Chron. n.s. 2: 588. 1883. Scaphosepalum verrucosum auct. non (Rchb. f.) Pfitzer. 1888: sensu G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 905. 2000.

Epiphyte or rarely a lithophyte; leaves to 15 cm long; inflorescence verrucose; rachis zigzag; flowering successively, perianth segments yellow-green, variously tinged or spotted with purple, tails usually dark purple; dorsal sepal 10–18 mm long. Locally common in cloud forests, rare on exposed sand-

Scaphyglottis 555

stone outcrops, 1300–2200 m; Bolívar (Auyán-tepui, Cerro Jaua, La Escalera to Cerro Venamo region, Macizo del Chimantá, Uaipán-tepui). Aragua, Carabobo, Distrito Federal, Miranda, Sucre, Zulia; Colombia, Guyana, Ecuador, Bolivia. ◆Fig. 491.

Fig. 491. Scaphosepalum breve

129. SCAPHYGLOTTIS Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 58. 1835. [Subtribe Laeliinae]. Cladobium Lindl., Introd. Nat. Syst. Bot. ed. 2, 446. 1836. Hexadesmia Brongn., Ann. Sci. Nat. Bot. sér. 2, 17: 44. 1842. Tetragamestus Rchb. f., Bonplandia (Hanover) 2: 27. 1854. Leaoa Schltr. & Porto, Arch. Jard. Bot. Rio de Janeiro, 3: 292. 1922. by Germán Carnevali and Ivón M. Ramírez-Morillo Mostly epiphytes, sometimes lithophytes or humicolous herbs, inconspicuous to large and conspicuous, usually cespitose, erect to pendulous. Roots thin or relatively thick. Rhizome abbreviate; stems heteroblastic (one internode), basally enveloped by few to several tubular, scarious sheaths that very rarely bear foliar blades, narrow-cylindric to narrow-pear-shaped, frequently ± dilated into narrowly fusiform pseudobulbs, sometimes variously stipitate, usually proliferous at apex with 1–several superposed, younger segments, each segment apically 1–3-leaved. Leaves conduplicate, articulate, coriaceous, usually linear, narrowly oblong-elliptic or elliptic, apex often obliquely emarginate and mucronate, sessile or rarely subpetiolate at base, usually congested at the apex of the stem, rarely subdistichous. Inflorescences terminal, usually 1-flowered, rarely racemose, abbreviate, generally fasciculate, emerging from a group of sheaths; floral bracts usually inconspicuous; pedicellate ovary terete, often relatively elongate, sometimes curved. Flowers mostly inconspicuous, less often ± showy, resupinate or not, usually short-lived. Perianth segments membranous to ± fleshy, white, greenish, yellowish or pink, sometimes spotted or streaked with red or purple, rarely lip or whole flower violet or bluish, subcampanulate to widely spreading; sepals subsimilar; dorsal sepal free, lateral sepals oblique, ± adnate to column foot with which basally they form a variously developed chin or mentum; petals subsimilar to sepals, usually shorter and narrower, rarely broader. Lip articulate to rigidly attached to the apex of the column foot, erect, ± spreading or recurved, variously clawed to subsessile, simple or variously lobed, frequently emarginate to 2-lobed apically; disk mostly ± ecallose. Column short to relatively elongate, erect or at angle with respect to pedicellate ovary, exauriculate to variously winged, basally produced into an inconspicuous to well-

556

O RCHIDACEAE

developed foot; anther terminal, operculate, incumbent; pollinia 4 or six, waxy, laterally flattened, with yellow caudicles; clinandrium entire or variously 3-lobed; rostellum transverse; stigma ventral. Capsules ovoid to ellipsoid. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 55 species, 18 species in Venezuela, 14 of these in the flora area. Some authors include Hexisea Lindl. and Reichenbachanthus Barb. Rodr. in Scaphyglottis, while some others segregate Cladobium Lindl. and Tetragamestus Rchb. f. Some species of Scaphyglottis have been reported to be pollinated by trigonid bees; many other species are self-pollinating or even cleistogamous. Key to the Species of Scaphyglottis 1. 1. 2(1).

2. 3(2).

3.

4(3).

4.

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

6.

Pedicellate ovary longer to little shorter than dorsal sepal, exposed above sheaths ................................................................................................... 2 Pedicellate ovary conspicuously shorter than sepals, or if longer, almost totally hidden by sheaths ...................................................................... 8 Pseudobulbs not abruptly long-stipitate, at most slightly attenuated basally (if stipitate, then pollinia 4 and column with finger-like projections below the stigmatic surface) ......................................................... 3 Pseudobulbs or stems abruptly long-stipitate (i.e., stalked; the stalk is at least 1/3 the total length of the stem); pollinia 6 ................................... 5 Lip simple (apically dilated but not lobed), flowers large for the genus; dorsal sepal 9–14 mm long, sepals and petals white with dense purple zones; lip pale purple with deep purple streaks or wholly purple; column erect with respect to pedicellate ovary; plants usually from intermediate or high-elevation cloud forests ............................... S. grandiflora Lip 3-lobed; flowers small with dorsal sepal 5–8.3 mm long, not resupinate, sepals and petals greenish, white or pink; lip greenish, violet or purple; column forming a sharp angle with respect to pedicellate ovary; plants usually from lowland rain forests .............................................. 4 Lip greenish or greenish white; column foot short (< 1/4 of the length of the column); central lobe of lip acute; older stems not stipitate, always green ........................................................................................ S. boliviensis Lip pink or violet; column foot relatively long (1/4–1/3 of column length); central lobe of lip truncate to emarginate; older stems commonly stipitate, usually purple-tinged ................................................... S. stellata Leaves 2–4(–5) times longer than wide, 15–40 mm wide; pseudobulbs usually apically 2-leaved ............................................................................. 6 Leaves 6–20 times longer than wide, 3–8 mm wide; pseudobulbs apically 1-leaved .................................................................................................. 7 Leaves green; sepals and petals 0.8–1.2 cm long, greenish or greenish purple with purple or maroon stripes; lip broader toward middle or at about the apical 1/3, acute or obtuse .............................................. S. bifida Leaves with purplish or brownish blotches; sepals and petals ca. 2 cm long, whitish or pink; lip broader toward apex, truncate or rounded ............................................................................................. S. dunstervillei

Scaphyglottis 557

7(5).

Plants shortly creeping to subcespitose; inflorescence stout, short, 1- or 2(3)-flowered; pedicels at most 2 times longer than the dorsal sepal; petals ca. 1 cm long, 3–4 times longer than wide; lip apex truncate or rounded, shortly apiculate ...................................................... S. fusiformis 7. Plants cespitose; inflorescence filiform, successively 3–5-flowered; pedicels at least 4 times as long as the dorsal sepal; petals 2.5–4 mm long, about as broad as long; lip apex deeply emarginate .......... S. sessilis 8(1). Flowers relatively large with dorsal sepal about 8 mm long; lip about 9.5 mm long; pedicel strongly arched downward ................... S. leucantha 8. Flowers small with dorsal sepal to 6.5 mm long; lip to 6.9 mm long (usually shorter); pedicel straight ................................................................ 9 9(8). Leaves (9–)12–22 mm wide; petals tenuously 3–5-veined ...................... 10 9. Leaves to 8 mm wide (usually narrower); petals 1-veined or rarely 3-veined ................................................................................................ 11 10(9). Lip apically truncate to rounded; calli antrorse .............................. S. felskyi 10. Lip apically acute to subrounded, apiculate; calli erect or ± retrorse .................................................................................................... S. modesta 11(9). Lip broader above the apical 1/3; disk callose; flowers solitary ............... 12 11. Lip broader in its lower 1/2; disk ecallose; flowers 1–8 in a fascicle ........ 13 12(11). Labellar claw < 1/3 total length of lip; petals oblique falcate; disk of lip bicallose; flowers purple; plants growing above 1300 m elevation .................................................................................... S. michelangeliorum 12. Labellar claw about 1/2 total length of lip; petals not or very slightly oblique; disk of lip unicallose; flowers white with a maroon column-apex; plants from 50–800 m elevation ................................................ S. prolifera 13(11). Flowers pink, rose or purple, very rarely white; lip spreading at apical 1/2, shortly clawed or sessile; pseudobulbs frequently stipitate, often purple-tinged; lateral sepals rounded to obtuse ................ S. graminifolia 13. Flowers white or rarely yellow; lip parallel to column to its apex, long clawed, acute to obtuse; pseudobulbs never stipitate, totally green; lateral sepals acute ............................................................................. S. sickii Scaphyglottis bifida (Rchb. f.) C. Schweinf., Bot. Mus. Leafl. 10: 27. 1941. —Hexadesmia bifida Rchb. f. in Saunders, Refug. Bot. 2: sub t. 113. 1878. Epiphyte; pseudobulbs 5–30 cm long, long-stalked; flowers greenish with purple on veins, in short inflorescences. Rain forests, 400–1400 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region), Amazonas (Río Ararí). Venezuelan Coastal Cordillera, Táchira; Mexico, Central America. Scaphyglottis boliviensis (Rolfe) B.R. Adams, Phytologia 64(10): 257. 1988. —Hexadesmia boliviensis Rolfe, Mem. Torrey Bot. Club, 6(1): 122. 1896. Scaphyglottis huebneri Schltr., Beih. Bot. Centralbl. 42(2): 95. 1925.

Epiphyte or rarely a lithophyte, arching or subpendulous, shade-loving; stems cylindric, 5–20 cm long; flowers subcampanulate, inconspicuous, with dorsal sepal 4–6.5 mm long; sepals and petals grayish green, column greenish with red-purple tinges. Locally common in rain forests, 50–600 m; Bolívar (Auyán-tepui), Amazonas (Coromoto, Río Casiquiare, Río Cataniapo, Río Yatúa). Zulia; Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil, Bolivia. Scaphyglottis dunstervillei (Garay) Foldats, Bol. Soc. Venez. Ci. Nat. 28: 255. 1969. —Hexadesmia dunstervillei Garay, Bot. Mus. Leafl. 18: 203. 1958. Epiphyte, medium-sized to large; pseudobulbs to 30 cm long, long-stalked; flowers large for the genus, rose-white with rose-

558

O RCHIDACEAE

brown veins. Cloud forests, 600–1300 m; Bolívar (Altiplanicie de Nuria, La Escalera to Cerro Venamo region). Miranda; St. Lucia. ◆Fig. 492. Scaphyglottis felskyi (Rchb. f.) Schltr., Repert. Spec. Nov. Regni Veg. Beih. 6: 66. 1919. —Ponera felskyi Rchb. f., Linnaea 41: 85. 1877. Epiphyte; flowers campanulate with sepals ca. 6 mm long; sepals and petals greenish with brownish stripes, lip whitish with brown spots. Rain forests, ca. 2700 m; Bolívar (Roraima-tepui). Guyana, Suriname. Scaphyglottis felskyi is an imperfectly known species. It has been reported in the literature for Roraima-tepui without locality or collector data. It is likely that this report is S. modesta, which is seemingly closest related to S. felskyi, and is relatively common in southeastern Bolívar. Scaphyglottis fusiformis (Griseb.) R.E. Schult., Bot. Mus. Leafl. 17: 205. 1956. —Hexadesmia fusiformis Griseb., Fl. Brit. W. I. 623. 1864. Epiphyte, rather small to medium-sized; rhizome creeping; pseudobulbs superposed; flowers small, greenish or whitish. Open forests or shrublands, 300–1400 m; Delta Amacuro, widespread in Bolívar and Amazonas. Venezuelan Coastal Cordillera, Andean foothills; Costa Rica, Colombia, Trinidad-Tobago, Guyana, Brazil. Scaphyglottis graminifolia (Ruiz & Pav.) Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 59. 1836. —Fernandezia graminifolia Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 1: 241. 1798. Scaphyglottis violacea (Lindl.) Lindl., Edwards’s Bot. Reg. 22: t. 1901. 1836. —Cladobium violaceum Lindl., Intr. Nat. Syst. Bot. ed. 2, 446. 1836. Epiphyte or lithophyte, erect to subpendulous; pseudobulbs frequently purpletinged, 5–15 cm tall; flowers inconspicuous, not widely spreading; dorsal sepal 3–4 mm long. Rain or cloud forests, ca. 50–1200 (–2000) m; Delta Amacuro (Río Amacuro, Serranía de Imataca), Bolívar (Altiplanicie de Nuria, widespread in the basin of the Río Caroní), Amazonas (Río Cataniapo). Aragua, Distrito Federal, Miranda, Yaracuy; Colombia, Guyana, Ecuador, Peru, Bolivia.

Scaphyglottis grandiflora Ames & C. Schweinf., Bull. Torrey Bot. Club 58: 349. 1931. Epiphyte, less often muscicolous on rocks, erect to subpendulous, mostly sun-loving; stems 4–20 cm long; flowers large for the genus with dorsal sepal 9–13.5 mm long, sepals and petals white with purple zones, pale purple with deep purple streaks or wholly deep purple, lip purple. Cloud forests or rarely rain forests, (700–)1400–2000 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Jaua-Sarisariñama massif, La Escalera to Cerro Venamo region, Macizo del Chimantá, Ptari-tepui, Roraima-tepui, Sierra Pakaraima), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Marahuaka, Cerro Parú, Sierra de la Neblina, Sierra Parima). Sucre; Amazonian Brazil, Guyana, French Guiana. Scaphyglottis leucantha Rchb. f., Linnaea 22: 856. 1850. —Ponera leucantha (Rchb. f.) Rchb. f., Bonplandia (Hanover) 2: 22. 1854. Scaphyglottis wercklei var. major C. Schweinf. Bot. Mus. Leafl. 4(7): 117. 1937. Epiphyte, erect to arching; secondary stems narrowly cylindric, not stipitate; flowers ± showy for the genus but held away from view due to downward torsion of pedicellate ovary, sepals and petals white at base, grading to pale green at apex; disk of lip with strong marks of light purple and some tinges of the same color or of violet or bluish, column green at base, pink at apex. Cloud forests, 1200–1300 m; Bolívar (La Escalera to Cerro Venamo region). Mérida, Trujillo; Central America, Andean Colombia, Ecuador, and Peru. Scaphyglottis michelangeliorum Carnevali & Steyerm., Phytologia 55: 289. 1984. Epiphyte, small for the genus; pseudobulbs 2–4.5 cm long, shortly stipitate; flowers inconspicuous, subcampanulate, with dorsal sepal ca. 38 mm long, sepals and petals entirely purple. Cloud forests, ca. 1500 m; Amazonas (Cerro Marahuaka). Endemic. Scaphyglottis modesta (Rchb. f.) Schltr., Repert. Spec. Nov. Regni Veg. 23: 46. 1926. —Tetragamestus modestus Rchb. f., Bonplandia (Hanover) 2: 27. 1854.

Scaphyglottis 559

Fig. 492. Scaphyglottis dunstervillei

Epiphyte or rarely a lithophyte or muscicolous herb, large for the genus, shade-loving; roots thick; stems 5–30 cm long, not stipitate, erect to arching; flowers inconspicuous, with dorsal sepal 5–6 mm long, often cleistogamous; perianth segments yellowish with purple streaks or maroon tinges. Locally common in rain forests or cloud forests, 600–1000 m; Bolívar (Altiplanicie de Nuria, La Escalera to Cerro Venamo region, basin of the upper Río Caroní), Amazonas (Cerro Aratitiyope, Cerro Huachamacari). Falcón, Miranda, Sucre; West Indies, Panama, Guyana, Brazil, Colombia, Ecuador. Scaphyglottis prolifera Cogn. in Mart., Fl. Bras. 3(5): 75. 1898. Scaphyglottis cuneata Schltr., Beih. Bot. Centralbl. 36(2): 398. 1918. Epiphyte; secondary stems not stipitate, 4–11(–20) cm long; flowers resupinate, solitary, inconspicuous with dorsal sepal 3–6 mm long; sepals and petals with a faint pink midvein; lip white; column cream. Rain forests, 100–800 m; Bolívar (near Santa Elena de Uairén). Zulia; Central America, West Indies, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia.

Fig. 493. Scaphyglottis sickii

Scaphyglottis sessilis (Rchb. f.) Foldats, Acta Bot. Venez. 3: 398. 1968. —Hexadesmia sessilis Rchb. f., Linnaea 42: 84. 1877. Scaphyglottis reedii (Rchb. f.) Ames, Amer. Orchid Soc. Bull. 10: 49. 1941. —Leaoa reedii (Rchb. f.) Garay, Arch. Jard. Bot. Rio de Janeiro 13: 45. 1955. Epiphyte, small to large; pseudobulbs 1foliate; flowers small, greenish with brown or purple tinges, especially on lip. Rain forests, 200–800 m; Bolívar, Amazonas. Venezuelan Coastal Cordillera, Andean foothills; Mexico, Central America, Colombia, Guyana, Peru, Brazil. Scaphyglottis sickii Pabst, Orquídea (Mexico) 18: 7. 1956. Scaphyglottis propinqua auct. non C. Schweinf. 1955; Foldats in Lasser, Fl. Venez. 15(3): 104. 1970. Epiphyte, rarely a lithophyte; secondary stems not stipitate, 3–10(–18) cm long; flowers solitary, inconspicuous with dorsal sepal 2.7–4 mm long; perianth segments not opening widely. Rain forests, 50–500(–800) m; widespread in Delta Amacuro, Bolívar, and Amazonas. Apure, Miranda; Panama,

560

O RCHIDACEAE

Guyana, French Guiana, Amazonian Ecuador, Peru, and Brazil. ◆Fig. 493. Scaphyglottis stellata Lodd. ex Lindl., Edwards’s Bot. Reg. 25: misc. 44. 1839. Scaphyglottis amethystina (Rchb. f.) Schltr., Beih. Bot. Centralbl. 36(2): 456. 1918. —Ponera amethystina Rchb. f. in Saunders, Refug. Bot. 2: pl. 93. 1869. Epiphyte, rarely a lithophyte, erect to arching; pseudobulbs often purple or redtinged, 4–12 cm long; inflorescences fascicu-

late; flowers not resupinate with dorsal sepal 5–8.5 mm long; sepals and petals pink to purple or rarely white; lip pink to violet or purple. Rain forests, frequently in rather open places, 50–400(–1200) m; Bolívar (basin of the upper Río Caroní, basin of the lower Río Caura, basin of the Río Paragua, Río Parguaza), Amazonas (Cerro Aratitiyope, Cerro Parú, basin of Río Cataniapo). Guatemala, Nicaragua, Costa Rica, Panama, French Guiana, Amazonian Colombia, Ecuador, Peru, and Brazil.

130. SCELOCHILUS Klotzsch,, Allg. Gartenzeitung 9: 261. 1841. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, usually growing on twigs, cespitose, erect to pendulous, usually sun-loving. Rhizome abbreviate; pseudobulbs inconspicuous to almost absent, usually subterete, enveloped by several imbricate sheaths from which the innermost 1 or 2 may have leaf blades, apically 1-leaved. Leaves conduplicate, articulate, usually oblong or oblong-elliptic, acute, basally shortly pseudopetiolate. Inflorescences lateral, from pseudobulb base, erect, arching to pendulous, shorter to much longer than leaves, usually a lax, few-flowered raceme, sometimes 1–few lateral branches arise from older internodes of the peduncle; peduncle remotely few-sheathed; floral bracts inconspicuous, tubular to concave. Flowers long-pedicellate, nonresupinate, shortlived, usually yellow or yellow-green with red, maroon, brown, or purple streaks or blotches. Perianth segments membranous to subfleshy, parallel to column or the apical 1/2 spreading; dorsal sepal free, concave; lateral sepals partially to totally connate into a ± concave or flat synsepal; synsepal subequal to dorsal sepal but often somewhat broader, margins ± erect, basally produced into a prominent, retrorse, cylindric spur. Petals parallel to column, subequal to dorsal sepal or broader. Lip erect, basally with a pair of short, narrow spurs that fit into the sepal spur, lamina simple, pandurate to ± 3-lobed, apically usually rounded or emarginate; disk variously callose, usually with 2 parallel, longitudinal keels. Column erect, relatively thin, subcylindric to subclavate, exauriculate or with very small wings, almost footless. Anther terminal, operculate, incumbent, with reduced partitions; clinandrium shallow; pollinia 2, waxy, subglobose, sulcate, almost without tegula and with a small viscidium; rostellum transverse; stigma ventral. Southern Mexico, Central America, Colombia, Venezuela, French Guiana, Ecuador, Peru; ca. 20 species, 3 or 4 in Venezuela, 2 of these in the flora area. Key to the Species of Scelochilus 1.

1.

Leaves 3–5 times longer than wide; inflorescences several-flowered; petals as wide as or slightly wider than dorsal sepal; lip 4–5 times longer than wide, oblong; plants from elevations above 1500 m ............ S. ottonis Leaves 8–10 times longer than wide; inflorescences 1-flowered; petals conspicuously broader than dorsal sepal; lip 2–3 times longer than wide, pandurate; plants from elevations below 500 m ..... S. paraguaensis

Schomburgkia 561

Scelochilus ottonis Klotzsch, Allg. Gartenzeitung 9: 261. 1841. Epiphyte; leaves 8–14 cm long; inflorescences ± equal to the leaves; flowers with perianth segments golden yellow, striped with maroon, lip golden yellow with maroon blotches or zones, 12–17 mm long. Cloud forests, ca. 1800 m; Amazonas (Sierra de la Neblina). Aragua, Distrito Federal, Carabobo, Mérida, Trujillo, Yaracuy.

Fig. 494. Scelochilus paraguaensis

Scelochilus paraguaensis Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 5: 268. 1972. Epiphyte; leaves 5–11 cm long, inflorescences shorter than leaves; flowers with pink perianth segments; lip white grading to pale pink or cream toward apex, 11–14 mm long; column white. Rain forests, 300–400 m; Bolívar (El Crucero at upper Río Paragua). Endemic. ◆Fig. 494.

131. SCHOMBURGKIA Lindl., Sert. Orchid. t. 10. 1838. —Laelia sect. Schomburgkia (Lindl.) L.O. Williams, Darwiniana 5: 75. 1941. [Subtribe Laeliinae]. Amalia Rchb. f. in Heynh., Nom. Bot. Hort. 52. 1841. by Germán Carnevali and Ivón M. Ramírez-Morillo Usually epiphytes, often lithophytes or muscicolous on rocks, mostly sun-loving, cespitose, large and showy. Rhizome shortly creeping, thick; pseudobulbs heteroblastic (one internode), erect, fusiform, ± stipitate, ± 4-angular, strongly ribbed, apically 1–3(4)-leaved, basally enveloped by several imbricate, scarious, not leafbearing sheaths. Leaves conduplicate, articulate, thickly coriaceous, sessile, oblong to elliptic, acute or emarginate apically. Inflorescences terminal, solitary, usually much longer than leaves, erect or rarely ascendent, pseudoumbellate, long-pedunculate; peduncle terete, many-articulate, enveloped by several sheaths, often purpletinged; sheaths ± imbricate, tubular, scarious, dorsally keeled, acuminate; rachis several times shorter than the peduncle, erect, often purple-tinged; floral bracts conspicuous, shorter than pedicellate ovary, linear-subulate to linear-oblong, usually wilting at anthesis and becoming pendulous; pedicellate ovary elongate, terete, perpendicular to rachis, white, pink, or purple. Flowers showy, sometimes cleistogamous, resupinate, spiral on rachis, opening simultaneously; perianth segments free, similar, subfleshy, widely spreading in noncleistogamous species, strongly undulate in most species, mostly dark purple or dark purple-brown but in some species pink, white, or yellow to brown with a marginal yellow band, usually shiny, narrowly oblong to narrowly oblong-elliptic, acute to rounded. Lip connate in the basal 1/5 with the basal margins of the column, above this articulate, shorter than sepals and petals, purplish in the species with dark purple perianth, whitish to yellowish in the species with white, or brown to yellow perianth, simple to sharply 3-lobed; lateral lobes erect and enfolding the column, rarely almost flat; central lobe larger than lat-

562

O RCHIDACEAE

erals, parallel to column or often spreading, clawed or sessile on the basal portion of lip; disk usually arched, with a callus composed of several raised, parallel keels that are often yellow or orange. Column erect, arched, concave, winged along all its margins, often white or purplish, with a very short foot at base; anther terminal or subventral, incumbent, operculate, imperfectly 4 or 8-locular; pollinia 8, subequal or 2 pairs larger than the others, waxy, yellow, laterally compressed, with well-developed caudicles; clinandrium lobed or crenate; rostellum transverse; stigma ventral, entire. Capsules ellipsoid. Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, northern Brazil, Bolivia; ca. 15 species, ca. 6 in Venezuela, 3 of these in the flora area. Key to the Species of Schomburgkia 1.

1.

2(1).

2.

Sepals and petals yellowish to chestnut brown, or pinkish brown, always with yellow margins on the petals; lip with yellow lateral lobes, white disk and pinkish central lobe, central lobe strongly reflexed; sepals 22– 28 mm long; rachis of the inflorescence green or white; flowers often cleistogamous . ....................................................................... S. marginata Sepals and petals dark purple-maroon, purple-brown, or less often pale purple, concolorous; lip entirely purple, sometimes with some white on the disk, central lobe ± erect; sepals (17–)27–36 mm long; rachis of the inflorescence pink or purple; flowers always chasmogamous .............. 2 Central lobe of lip sessile on the lateral lobes or very shortly clawed, about as long as wide; midlobe with a callosity composed of 3 conspicuously raised keels with 2 inconspicuous keels flanking them ............... S. heidii Central lobe of lip broadly but conspicuously clawed, 1.5–2 times longer than wide; midlobe with a callosity composed of 5 conspicuously raised keels with 2 inconspicuous keels flanking them ..................... S. undulata

Schomburgkia heidii Carnevali, Ernstia 10: 2. 1982. Schomburgkia rosea auct. non Linden ex Lindl. 1845: sensu Foldats in Lasser, Fl. Venez. 15(3): 462. 1970. Lithophyte, less often an epiphyte; sepals and petals wine purple, less often pale purple, lip purple. Locally common on igneous outcrops under the shelter of trees or shrubs in semideciduous forests, 50–200 m; Bolívar (Piedra Marimare), Amazonas (near Puerto Ayacucho). Colombia (Vichada). ◆Fig. 495. There is some evidence of introgression between Schomburgkia heidii and S. undulata near Puerto Ayacucho.

Schomburgkia marginata Lindl., Sert. Orch. sub t. 10, t. 13. 1838. —Laelia marginata (Lindl.) L.O. Williams, Darwiniana 5: 76. 1941. Schomburgkia crispa auct. non Lindl. 1838: sensu Foldats in Lasser, Fl. Venez. 15(3): 456. 1970. Schomburgkia weberbaueriana auct. non Kraenzl. 1906: sensu Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 1: 390. 1959. Epiphyte. Locally common in semideciduous or rain forests, 50–600(–800) m; Bolívar (Altiplanicie de Nuria, near Tumeremo, near Upata, near El Pao, near Santa Elena de Uairén, Voituco). Monagas; Guyana, Suriname, northern Brazil.

Schomburgkia 563

Schomburgkia undulata Lindl., Edwards’s Bot. Reg. 30: misc. 13, sub t. 23. 1844. —Laelia undulata (Lindl.) L.O. Williams, Darwiniana 5: 76. 1941. Epiphyte or lithophyte; sepals and petals dark purple-maroon, lip purple or pinkpurple. Semideciduous forests, often on igneous outcrops, 200–300 m; Bolívar (Cerro Bolívar, near Ciudad Piar). Widespread in northern Venezuela in areas with a marked dry season, but absent from the western coast of Lago de Maracaibo; Colombia, Trinidad-Tobago.

Fig. 495. Schomburgkia heidii

564

O RCHIDACEAE

132. SCUTICARIA Lindl., Edwards’s Bot. Reg. 29: misc. 14. 1843. [Subtribe Zygopetalinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Mostly epiphytes, rarely muscicolous on rocks or lithophytes, erect or pendulous, cespitose or creeping, sun- or shade-loving. Roots fibrous, elongate, sometimes reddish or brownish. Rhizome abbreviate or creeping, relatively thick; pseudobulbs inconspicuous, heteroblastic (one internode), cylindric, fusiform or narrow-pearshaped, apically 1-leaved, totally enveloped by scarious, tubular, imbricate sheaths. Leaves terete, articulate, very elongate, linear, pendulous or erect in 1 species, sometimes purplish. Inflorescences lateral from the base of the pseudobulbs, 1-flowered or 2–4-flowered racemes, always much shorter than the leaves, erect or horizontal to nodding; peduncle abbreviate or rarely longer than leaves, enveloped by scarious, tubular sheaths, or remotely sheathed; rachis very abbreviate; floral bracts inconspicuous; pedicellate ovary terete, longer than the sepals. Flowers resupinate, large and showy, fleshy, long-lasting, subcampanulate or with widely spreading perianth segments; perianth segments yellow or greenish yellow with reddish or brownish spots or blotches, and a cream or yellowish, red streaked lip, or completely maroon or reddish and then lip white with reddish purple lines; sepals free, subequal or the lateral broader; lateral sepals inserted on the margins of the column foot and forming a conspicuous mentum; petals narrower than the sepals. Lip articulate with column foot, erect, much broader than the sepals, 3-lobed; lateral lobes erect and loosely embracing the column; central lobe smaller than laterals, flat, rounded or emarginate at apex; disk with a conspicuous, fleshy, several keeled callus. Column elongate, relatively thick, erect, terete, wingless, basally produced into a ± developed foot; anther terminal, operculate, incumbent, very convex, 1-locular or imperfectly 2-locular; pollinia 4, waxy, compressed, in 2 dissimilar pairs in which the anterior pair is larger than the posterior, tegula semilunate, thin, viscidium semilunate, small; clinandrium truncate, shallow; rostellum transverse; stigma ventral, entire. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 5 species, 1 in Venezuela. Scuticaria steelei (Hook.) Lindl., Edwards’s Bot. Reg. 29: misc. 14. 1843. —Maxillaria steelei Hook., Bot. Mag. 64: t. 3573. 1837. Epiphyte, pendulous; leaves 5–10 mm thick, 50–160(–200) cm long; flowers very fragrant, yellow with reddish or maroon spots; dorsal sepal 35–50 mm long; lip whitish externally, internally whitish or cream with transverse reddish or maroon stripes on

lateral lobes, callus bright yellow or orange. Locally common in rain forests, frequently on river edges, sometimes in shrublands, 100–900(–1200) m; Bolívar (Cerro Guaiquinima, near El Cajón, near El Paují, Río Akaruai, Río Carrao, Río Uaiparú), Amazonas (near Puerto Ayacucho, Río Atabapo, Río Guainía, Río Pavone, Río Sipapo). Amazonian Colombia and Brazil; Guyana, Suriname, French Guiana. ◆Fig. 496.

133. SELENIPEDIUM Rchb. f., Xenia Orchid. 1: 3, t. 2. 1854; emend. Pfitzer in Engl., Pflanzenr. IV. 50(Heft 12): 27. 1903. [Subfamily Cypripedioideae]. Cypripedium Lindl., Gen. Sp. Orchid. Pl. 525. 1840, pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrials, erect or rarely arched-pendent, cespitose, sun-loving or rarely shade-loving. Roots fibrous, branched. Rhizome subterranean, horizontal, shortly

Selenipedium 565

Fig. 496. Scuticaria steelei

Fig. 497. Selenipedium steyermarkii

creeping, thick. Stems cane-like, terete, simple or ± branched, often several meters long, enveloped by tubular, imbricate sheaths. Leaves plicate, vernation convolute, nonarticulate, distichous or subspiralled, thin, puberulent to pilose; blades linearlanceolate to elliptic. Inflorescences terminal at the apices of main stem or of branches, racemose or rarely with 1 lateral branch, pubescent, often glandular-pubescent, usually longer than leaves, erect, arched to pendulous; peduncle short, loosely enveloped by several sheaths, these often subfoliaceus; rachis spirally or subdistichously few- to many-flowered, lengthening with age, straight or ± flexuose; floral bracts conspicuous, subfoliaceous; pedicellate ovary elongate, terete, pubescent, horizontally spreading or subpendulous. Flowers relatively showy, resupinate, spreading or nodding, with spreading perianth segments, short-lived, opening in succession; sepals and petals membranous, variously pubescent, usually yellowish or greenish with red-brown spots or blotches; dorsal sepal free, elliptic, lateral sepals connate into a synsepal which is subequal to dorsal sepal, or ± narrower, apex bidentate or bifid; petals free, pendulous, much narrower than sepals, usually linear-oblong, apex acute or subtruncate. Lip fleshier and larger than the other perianth segments, slipper-like, inflated, with a cuneate base, the basal lobes or margins are involute and the margin around the opening incurved, variously pubescent or glabrescent. Column short, cylindric; anthers 2, situated on both sides of confluent stigmas; third anther modified into a shield-like staminode which is usually held perpendicular to the column and surpasses in length the stigmas; pollen in monads, powdery and viscose when mature; fertile stigmas 3, confluent into a triangular or suborbicular, sulcate body. Ovary 3-locular with axile placentation. Capsules ± fleshy, cylindric or fusiform, crowned with the persistent perianth.

566

O RCHIDACEAE

Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Brazil; 5 or 6 species, 2 in Venezuela, both in the flora area. Key to the Species of Selenipedium 1. 1.

Leaves to 6 times longer than wide; pedicellate ovary at least twice longer than its bract ........................................................................ S. palmifolium Leaves > 8 times longer than wide; pedicellate ovary shorter to 1.5 times longer than its bract ........................................................... S. steyermarkii

Selenipedium palmifolium (Lindl.) Rchb. f., Xenia Orchid. 1: 5, t. 2, figs. 6–9. 1854. —Cypripedium palmifolium Lindl., Gen. Sp. Orchid. Pl. 527. 1840. Terrestrial 50–200 cm tall; leaves 10–25 × 2–7 cm; sepals and petals translucent purple-brown; lip 23–27 mm long, orangebrown with some yellow at apex, the bottom violet, the folded-in margins violet; column greenish with brown tinges; anthers green; stigma light green. Rain forests, often in marshy areas, 200–400 m; Bolívar (La Escalera to Cerro Venamo region, Río Chicanán). Trinidad-Tobago, Guyana, Suriname, French Guiana, Amazonian Brazil.

Selenipedium steyermarkii Foldats, Bot. Soc. Venez. Ci. Nat. 21: 254, fig. 1. 1961. Terrestrial 150–300 cm tall, sun-loving; leaves 10–20 × 1–2.3 cm; sepals greenish brown without, brownish purple within, petals deep green; lip 2–3.6 cm long, pale yellow or pale brown at base grading to darker yellow-brown at apex, upper part of apex light bronze, 2 dark brownish purle marks flank either side of widest part of opening. Cloud forests, often in marshy areas, 900–1500 m; Bolívar (Amaruay-tepui, Auyán-tepui, Carrao-tepui, La Escalera to Cerro Venamo region). Guyana. ◆Fig. 497.

134. SIEVEKINGIA Rchb. f., Beitr. Syst. Pflanzenk. 3. 1871. [Subtribe Stanhopeinae]. by Gustavo A. Romero-González Cespitose epiphytes. Stem modified into ovoid, sulcate pseudobulbs of 1 internode, when young entirely concealed by distichous, scarious sheaths. Leaves 1(2), narrowly elliptic to narrowly ovate, acute, plicate, articulate, petiolate. Inflorescences 1–several, lateral, arising from the base of the pseudobulbs, pendent, racemose, few- to many-flowered. Flowers resupinate, often showy; floral bracts concave, narrowly ovate, shorter than the pubescent, pedicellate ovary. Sepals and petals membranous, spreading; sepals similar, narrowly to broadly ovate, acute; petals similar to the sepals but much narrower, the margins entire or fimbriate. Lip sessile, adnate to the column base, entire or 3-lobed, shallowly concave; disk with keels and/ or tooth-like calli. Column slender, elongate, recurved, semiterete, apically clavate and/or winged, footless; anther terminal, operculate, incumbent, 1-locular; pollinarium with 2 pollinia, yellow, narrowly obovate, cleft, cartilaginous, and dorsoventrally flattened, the tegula short, obtriangular, the viscidium linear, hooked. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 15 species, 1 in Venezuela. Sievekingia jenmanii Rchb. f., Flora 69: 450. 1886. Sievekingia dunstervilleorum Foldats, Acta Bot. Venez. 3: 399. 1968.

Vegetatively similar to Stanhopea; inflorescence to 8-flowered; flowers yellow; sepals and petals hyaline-white; lip and column light yellow; disk with 7–9 filiform (some-

Stigmatostalix 567

times bifid) appendices. Wet forests, 100– 1500 m; Bolívar (Cerro Venamo, Uei-tepui), Amazonas (Caño Iguana). Guyana. ◆Fig. 498. The report from Caño Iguana is based on a pollinarium found on a male euglossine bee (Euglossa sp.).

Fig. 498. Sievekingia jenmanii

135. SIGMATOSTALIX Rchb. f., Bot. Zeitung (Berlin) 10: 769. 1852. [Subtribe Oncidiinae]. Petalocentrum Schltr., Repert. Spec. Nov. Regni Veg. 15: 144. 1918. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, cespitose, erect. Rhizome abbreviate. Pseudobulbs aggregate, usually strongly flattened and ancipital, ellipsoid, oblong, to orbicular, 1-foliate to rarely bifoliate, when bifoliate, one of the leaves usually much larger than the other, pseudobulb clothed by several sheaths of which the 1–2 innermost are usually leaf bearing. Leaves conduplicate, articulate with their sheaths, subcoriaceous to coriaceous, usually narrowly elliptic or narrowly oblong-elliptic, basally sessile to subpetiolate. Inflorescences originating from pseudobulb base and emerging between the axils of the leaf sheaths, solitary or several from each pseudobulb, few- to laxly many-flowered, usually paniculate with abbreviate lateral branches, as long as or longer than leaves; branches basally clothed by few to several sheaths, flowers successive on each branch. Flowers small to medium-small, resupinate, with widely spreading perianth segments, usually yellow with brown or maroon bars or stripes at base of sepals and petals, these usually smaller than lip. Sepals free, or the later-

568

O RCHIDACEAE

als shortly connate at base, subequal. Petals similar to sepals or slightly different. Lip sessile to clawed, simple to lobed, sometimes of very complex morphology, with a callus-like, open oil gland near base, yellow with brown or purple zones, rarely entirely purplish. Column slender, elongate, dilated apically, usually curved in a swan’s neck fashion. Anther terminal, operculate, incumbent, often rostrate, unilocular; pollinia 2, cartilaginous, with a relatively broad, elongate tegula and a small viscidium. Stigma ventral, situated toward apex of column; rostellum conspicuous. Southern Mexico, Central America, Colombia, Venezuela, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 35 species, 2 in Venezuela, both in the flora area. Key to the Species of Sigmatostalix 1.

1.

Lip shallowly compressed below middle, dilated to a ± 4-parted apical lobe, base cuneate; sepals 4–5 times as long as broad; stipitate portion of column as long as or shorter than apical thickened zone; leaves 1– 2 times as long as the pseudobulbs ....................................... S. amazonica Lip sharply constricted at middle, dilated to a broadly divergent 2-lobed apex base truncate to rounded, sepals 2–3 times as long as broad; stipitate portion of column 2–3 times longer than thickened apical zone; leaves at least 4 times as long as pseudobulbs ......................... S. huebneri

Sigmatostalix amazonica Schltr., Beih. Bot. Centralbl. 42(2): 148. 1925. Epiphyte, medium-sized; leaves carinate on the backside; inflorescence paniculate with abbreviate, unifloral branches, arched or erect; flowers with sepals light maroon with yellow apices and brownish red spots, the lip yellow with maroon base, anther clear yellow. Rain forest, growing in the shade, 100–200 m; Amazonas (Paso de Ganado, Río Baría). Colombia, Ecuador, Peru, Bolivia. Sigmatostalix huebneri Mansf., Repert. Spec. Nov. Regni Veg. 36: 62. 1934. Epiphyte, small; inflorescence erect to erect-spreading; flowers with sepals and petals hyaline yellowish with brownish bands, lip solid yellow; column yellow, becoming yellowish brown toward base. Rain forests, 100–200 m; Amazonas (Río Baría, Río Guainía). Brazil. ◆Fig. 499.

Fig. 499. Sigmatostalix huebneri

136. SOBRALIA Ruiz & Pav., Fl. Peruv. Prodr. 720. 1794. [Subtribe Sobraliinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, lithophytes, or terrestrial herbs, small to very large, cespitose. Rhizome short. Secondary stems terete or laterally compressed, erect to pendulous,

Sobralia 569

simple or branched, partially or totally covered by leaf sheaths. Leaves distichous, plicate, sessile on vaginate sheath, commonly coriaceous or rigid, rarely thin, (mostly elliptic or ovate-elliptic, sometimes linear); sheaths tubular, tightly fitting the stem or funnel-shaped and rather loose, glabrous, verruculose, rugose or variously pubescent. Inflorescences terminal or axillary, sessile or pedunculate; peduncle terete, flattened or angled, racemose or paniculate. Floral bracts cucullate, minute to rather large. Flowers medium-small to very large and showy, mostly white, yellow or purple, rarely green, usually fugaceous, resupinate. Sepals and petals similar, free, membranaceous to rarely subfleshy, spreading, or campanulate, petals occasionally broader. Lip entire or more rarely lobed, usually more conspicuous than other perianth segments, basally enveloping the column, with a spreading lamina above variously ornate with bristles, keels or toothed crests. Column footless, slender, elongate, thinly-winged, the wings terminated in a pair of falcate processes on sides of stigma; stigma reniform, transverse; rostellum large, convex. Anther terminal, incumbent, operculate, bilocular. Pollinia 8, 4 in each locule, powderygranulose, rarely waxy. Capsule oblong or linear-oblong. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 100 species, ca. 27 in Venezuela, 15 of these in the flora area The northwestern Andean countries and southern Central America are particularly species-rich but there is a secondary center of diversity in the Guayana Highlands with several endemic species. The systematics of the genus are poorly understood because the flowers are fugacious and preserve poorly in herbarium specimens, hence, critical floral details, often readily observable in fresh material, are usually difficult if not impossible to recover from pressed material. Key to the Species of Sobralia 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(1). 5. 6(5). 6. 7(6).

Inflorescences axillary from upper stem internodes ................................ 2 Inflorescences terminal, 1-flowered to racemose ...................................... 5 Leaves linear or linear-elliptic, 4–8 mm wide, 5 or more times longer than wide; plants always growing terrestrially at river edges .... S. stenophylla Leaves elliptic or ovate-elliptic, 10–50 mm wide, 5–7 times longer than wide; plants not growing (or very rarely) at river edges ...................... 3 Stems freely branching in the upper 1/2; inflorescences 1-flowered; plants epiphytic or subterrestrial in cloud forest ......................... S. oliva-estevae Stems simple; inflorescences 2–7-flowered; plants terrestrial ................. 4 Lip broadest at middle or below, ± 3-lobed; petals of approximately the same width as the sepals ........................................................ S. dichotoma Lip broadest at the apical 1/3, simple; petals broader than the sepals .....................................................................................................S. speciosa Inflorescences with an evident peduncle or rachis, 1 to several-flowered ................................................................................................................ 6 Inflorescences without an evident peduncle, 1-flowered or successively 1-flowered ............................................................................................. 11 Stems to 50 cm tall; dorsal sepal < 4 cm long ........................................... 7 Stems > 80 cm (usually much more); dorsal sepal > 0.4–5 cm ................. 8 Peduncle elongate, laterally flattened; lip simple ........................ S. fragrans

570

O RCHIDACEAE

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

9.

10(9).

10.

11(5).

11.

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

13.

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

Peduncle abbreviate, cylindrical; lip simple or lobed ............................... 8 Leaves linear or linear-elliptic 4–8 mm wide, at least 15 times longer than wide ....................................................................................... S. stenophylla Leaves broader, narrowly elliptic to elliptic, at least 10 mm wide, 5– 10 times longer than wide ..................................................................... 9 Flowers purple with white and yellow in lip throat; leaves 3–5 times longer than wide; most inflorescences axillary; plants usually growing in tepui bog or swamp associations, rarely on sandstone .........S. speciosa Flowers white with yellow in lip throat; leaves 6 or more times longer than wide; plants growing usually in sandy soil or over sandstone outcrops; all inflorescences terminal ........................................................ 10 Lip with a bright yellow disk ornamented with conspicuous bright yelloworange veins and pseudopollen; flowers with waxy texture; petals and sepals widely spreading to recurved-reflexed; plants usually growing on sand or sandstone outcrops ............................................... S. liliastrum Lip with a pale yellow disk, keels elevated, of the same pale yellow color in the lower 3/4, apically white and lacking pseudopollen; flowers hyaline; petals and sepals not widely spreading, forming a subcampanulate flower; plants usually growing on granitic outcrops ........ S. granitica Leaf sheaths inflated (funnel-shaped) toward their apices, not tightly clasping the stems; plants terrestrial, usually growing on sand or sandstone outcrops, flowers with white, cream, or greenish sepals and purple petals and lip ...................................................... S. infundibuligera Leaf sheaths tubular, tightly clasping the stems; plants growing epiphytic or terrestrial over a variety of substrates; flowers totally purple, yellow, or white, not with the above combination of colors ............................ 12 Dorsal sepal < 4 cm long; lip with an apical lobe; plants usually epiphytic with stems often < 60 cm long ............................................................. 13 Dorsal sepal > 4.5 cm long; lip simple; plants terrestrial or epiphytic and then with purple flowers; stems > 60 cm (usually much longer) ....... 14 Leaves 17–25 cm long, 5–12 times longer than wide, long-acuminate; flowers white or pale cream-colored, with or without area purpled marked lip; inflorescences eventually developed into a twisted cone composed of umbricating bracts, the inflorescence elongates as it grows producing successive, individual flowers ................................... S. candida Leaves 10–15 cm long, 2.5–5 times longer than wide, shortly acuminate; flowers creamy-white or yellow, with an unmarked lip; inflorescences never developed into a twisted cone composed of imbricating bracts, the inflorescence never elongates and could have to 3 flowers open simultaneously ....................................................................... S. suaveolens Mature stems branched; flowers purple; plants epiphytic or lithophytic ...................................................................................................... S. sessilis Stems unbranched; flowers white, yellow, or rose-purple; plants terrestrial or epiphytic .................................................................................. 15 Leaf-bearing bracts subtending the inflorescence 3.5–5 times longer than wide, stiff; leaves narrowly elliptic, 4–6 times longer than wide; flowers with deep rose-purple sepals, petals paler rose, lip pale rose-purple with the center of the lamina yellow; plants terrestrial in open places ..................................................................................................... S. violacea

Sobralia 571

15.

Leaf-bearing bracts subtending the inflorescence 1.5–3 times longer than wide, foliaceous; leaves 2–3 times longer than wide; flowers white, yellowish, or entirely bright purple; plants epiphytic or terrestrial ....... 16 16(15). Margin of lip and the margins of the apical 1/2 of petals erose-fimbriate; leaves broadly elliptic, abruptly acuminate, 2–2.9 times longer than wide; plants growing in exposed positions .............................. S. fimbriata 16. Margin of lip and apical margins of petals smooth or wavy; leaves 3.2– 5 times longer than wide; plants growing mostly inside the forest, terrestrial or epiphytic ............................................................................. 17 17(16). Stems conspicuosly flattened in the final internode before the inflorescence; the whole inflorescence hidden under the sheath of the last leafbearing sheath, no scarious sheaths or bracts are displayed; the last leaf-bearing sheath 1.5–2 times longer than wide; plants usually epiphytic in lowland rain forests 80–500(–1350) m ................ S. macrophylla 17. Stems terete or only slightly flattened in the final internode before the inflorescence; only about 1/2 of the inflorescence hidden under the sheath of the last leaf-bearing sheath and scarious bracts and sheaths are displayed; the last leaf-bearing sheath 3–4 times longer than wide; plants terrestrial or low-epiphytic in rain or cloud forests (90–)300– 1300 m ........................................................................................... S. valida Sobralia candida (Poepp. & Endl.) Rchb. f., Fl. Serres Jard. Eur. 8: 247. 1853. —Cyathoglottis candida Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 56. 1836. Epiphyte, rarely subterrestrial, small to medium-sized for the genus, erect to pendent, cespitose; stems to 1 m long; inflorescence a cone-like bracted structure; flowers small for the genus, opening widely to campanulate; sepals and petals whitish, or pale cream color, with green tinged apices; lip white in lower 1/2, apically pale yellow with 2 or 3 bands of dark brown or purple spots; dorsal sepal to 32 mm long. Rain or cloud forests, 1100–1700 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region, Macizo del Chimantá). Ecuador, Peru. Sobralia dichotoma Ruiz & Pav., Fl. Peruv. Prodr. 1: 232, pl. 26. 1798. Sobralia cattleya Rchb. f., Gard. Chron. n.s. 7: 72. 1877. Terrestrial herb, stout; stems to 3 m tall; leaves 20–30 cm long; inflorescences with to 20 flowers; flowers purple or rose, often with brownish hues, with dark purple and yellow on the lip. Granitic outcrops, 100–200 m; Amazonas (Piedra Cocuy). Táchira; Colombia, Ecuador, Peru, Bolivia. The presence of Sobralia dichotoma in the flora area is unlikely. The species is known

only in the Andean portions of South America and has been reported from the flora area from an unnumbered collection by Spruce, which was not located for this treatment. In addition, a collection of Sobralia liliastrum Lindl. at AMES, also by Spruce and misidentified as S. dichotoma, from Río Casiquiare, suggests that Spruce probably also misidentified the Piedra Cocuy collection. It could also have been mistaken with any of the species of Epistephium growing in the Casiquiare region (E. hernandii Garay, E. sclerophyllum Lindl., etc.). These species display large showy purple flowers and occasionally bear lateral inflorescences when the plants are mature. Sobralia fimbriata Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 54. 1836. Terrestrial or rarely an epiphyte; stems to 1 m tall; flowers medium-sized for the genus, opening well; sepals and petals white; lip white with deep yellow and orange in center; lateral sepal 45–55 mm long. Rain or cloud forests, 700–1400 m; Bolívar (Altiplanicie de Nuria, Auyán-tepui), Amazonas (Cerro Aracamuni, Sierra Parima). Amazonian Brazil, Ecuador, Peru. ◆Fig. 500. Sobralia fragrans Lindl., Gard. Chron. 1853: 598. 1853.

572

O RCHIDACEAE

Epiphyte, small for the genus; stems 18– 50 cm long, erect to arching; inflorescence elongate with a laterally compressed peduncle; flowers small for the genus, white or whitish-flesh-colored; dorsal sepal 30–41 mm long. Rain or cloud forests, 50–1500 m; Bolívar (Cerro Sipapo, La Escalera to Cerro Venamo region, Río Caroní, lower Río Caura basin), Amazonas (upper Río Orinoco). Andean foothills (Apure, Táchira); Mexico, Guatemala, Belize, Honduras, Costa Rica, Panama, Colombia, Paraguay.

tepui tops, flowering through the year, each population has its own season, (700–)1200– 1900(–2200) m; widespread in Delta Amacuro, Bolívar, and Amazonas. Colombia, Guyana, Brazil. ◆Fig. 502. Populations from the Río Negro region of Amazonian Brazil and the Vaupes region of Colombia cited in the literature as belonging to Sobralia infundibuligera have white flowers and the lower surface of the leaves are purple, suggesting their membership in a different, probably undescribed species.

Sobralia granitica G.A. Romero & Carnevali, Harvard Pap. Bot. 5: 184. 2000. Terrestrial or lithophyte, vegetatively indistinguishable from the more common Sobralia liliastrum but usually with smaller plants and flowers, sun-loving; flowers white, callus dull clear yellow. Granitic outcrops or shrublands over sandy soils, 50– 200(–700) m; Bolívar (Serranía de los Pijiguaos), Amazonas (La Esmeralda, Río Autana, Río Cataniapo, Río Parguaza, Río Sipapo, San Carlos de Río Negro-Solano road). Although Sobralia granitica is readily distinguishable in the field from the closely related S. liliastrum by the characters described in the key, the distortion of the flowers upon pressing and dehydration, renders the two taxa almost impossible to recognize in herbarium specimens. Thus, the distributions of each of the members of this species pair are currently difficult to ascertain and their resolution awaits extensive field work. However, the evidence at hand seems to indicate that S. granitica is restricted to the lowlands of the Venezuelan Amazonas and the Parguaza region of Bolívar, probably extending into neighboring Colombia.

Sobralia liliastrum Lindl., Gen. Sp. Orchid. Pl. 177. 1833. —Cattleya liliastrum (Lindl.) Beer, Prakt. Stud. Orchid. 212. 1854. Sobralia elisabethae R.H. Schomb., Verh. Vereins Beförd Gartenbaues Königl. Preuss. Staaten 15: 137, t. 1, 2. 1841. Terrestrial, or very rarely a subepiphyte, rather large for the genus, erect, sun-loving; stems 1–3 m tall; flowers medium-sized to rather large for the genus, opening well; sepals and petals white, lip white with yellow in the throat, veins usually deep yellow or orange, rarely maroonish; sepals 5–7 cm long. Sandstone to granitic outcrops or sandy soils, mostly in open places, 50–1400(–1800) m; widespread in Bolívar and Amazonas. Colombia, Guyana, Suriname, French Guiana, Brazil (Amazonas, Pará, Roraima, Bahia). ◆Fig. 503.

Sobralia infundibuligera Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 3: 290. 1965. Sobralia liliastrum var. rosea Lindl., Fol. Orchid., Sobralia 4. 1854. Terrestrial to 2.5 m tall erect, sun-loving; leaves 20–30 × 3–6 cm; flowers not widely spreading, sepals and petals pale green, yellowish or cream, basally white, sometimes apically purple-tinged or very rarely wholly pale purple; lip deep purple-rose or lavender. Open rocky or swampy places, mostly at

Sobralia macrophylla Rchb. f., Bot. Zeitung (Berlin) 10: 713. 1852. Usually an epiphyte, medium-sized for the genus, cespitose, erect to arching; flowers campanulate, yellow or white, usually with paler margins, lip white or pale yellow with deep yellow in throat. Rain forests or very rarely cloud forests, 50–500(–1400) m; Bolívar (Urimán), Amazonas (Casiquiare, Río Cataniapo basin, upper Río Orinoco, Siapa, Sierra Tapirapecó). Andean foothills (Apure, Táchira); Costa Rica, Panama, Colombia, Ecuador, Brazil (Amazonian states). Sobralia oliva-estevae Carnevali & I. Ramírez, Ann. Missouri Bot. Gard. 77: 556. 1990. Epiphyte or subterrestrial, medium-sized to rather large, branched; inflorescences axillary, 1-flowered; flowers widely spreading,

Sobralia 573

bright purple with yellow and white in lip. Cloud forests, 1100–1300 m; Bolívar (La Escalera to Cerro Venamo region). Guyana. ◆Fig. 505. Sobralia oliva-estevae is closely related to S. speciosa C. Schweinf. but is epiphytic, branching, and the lateral inflorescences are 1-flowered. Sobralia sessilis Lindl., Edwards’s Bot. Reg. 27: misc. 3. 1841. Sobralia yauaperyensis Barb. Rodr. in Vellosia ed. 2, 1: 131. 1891. Sobralia violacea auct. non Linden & Rchb. f. 1846: sensu Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 1: 401. 1959, pro parte. Epiphyte, more rarely terrestrial or lithophyte, medium-sized to rather large for the genus, branching in the upper 1/2 of the stems, branches rooting or not; flowers sessile, with sepals spreading widely and petals held perpendicular to the dorsal sepal, deep or pale purple with a white or pale yellow zone or stripe at the apex of the lip throat, medium-sized for the genus with sepals 5–6 cm long. Rain forests or low elevation cloud forests, 50–900 m; Delta Amacuro (Río Cuyubini), Bolívar (Auyán-tepui, headwaters of Río Chicanán), Amazonas (near Puerto Ayacucho, Río Baría and Río Siapa). Coastal Range (Miranda), Andean foothills (Lara, Barinas, Zulia); Amazonian Brazil (Amazonas) and Colombia, Guyana, Suriname. ◆Fig. 501. The specimen Steyermark 89426 is placed hesitantly under this species because the flowers are described as being white. Sobralia sessilis belongs in a rather complex group that includes taxa such as Sobralia decora Lindl., S. fenzliana Rchb. f., S. fruticetorum Schltr., and others, all characterized by epiphytic plants with branching stems, and purple, sessile, solitary flowers. The flowers emerge from a condensed inflorescence with 1 or 2 bracts with small foliar blades. Extensive sampling of freshly preserved flowers and careful field and greenhouse work with live plants is required to adequately address the question of how many taxa are involved in this complex. The complex spreads throughout most of the Neotropics. The flowers in this complex are

particularly difficult to reconstruct from herbarium specimens. Sobralia speciosa C. Schweinf., Bot. Mus. Leafl. 19: 198, pl. 28. 1961. Terrestrial 1–2 m tall, erect, mediumsized to rather large for the genus; leaves 12–15 cm long; inflorescences axillary, 2–7flowered; flowers successive, deep purple with white in lip throat, yellow keels, and deep purple veins in back of the throat, rather large for the genus with sepals 5–8 cm long, opening well. Open boggy or rocky places at tepui summits or slopes, (500–) 1500–1800 m; Amazonas (Cerro Aracamuni, Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). Sobralia stenophylla Lindl., Fol. Orchid., Sobralia 2. 1854. Terrestrial to subaquatic or rheophyte, sun-loving, medium-sized for the genus, densely cespitose, erect to arching; leaves linear, 4–8 mm wide; inflorescences mostly axillary; flowers widely spreading to campanulate, purple with yellow and white on lip, small for the genus with sepals 3.5–3.9 cm long. Locally common along the margins of fast-flowing, black-water rivers, over sandstone or sandy soils, (400)–700–1700 m; Bolívar (upper Río Caroní basin). Guyana, Suriname, French Guiana, adjacent northern Brazil. ◆Fig. 504. Sobralia suaveolens Rchb. f., Gard. Chron. n.s. 9: 622. 1878. Usually an epiphyte, but sometimes terrestrial or a lithophyte, small for the genus, erect to arching; flowers campanulate, small for the genus, usually in pairs, creamy-white or yellowish. Rain or cloud forests, 200–1200 m; Bolívar (Cerro Guaiquinima, La Escalera to Cerro Venamo region, lower Río Caura). Andean foothills (Barinas); Panama, Amazonian Brazil, French Guiana. The Guayanan populations of this species have narrower leaves than the plants from the Andean foothills and Panama. Sobralia valida Rolfe, Bull. Misc. Inform. Kew 1909: 65. 1909. Sobralia biflora auct. non Ruiz & Pav. 1798: sensu Schweinf., Mem. New. York Bot. Gard. 14(3): 87. 1967.

574

O RCHIDACEAE

Sobralia sessilis auct. non Lindl. 1841: sensu Foldats in Lasser, Fl. Venez. 15(1): 189. 1970. Usually terrestrial, sometimes a low epiphyte, erect to arching, medium-sized for the genus; leaves to 25 cm long and 10 cm wide; flowers medium-sized for the genus, campanulate or with widely spreading segments, white or very rarely pale pink with some yellow on lip. Rain or cloud forests, (400–)700– 1500 m; a common species in the whole Venezuelan Guayana in suitable places, particularly in the basin of the Río Caroní. Panama, Colombia, Guyana, Ecuador, Brazil (Amazonian states), Bolivia. ◆Fig. 506.

Fig. 500. Sobralia fimbriata

Sobralia violacea Linden ex Lindl., Orchid. Linden. 26. 1846. —Cattleya violacea (Linden ex Lindl.) Beer, Prakt. Stud. Orchid. 215. 1854, non Rolfe 1889. Sobralia yauaperyensis auct. non Barb. Rodr. 1881: sensu Foldats in Lasser, Fl. Venez. 15(1): 199. 1970. Terrestrial, very rarely an epiphyte, medium-sized to rather large for the genus, erect, usually sun-loving, cespitose; flowers widely spreading, medium-sized to rather large, rose pink, purple or white. Open places at the edges of cloud forests or on tepui summits, 700–1500 m; Bolívar (La Escalera to Cerro Venamo region), Amazonas (Cerro Aracamuni, Cerro Aratitiyope, Cerro Marahuaka, Cerro Duida, Cerro Parú, Cerro Yapacana, Sierra de la Neblina). Andes (Barinas, Falcón, Mérida, Lara, Trujillo, Portuguesa, Táchira), Coastal Range (Falcón); Colombia, Peru, Bolivia. The Peruvian and Bolivian collections currently assigned to this species might be referable to other species.

Fig. 501. Sobralia sessilis

Sobralia 575

Fig. 502. Sobralia infundibuligera

Fig. 503. Sobralia liliastrum

Fig. 504. Sobralia stenophylla

576

O RCHIDACEAE

Fig. 505. Sobralia oliva-estevae

Fig. 506. Sobralia valida

Stanhopea 577

137. SOLENIDIUM Lindl., Orchid. Linden. 15, no. 79. 1846. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, cespitose or shortly creeping, erect. Rhizome abbreviate, enveloped by scarious sheaths. Pseudobulbs pear-shaped to oblong, laterally compressed, apically 1- or 2-leaved, basally enveloped by several imbricate sheaths the innermost 1 or 2 may have foliar blades. Leaves conduplicate, articulate, coriaceous, erect, linear or linear-oblong, sessile or shortly pseudopetiolate. Inflorescence lateral, originating from the base of the mature pseudobulb or from the new growth, erect, racemose, lax or ± densely many-flowered, bearing flowers in the upper 1/2; peduncle shorter than leaves, remotely bracteate, terete; rachis straight or ± zigzag; floral bracts shorter than pedicellate ovary, inconspicuous; pedicellate ovary long, terete. Flowers resupinate, ± showy, perianth segments subfleshy, widely spreading, greenish with brown or maroon blotches. Sepals subequal, free, obovate to oblanceolate, rounded to obtuse, the laterals somewhat oblique; petals subequal to sepals. Lip erect, free from column, in the apical 1/2 expanded into a transversely oblong to suborbicular or subquadrate blade, basally broadly clawed, the claw with an elongate callus composed of 2 parallel keels or a convex platform which is divaricately bidentate at apex; callosities and claw pubescent. Column semiterete, as long as or shorter than the callosities, footless, apically or medially winged, wings auriculate or dolabriform; anther operculate, incumbent, 1-locular, apically produced into a tongue-like appendix; clinandrium shallow, entire to erose; pollinia 2, waxy, with an elongate tegula and a small, strap-like viscidium; rostellum transverse; stigma ventral. Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia; 2 species, 1 in Venezuela. Solenidium lunatum (Lindl.) Kraenzl. in Engl., Pflanzenr. IV 50(Heft. 80): 316. 1922. —Oncidium lunatum Lindl., Edwards’s Bot. Reg. 23: sub t. 1920. 1838. Epiphyte; leaves 5–13 cm long; inflorescences 9–25 cm long; dorsal sepal 8–11 mm long. Rain forests, 100–200 m; Bolívar (Río Erebato), Amazonas (near La Esmeralda). Guyana, Ecuador, Peru, northern and western Brazil, Bolivia. ◆Fig. 507.

Fig. 507. Solenidium lunatum

138. STANHOPEA Frost ex Hook., Bot. Mag. 56: t. 2948, 2949. 1829. [Subtribe Stanhopeinae]. Ceratochilus Lindl., Bot. Cab. 15: t. 1414. 1828, non Blume 1825. Stanhopeastrum Rchb. f., Bot. Zeitung (Berlin) 10: 927. 1852. by Gustavo A. Romero-González Cespitose epiphytes or sometimes terrestrial herbs. Stem modified into ovoid, sulcate pseudobulbs of 1 internode, when young entirely concealed by distichous, scarious sheaths. Leaf 1, narrowly to broadly ovate, acute to acuminate, plicate, articulate, petiolate. Inflorescences 1–several, lateral, arising from the base of the pseudobulbs, pendent, racemose, generally few-flowered. Flowers fleshy, morpho-

578

O RCHIDACEAE

logically complex, nonresupinate, often showy; floral bracts concave, narrowly to broadly ovate, shorter than the generally elongate pedicellate ovary. Sepals and petals membranous, spreading to reflexed; sepals narrowly to broadly elliptic, the lateral ones usually wider than the dorsal one, often basally connate; petals smaller than the sepals, oblong, acute, the margins undulate or not. Lip adnate to the base of the column, oblong, basally sacciform furnished with a short spur, with a transverse, antrorse, anteriorly excavate keel near the apex, the apex triangular, or lip continuous with the base of the column and divided into a hypochile, mesochile, and epichile; hypochile subglobose, excavate; mesochile short, with 2 fleshy, horn-like, arched processes; epichile generally articulate to the apex of the mesochile, flat, concave, or convex, apically entire, or 2- or 3-lobed. Column elongate, terete, generally dilated toward the apex, winged or not; anther terminal, operculate, incumbent, 1locular; pollinarium with 2 yellow pollinia, ovate, cleft, cartilaginous, and dorsoventrally flattened, the tegula short, ligulate or narrowly obtriangular, the viscidium subtriangular or crescent-shaped. Southern Mexico, Central America, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, northern Argentina; 40 species, 5 in Venezuela, 3 of these in the flora area. Key to the Species of Stanhopea 1. 1. 2(1). 2.

Mesochile of the lip with horn-like projections; flowers yellow, spotted with reddish purple markings ...................................................... S. wardii Mesochile of the lip without horn-like projections; flowers white ........... 2 Lip < 5 cm long, with 2 erect horns at the base ............................. S. candida Lip > 5.5 cm long, with 2 hooked horns at the base ................. S. grandiflora

Fig. 508. Stanhopea candida

Stelis 579

Stanhopea candida Barb. Rodr., Gen. Spec. Orchid. 1: 101. 1877. Stanhopea randii Rolfe, Bull. Misc. Inform. Kew 1894: 363. 1894. Epiphyte; racemes 2–5-flowered; flowers white; dorsal sepal to 5 cm long. Wet forests, 50–200 m; Amazonas (Isla Hormiga, San Antonio). Colombia, Ecuador, Peru, Brazil (Pará), Bolivia. ◆Fig. 508. Stanhopea grandiflora (Lodd.) Lindl., Gen. Sp. Orchid. Pl. 158. 1832. —Ceratachilus grandiflorus Lodd., Bot. Cab. 15: t. 1414. 1828 [1829]. Stanhopea eburnea Lindl., Edwards’s Bot. Reg. 18: t. 1529. 1832. Epiphyte; racemes 1- or 2-flowered; flowers white; dorsal sepal 7.5 cm long. Wet forests, 50–500 m; Delta Amacuro (Río Amacuro, Serranía de Imataca), Bolívar (Canaima, Cerro Bolívar area, Cuchari-tepui, Río

Caura, Río Paragua), Amazonas (Río Cataniapo, Río Orinoco, between Samariapo and San Fernando de Atabapo, Tamatama). Coastal Cordillera of Miranda, Sucre (Cerro Patao); Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil (Amazonas, Pará). Stanhopea wardii Lodd. ex Lindl., Sert. Orchid. t. 20. 1838. Stanhopea aurea Lodd. ex Lindl., Edwards’s Bot. Reg. 27: misc. 11. 1841. Stanhopea venusta Lindl., Edwards’s Bot. Reg. 27: misc. 11. 1841. Epiphyte; racemes 3–10-flowered; flowers yellow to greenish spotted with reddish purple. Wet forests, 600–1800 m; Bolívar (fide Dunsterville, locality not specified). Venezuelan Coastal Cordillera, Zulia (Perijá); Central America, Colombia.

139. STELIS Sw., J. Bot. (Schrader) 1799: 239. 1799 [1800], nom. cons. [Subtribe Pleurothallidinae]. Humboldtia Ruiz & Pav., Fl. Peruv. Chil. Prod. 121, t. 27, 1794, nom cons., non Vahl 1794. Dialissa Lindl., Ann. Mag. Nat. Hist. 15: 107. 1845. Apatostelis Garay, Bot. Mus. Leafl. 27: 185. 1980. by Germán Carnevali and Ivón M. Ramírez-Morillo Mostly epiphytes, but many taxa secondarily terrestrial or lithophytic, cespitose or creeping, minute to fairly large for the Pleurothallidinae. Rhizome prostrate or ascendent, sometimes distinctly pendent, often branching and then plants mat-forming. Stems almost absent to very elongated, shorter than to much longer than the single apical leaf, terete, covered partially or totally by sheaths, in many species apically proliferous (then producing chains of superposed stem). Leaves single on top of the stems, articulate, thinly coriaceous to fleshy or even subterete, usually elliptic or elliptic-obovate, but sometimes linear or oblong, sessile to conspicuously subpetiolate. Inflorescences racemes, few- to many-flowered, solitary to several, simultaneous or successive, originating on or just below apex of the stem from a swollen annulus, basally often subtended by a spathe, rachis straight or zigzag, rarely very abbreviate and then inflorescences umbellate. Flowers opening successively or simultaneously, often sensitive to light or temperature and then opening and closing at particular times of the day, in response to the right stimulus; usually resupinate, spreading widely to campanulate, the triangular sepals are variously connate forming a triangular shaped-flower made of 3 triangles fused at base, the central portion of the triangle is occupied by an apparatus conformed by the column, the 2 petals and the lip. Perianths segments membranaceous to thinly coriaceous, usually white, green, yellow, or purple, often variously pubescent. Sepals connate basally, 1-, 3-, 5-, 7-, or 9-veined, similar or the laterals fused together to form a bila-

580

O RCHIDACEAE

biate flower, or the dorsal sepal larger than the other 2; petals usually much smaller than the sepals, fleshy, transverse, marginally thickened, fitting snugly in the shallow tube formed by the base of the sepals and closely flanking the lip and column. Lip small, thick, usually shaped like 1/4 of a sphere, the uppermost plane of the lip resting beneath the column and often bearing a glenion (a small area with a shiny surface that appears wet) and one or few keels or ridges, the lip basally sessile on the column. Column short and broad, apically dilated. Anther apical, incumbent, operculate. Pollinia 2, usually pear-shaped. Stigma apical, tranverse, 1- or 2-lobed, when 2-lobed the lateral lobes protrude to either side of the anther under the flap on the rostellum, the stigma lobes often so elongated that the receptive surfaces are held far to either side of the anther. Capsules ellipsoid. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 500–600 species, ca. 80 in Venezuela, 23 of these in the flora area. Stelis is a neotropical genus currently undergoing revision. It ranges from Mexico to southeastern Brazil, but is particularly diverse in the Andean countries and in southern Central America. Stelis is characterized by the ± triangular structure of the small, complex flowers. It is a difficult genus in that important diagnostic characters of the flowers are many times irretrievably lost upon pressing and drying. These flower characters, particularly those of the fleshy lip and petals, are also difficult to describe and interpret. All these factors have rendered taxonomists of the past reluctant to study the genus. Some of the older names (e.g., Stelis argentata Lindl.) have indiscriminately been applied to a large assortment of superficially similar entities, which in many floristic treatments resulted in the circumscription of widespread, variable, mostly polyphyletic taxa. The unraveling of these morphologic and biogeographical problems is currently underway by modern Stelis students. The key in this treatment was written based mostly on vegetative and inflorescence features, but good floral material is often essential for positive determinations of some of the taxa. The flower characters used (number of veins of the sepals and petals, shape of the sepal cup, apicule of the lip) are usually easy to retrieve from good rehydrated material. Good rehydrations of pressed flowers (if they have not been too severely flattened) can be obtained by soaking the flowers overnight in concentrated ammonia and then rinsing them in water. Sometimes, good results are obtained by boiling the flowers in water for 1–2 minutes and letting them rest for at least 30 minutes. Fresh or pickled flowers are always the best choice. Besides the entities included in this treatment, there is evidence indicating the presence of several other taxa of the genus in the flora area, but in some cases the material is insufficient for positive determination. The identification of some of the Venezuelan Guayana Stelis taxa is only tentative in lieu of a good revision of the species of the genus. The approach taken by the authors of this treatment has been in most cases conservative in applying names used by previous authors (Foldats, Dunsterville, and Garay) to some of the taxa, but in some cases the identifications are probably inaccurate (e.g., Stelis effusa is a Costa Rican taxon, unlikely to occur in the Pantepui area). Until the species circumscription and geographical ranges of the taxa of the genus are better understood, this treatment (and most other regional treatments of Stelis) will remain tentative.

Stelis 581

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

2.

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

5.

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

7.

8(7).

8.

9(3). 9.

Flowers bilabiate, i.e., with lateral sepals totally connate into a synsepal which is subequal to dorsal sepal .......................................................... 2 Flowers stellate, i.e., with the 3 sepals equally connate and subequal in shape and size ........................................................................................ 3 Plants small to medium-sized; stems 3.5–9.5 cm long; dorsal sepal 6 mm or less long; petals basally broadly cuneate, in natural position touching their margins over column; rhizome abbreviate; lip when viewed from above truncate ..................................................................... S. aviceps Plants large; stems 10–20 cm long; petals basally cordate, in natural position overlapping behind the column; rhizome ± elongate; lip when viewed from above broadly obtuse .............................................S. maxima Dorsal sepal 4- or 5-veined ........................................................................ 4 Dorsal sepal 3-veined ................................................................................. 9 Free portion of lateral sepals shorter than its width at base (at the indentation point) ........................................................................................... 5 Free portion of the flattened lateral sepals longer than its width at base (at the indentation point) ...................................................................... 7 Adult leaves to 30 mm long; inflorescences arching to suberect (erect when young), successively flowered, continually lengthening; rachis very lax on old inflorescences, sublax and strongly zigzag on short, young inflorescences; floral bracts relatively inconspicuous ................... S. tolimensis Adult leaves at least 50 mm long (usually much longer, to 120 mm); inflorescences erect, straight, simultaneously flowered and not continuously lengthening rachis straight and dense; floral bracts relatively conspicuous and large ............................................................................ 6 Floral bracts imbricate, covering completely the rachis; callus of lip smooth; apex of lip and petals smooth ......................................... S. bangii Floral bracts lax to subimbricate, leaving portions of rachis naked; callus of lip granulose; apex of lip and petals granular-verruculose ..... S. zonata Leaves fleshy-coriaceous, semiterete; petals tranversely callose, the apex extending into a fleshy protrusion; lip apically with a transverse callus, not apiculate ................................................................................... S. effusa Leaves coriaceous, conduplicate; petals neither tranversely callose, nor with the apex extending into a fleshy protrusion; lip without an apically transverse callus, apiculate at apex ........................................ 8 Rhizome abbreviate; leaves gradually tapering toward base, lamina obovate-elliptic; inflorescences usually much longer than the subtending leaf; dorsal sepals similar to laterals; petals with a granular-verrucose apical surface ........................................................................... S. argentata Rhizome shortly but distinctly creeping or ascending; leaves abruptly petiolate, lamina broadly elliptic; inflorescences shorter to ± equal to the subtending leaf; dorsal sepal unequal to lateral ones; petals with a smooth apical surface ............................................................. S. tridentata Lip with a small, upturned apicule ......................................................... 10 Lip without a small, upturned apicule .................................................... 13

582

O RCHIDACEAE

10(9). 10.

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

12.

13(9). 13. 14(13). 14. 15(14).

15.

16(15). 16.

17(16).

17. 18(17).

Petals 1-veined, inflorescences 2–4 times longer than the leaves; leaves (including petiole) to 24 mm long (usually shorter) ............. S. guianensis Petals 3-veined, inflorescences shorter than to 2 times longer than the leaves (when 2 times longer than the leaves then sepals at least 5 mm long), leaves (including petiole) at least 30 mm long (usually longer) .............................................................................................................. 11 Inflorescences much surpassing the leaves, free portions of sepals at least 5 mm long ................................................................................. S. argentata Inflorescences shorter or a little longer than the leaves (to 1.5 times longer); free portions of the sepals 1–2.5 mm long ............................. 12 Leaves gradually tapering toward base, lamina narrowly elliptic-obovate, < 12 mm wide; rhizome abbreviate, plants cespitose; sepals flat or convex, pointing forward or spreading in natural position, the ventral face glabrous; dorsal sepal similar to lateral ones; plants usually from intermediate-elevation cloud forests ................................................ S. pygmaea Leaves abruptly petiolate, lamina broadly elliptic, at least 15 mm wide; rhizome shortly but distinctly creeping; sepals convex, somewhat reflexed in natural position, the ventral face pubescent; dorsal sepal unequal to laterals (a little longer and narrower); plants from low-elevation rainforests ........................................................................ S. tridentata Petals 1-veined ......................................................................................... 14 Petals 3-veined ......................................................................................... 21 Rhizome distinctly creeping and branching, plants mat-forming; leaves linear to linear-oblanceolate ......................................................... S. pusilla Rhizome abbreviate, usually not branching, plants cespitose; leaves broader than linear .............................................................................. 15 Inflorescences 2–5 times longer than subtending leaf (the peduncle always longer than the subtending leaf); leaves to 21 mm long (usually shorter); free portion of dorsal sepal broader than its length (to about as long) ..................................................................................... S. latisepala Inflorescences shorter than to about as long as (rarely somewhat longer than) subtending leaf (the peduncle always shorter than the subtending leaf); leaves at least 30 mm long (usually longer); free portion of dorsal sepal longer than its width at indentation point ..................... 16 Stems (2–)2.5–5 times shorter than leaves; when young totally enclosed by tubular sheaths; inflorescences produced one at a time ............... 17 Stems longer to 1.5 times shorter than leaves, not totally enclosed by tubular sheaths when young; inflorescences usually produced 2–4 at a time ....................................................................................................... 19 Flowers pink to purple; inflorescences prostrate or arching; apex of sepals with a tuft of long, conspicuous, sublanate, white hairs; petals glabrous; column with a ventral, simple stigmatic surface .............. S. garayi Flowers pale green; inflorescences erect; apex of sepals glabrous; column with an apical, 2-lobed stigmatic surface ............................................ 18 Inflorescences usually shorter than leaves, rarely as long; sepals lanceolate, longer than wide, with margins not marginate, reflexed in natural position; apices of petals and callus of the lip pubescent ............................................................................................ S. santiagoensis

Stelis 583

18.

19(16). 19. 20(19). 20. 21(13).

21.

22(21).

22.

23(22).

23.

24(23).

24.

Inflorescences longer than the leaves, rarely only as long; sepals broadly ovate, as broad as long or broader than long, margins concave or at least not reflexed, marginate; apices of petals and callus of the lip glabrous .......................................................................... S. ophioglossoides Lateral sepals dorsally ecarinate; sepals broader toward the base or at the basal 1/3; lip obtuse to subacute, basally pubescent ............ S. minimiflora Lateral sepals dorsally carinate to winged; sepals broader toward the middle; lip truncate, totally glabrous .................................................. 20 Lip with a smooth, concave ventral face; stigmatic surface single, ventral ......................................................................................................... S. alata Lip with a deeply sulcate, convex ventral face; stigmatic surfaces 2, apical, protruding at each side of the anther .................................. S. fendleri Dorsal sepal longer than the lateral ones, hence sepals forming an inaequilateral triangle, their internal surface verruculose; petals transversely rhombic, their superior and inferior margins acute, 2– 3 times wider than long ............................................................. S. fraterna Dorsal sepal subequal to the lateral ones, hence sepals forming an equilateral triangle, their internal surface smooth or inconspicuously verruculose to pubescent; petals usually subquadrate to obovate, about as long as wide or longer than wide, their superior and inferior margins flat or obtuse ........................................................................................ 22 Leaves linear oblanceolate,10–12 times longer than wide; sepals acute; lip longer than wide, in ventral outline subquadrate, in cross section hemicylindric, basally pubescent, with an apicule pointing frontward ............................................................................................ S. schomburgkii Leaves obovate to narrowly-elliptic, 2–7(–10) times longer than wide; sepals obtuse to broadly obtuse; lip as long as wide or wider than long, usually like 1/4 of a hemisphere or broadly oblong, concave, basally glabrous, apically rounded or truncate .................................................... 23 Peduncle of inflorescence much longer than subtending leaf, at least twice as long; stems 3–13 mm long, leaves obovate or obovate-elliptic, 2– 3 times longer than wide; lip broadly oblong, concave .............. S. obovata Peduncle of inflorescence usually much shorter than subtending leaf, rarely about as long; stems (10–)20-100 mm long; leaves narrowly elliptic (3–)5–10 times longer than wide; lip like 1/4 of a hemisphere ...... 24 The peduncle is < 1/6 the total length of the inflorescence; internodes of the rachis about twice as long as the floral bracts; leaves 4–20 mm wide; lip in lateral view about as thick as long or thicker than long ..... S. cucullata The peduncle is at least 1/3 of the total length of the inflorescence; internodes of the rachis at least 3 times longer than the floral bracts; leaves 3–7 mm wide; lip in lateral view longer than thick .............. S. intermedia

Stelis alata Lindl., Fol. Orchid., Stelis 18. 1859. —Apatostelis alata (Lindl.) Garay, Bot. Mus. Leafl. 27: 187. 1979. Stelis tenuilabris Lindl., Fol. Orchid., Stelis 18. 1859. Epiphyte, usually in exposed positions; leaves 20–50 mm long, longer than stems; in-

florescences shorter than or rarely as long as the subtending leaf; flowers greenish yellow or rarely with pale lavender tinges; dorsal sepal 1.5–3 mm long. Cloud forests, (700–) 1300–2400 m; Bolívar (Amaruay-tepui, Jaua-Sarisariñama massif, La Escalera to Cerro Venamo region, Macizo del Chimantá),

584

O RCHIDACEAE

Amazonas (Cerro Marahuaka, Cerro Yutajé). Aragua, Distrito Federal, Mérida, Yaracuy; Colombia, Ecuador, Guyana. If Stelis tenuilabris Lindl. is considered distinct from S. alata, a collection from Cerro Jaua (Dunsterville & E. Dunsterville 1320) represents S. tenuilabris. Stelis argentata Lindl., Edwards’s Bot. Reg. 28: misc. 64. 1842. Epiphyte; leaves 65–100 mm long, 7–16 mm wide; inflorescences much surpassing the leaves, to 25 cm long; flowers pale greenish maroon or entirely pale maroon, sepals sometimes hairy on the ventral surface; dorsal sepal 100–800(–1300) m; Delta Amacuro (Serranía de Imataca), Bolívar (La Escalera to Cerro Venamo region, Río Canaracuni, Río Paragua). Anzoátegui, Apure, Aragua, Distrito Federal, Mérida, Monagas; Costa Rica, Panama, widespread in South America. As here circumscribed, Stelis argentata is an extremely variable species, with a wide ecological and geographical range. It may eventually prove to encompass several closely related species. Stelis aviceps Lindl., Fol. Orchid., Stelis 16. 1859. Stelis cupuligera auct. non Rchb. f. 1854: sensu Dunst. & Garay, Venez. Orchid. Ill. 5: 294. 1976. Epiphyte; leaves 40–100 mm long, 7–16 mm wide; inflorescences usually surpassing the subtending leaf; flowers dull yellow or pale greenish; dorsal sepal 5–7 mm long. Rain or low-elevation cloud forests, (600–) 1000–1800 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region, Macizo del Chimantá [Abacapá-tepui], Ptari-tepui), Amazonas (Cerro Cuao). Ecuador. Stelis bangii Rolfe, Mem. Torrey Bot. Club. 4: 260. 1895. Stelis tristyla auct. non Lindl. 1838: sensu Foldats in Lasser, Fl. Venez. 15(2): 165. 1970; Dunst. & Garay, Orchids Venez. Ill. Field Guide 1000. 1979. Epiphyte, rarely a lithophyte; leaves 3–15 cm long, 8–30 mm wide; inflorescences about as long as or longer than the leaves, with several flowers opening simultaneously; flowers dark purple, or greenish purple, sen-

sitive, opening and closing in response to environmental cues, free portion of the sepals 5–7 mm long, 4–6 mm wide. Rain or cloud forests, 800–1600(–1900) m; Bolívar (La Escalera to Cerro Venamo region, Sierra de Maigualida, widespread on tepuis), Amazonas (Cerro Aracamuni). Aragua, Distrito Federal, Mérida, Miranda, Monagas; Colombia, Bolivia, Trinidad, Guyana, Brazil (northern states). The status of Stelis bangii and its relationships with S. zonata are still unresolved. There seems to be only one species with broad, 5-veined, long-connate sepals on an intermediate to large-sized plant in the Guayanan area. This taxon or group of taxa are also characterized by distichous, closely set floral bracts. Stelis zonata has been separated from this taxon by Foldats (as S. grossilabris) by the basal portion of the callus being granulose. The texts and the illustrations published by Dunsterville and Garay seem to suggest that there are two entities based on the closeness of the floral bracts and in the shape of the expanded sepal cup. The study of the abundant available material from the flora area, however, indicates that these two characters are highly variable and that we could be dealing with just one species. The degree of granulosity of the lip callus is also variable and does not seem to correlate with the variation of the other 2 characters. Until a thorough revision of this complex of species of Stelis (also including S. muscifera Rchb. f., S. acuifera Lindl., S. tristyla Lindl., and others) is carefully undertaken, we have approached this complex conservatively. We follow Garay in the application of the name S. bangii to the entity with densely imbricate floral bracts. Stelis cucullata Ames, Sched. Orchid. 6: 52. 1923. Epiphyte; leaves 2–12 cm long, 4–20 mm wide; inflorescences as long as or longer than the subtending leaves; flowers yellow, with widely spreading sepals; sepals pubescent or glabrous; dorsal sepal 1.3–2 mm long. Rain or cloud forests, 100–1500(–2000) m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region, Río Toro), Amazonas (Río Cataniapo, Tamatama). Aragua, Monagas, Táchira; Costa Rica, Nicaragua.

Stelis 585

Stelis effusa Schltr., Repert. Spec. Nov. Regni. Veg. 3: 247. 1907. Epiphyte; leaves longer or as long as the subtending stems, very fleshy, 2–4.8 cm long, 3–10 mm wide; inflorescences solitary, much longer than the subtending leaves; flowers opening in slow succession, yellow or purplish; dorsal sepal 1.8–3 mm long. Cloud forests, 1200–2100 m; Bolívar (Jaua-Sarisariñama massif, La Escalera to Cerro Venamo region). Costa Rica. ◆Fig. 509. The Venezuelan Guayana material of Stelis effusa appears to differ from the type (Costa Rican material) in having 3-veined sepals (instead of 5-veined) and fleshier leaves. It probably represents an undescribed taxon. Stelis fendleri Lindl., Fol. Orchid., Stelis 3. 1859. Epiphyte; leaves 5–12 cm long, 8–19 mm wide, shorter or as long as the stems; inflorescences solitary or fasciculate (to 7), about as long as the subtending leaves, rarely longer; flowers opening simultaneously, yellowish; dorsal sepal 1.3–2.3 mm long. Rain forests, 400–600 m; Bolívar (Río Icabarú). Aragua, Distrito Federal, Mérida; Colombia. Stelis fendleri is locally common in places in the Venezuelan Coastal Range, where it usually inhabits cloud forests at 1500– 2500 m. Stelis fraterna Lindl., Fol. Orchid., Stelis 14. 1859. Stelis papaquerensis auct. non Rchb. f. 1849: sensu Foldats in Lasser, Fl. Venez. 15(2): 138. 1970; Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 3: 305. 1965. Epiphyte; leaves 4–16 cm long, 8–35 mm wide, longer or as long as the stems; inflorescences solitary, usually longer than the subtending leaf; several flowers opening simultaneously, greenish or purplish; dorsal sepal 1.8–3.1 mm long. Rain forests, 100– 500 m; Bolívar (Cerro Guaiquinima, La Escalera to Cerro Venamo region, Roraimatepui foothills), Amazonas (near Puerto Ayacucho). Táchira; Trinidad, Guyana, Brazil (Amazonas). Stelis garayi (Dunst.) Carnevali & I. Ramírez, Ann. Missouri Bot. Gard. 77:

556. 1990. —Apatostelis garayi Dunst., Amer. Orchid Soc. Bull. 50: 1075. 1981. Epiphyte growing low on trees; leaves 5–7 cm long; inflorescence arching or pendent with 20–35 flowers opening simultaneously; perianth segments pink to purple; dorsal sepal 1–1.2 mm long. Rain forests, 200–300 m; Bolívar (Salto Paraván on the Río Yuruán), Amazonas (Río Autana). Ecuador. Stelis guianensis Rolfe, Trans. Linn Soc. Bot. ser. 2, 6: 59. 1901. Epiphyte; leaves 1.5–2.5 cm long, 2–5 mm wide; inflorescences 40–70 mm long, much surpassing the leaves; flowers deep magenta; dorsal sepal 2.5–3 mm long. Rain or cloud forests, 100–1400 m; Bolívar (La Escalera to Cerro Venamo region, Roraimatepui). Adjacent Guyana. Stelis intermedia Poepp. & Endl., Nov. Gen. Sp. Pl. 1: 46. 1836. Epiphyte, rarely a lithophyte; leaves 30– 50 mm long, 2–5 mm wide, shorter or as long as the stem; inflorescences shorter to a little longer than the subtending leaves; flowers dull yellow to orange-yellow; dorsal sepal 1– 1.7 mm long. Cloud forests, 1000–2000 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region, Macizo del Chimantá [Abacapá-tepui]). Peru. The Guayanan material could represent a different taxon than the Peruvian material of Stelis intermedia. Stelis latisepala C. Schweinf., Bot. Mus. Leafl. 19: 199. 1961. Epiphyte; leaves 8–20 mm long, 3–5 mm wide, much longer than the stems; inflorescences 25–45 mm long, much surpassing the subtending leaf; scape and pedicels purple or brown-purple, flowers totally purple, brownish purple, coppery brick red with tawny yellow margins; dorsal sepal 2.2–2.6 mm long. Cloud forests or shrublands, 1600–2200 m; Bolívar (Auyán-tepui, Kamarkawarai-tepui, Macizo del Chimantá [Apacará-tepui]). Endemic. ◆Fig. 510. Stelis maxima Lindl., Ann. Mag. Nat. Hist. 15: 106. 1845. Epiphyte or lithophyte; leaves 7–12 cm long, 10–25 mm wide, much shorter than the

586

O RCHIDACEAE

stem; inflorescences approximately as long as the leaves, flowers opening in slow succession, 5–7 open simultaneously at any given time; flowers bilabiate, yellow-white or greenish, nonresupinate; dorsal sepal 7–12 mm long. Locally common in tepui bogs or dwarf forests, 1700–2200 m; Amazonas (Sierra de la Neblina). Endemic. Stelis minimiflora Schltr., Repert. Spec. Nov. Regni Veg. 27: 31: 1924. —Apatostelis minimiflora (Schltr.) Garay, Bot. Mus. Leafl. 27: 189. 1979. Stelis braccata auct. non Rchb. f. & Warsz. 1854: sensu Foldats in Lasser, Fl. Venez. 15(2): 76. 1970; Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 1: 417. 1959. Epiphyte; leaves succulent, often subterete, 5–10 cm long, 5–15 mm wide; flowers yellowish or greenish; dorsal sepal 1.2–2 mm long; inflorescences 1–4 simultaneous with many flowers opening at a time, shorter or about as long as the subtending leaf. Cloud forests, ca. 1100 m; Bolívar (La Escalera to Cerro Venamo region). Aragua, Miranda, Yaracuy. Ecuador, Colombia. Stelis minimiflora is locally very common in the Venezuelan Coastal Range, but has only been collected once in the flora area. Stelis obovata C. Schweinf., Bot. Mus. Leafl. 19: 202, pl. 29. 1961. Stelis maderoi auct. non Schltr. 1920: sensu G.A. Romero & Carnevali, Orchids Venez. Ill. Field Guide 1030. 2000. Epiphyte; leaves 15–25(–35) mm long, usually much longer, rarely as long as the stems; inflorescences 30–70 mm long, much surpassing the subtending leaves; flowers cream-colored or yellow; dorsal sepal 1.5–2 mm long. Cloud forests or shrublands, (1400–)1720–2000; Bolívar (Auyán-tepui, Macizo del Chimantá), Amazonas (Cerro Huachamacari, Cerro Yutajé, Sierra Parima). Endemic. Stelis ophioglossoides (Jacq.) Jacq., J. Bot. (Schrader) 2(4): 239, t. 2, fig. 3a–g, 1799. —Epidendrum ophioglossoides Jacq., Enum. Syst. Pl. 29. 1760. Stelis trigoniflora (Sw.) Garay, Bot. Mus. Leafl. 26: 25. 1978. —Epidendrum trigoniflorum Sw., Prodr. 125. 1788.

Epiphyte; leaves 6–12 cm long, 7–9 mm wide, longer than the stem; inflorescences straight, longer than the subtending leaf, rarely as long, flowers opening several at a time in response to some environmental cue, light green-yellow; dorsal sepal 2.5–2.7 mm long. Rain forests, 200–400 m; Bolívar (Caño Pablo, north of Ichún, upper Río Caura, upper Río Paragua, Santa María de Erebato). Guyana, West Indies, Trinidad-Tobago. ◆Fig. 512. This species is at the center of a dispute on the typification of the genus Stelis and was the first species described that is referable to the genus, although it was described in Epidendrum. Stelis pusilla H.B.K., Nov. Gen. Sp. Pl. (quarto ed.) 1: 361. 1816. Epiphyte; leaves 20–25 mm long, 5–6 mm wide, about as long as or longer than the stem; inflorescences 20–40 mm long, longer than subtending leaf, flowers pale green or yellow; dorsal sepal ca. 2 mm long. Cloud forests, 1200–1300 m; Bolívar (20 km east of San José de Kayamá, Sierra de Maigualida). Mérida, Táchira, Trujillo; Colombia, Ecuador, Peru, Bolivia. Stelis pygmaea Cogn. in Urb., Symb. Antill. 6: 390. 1910. Stelis trichorrachis auct. non Rchb. f. 1855: sensu Dunst. & Garay, Venez. Orchid. Ill. 4: 321. 1966. Epiphyte, usually growing on twigs or small branches; leaves 15–50 mm long, at maturity slightly longer than subtending stem; inflorescences shorter to a little longer than leaves; flowers pale green or greenish yellow, campanulate, often cleistogamous; free portion of dorsal sepal 0.8–1.2 mm long. Rain or low-elevation cloud forests, 300– 1000 m; Bolívar (La Escalera to Cerro Venamo region). Aragua, Falcón, Miranda; West Indies. ◆Fig. 511. Stelis santiagoensis Mansf., Notizbl. Bot. Gart. Berlin-Dahlem 10: 239. 1928. Epiphyte; leaves 60–120 × 6–9 mm, 2–3 times longer than its stem; inflorescences 40–70 mm long, shorter than the subtending leaves; flowers pale green; dorsal sepal 2–3 mm long. Rain forests or, rarely, low-elevation cloud forests, 200–800(–1200) m; Delta

Stelis 587

Amacuro (northeast of El Palmar), Bolívar (Cerro Guaiquinima, La Escalera to Cerro Venamo region), Amazonas (Río Pasimoni, Río Putaco). Aragua, Carabobo, Yaracuy; Amazon basin in Colombia, Ecuador, Peru, and Brazil. Stelis schomburgkii Fawc. & Rendle, J. Bot. 48: 108. 1910. Stelis aprica auct. non Lindl. 1836: sensu Dunst. & Garay, Orchids Venez. Ill. Field Guide 955. 1979. Epiphyte or lithophyte; leaves 50–90 mm long, 5–10 mm wide, shorter than the stems; inflorescences 70–120 mm long, a little longer than subtending leaves; flowers pale yellow or green; dorsal sepal 1–1.5 mm long. Cloud forests or tepui shrublands, 1800– 2700 m; Bolívar (Auyán-tepui, Macizo del Chimantá), Amazonas (Cerro Marahuaka, Sierra de la Neblina). Guyana, adjacent Brazil. Stelis tolimensis Schltr., Repert. Spec. Nov. Regni Veg. Beih. 27: 157. 1924. Epiphyte, often growing among mosses; leaves 15–20 mm long, 7–10 mm wide, slightly longer than the stem; inflorescences heavily zigzag, creeping, ascendent, or pendent, as long or longer than the subtending leaf; flowers successive, dull olive-green, dull purple to dark purple; dorsal sepal 1.5–2 mm long. Locally common in rain or cloud forests, 100–1400 m; Bolívar (La Escalera to Cerro Venamo region), Amazonas (Río Pasimoni). Colombia. The flora area material of Stelis tolimensis could prove to be a different species than that from Colombia. It is known from 2 populations in the Guayana area, one in cloud forests in the La Escalera to Cerro Venamo region, the other from lowland rain forests in southern Amazonas. The population from Río Pasimoni seems to include occasional plants with larger flowers on straight rachises, but otherwise it seems conspecific with the Bolívar plants. Stelis tridentata Lindl., Fol. Orch. Stelis 6. 1859. Epiphyte, creeping or ascending; leaves 2.5–5 × 1.3–2.4 cm, petiolate, the petioles 0.8–1.9 cm long; inflorescences shorter than to as long as the leaves, laxly-flowered; flowers yellow, sepals 1.6–3 mm long; apparently

rare in rain forests, no exact elevation known but likely from below 500 m; Amazonas (upper Río Orinoco). Costa Rica(?), Colombia, Ecuador. Stelis tridentata is known from a single collection in the flora area with “Upper Orinoco” given for the locality. The Venezuelan material does not seem to match the type material very closely and might belong to a different species. Stelis zonata Rchb. f., Gard. Chron. n.s. 20: 556. 1883. Stelis grossilabris auct. non Rchb. f. 1881: sensu Foldats in Lasser, Fl. Venez 15(2): 99. 1970; Dunst. & Garay, Venez. Orchid. Ill. 3: 303. 1965. Epiphyte, rarely a lithophyte; leaves 3.5– 11 cm long, 8–22 mm wide, longer or about as long as the stems; inflorescences straight, longer than the subtending leaf; flowers opening several at a time in response to an environmental cue, dull purple with a green margin or totally dull purple; dorsal sepal 4.5–5.5 cm long. Rain or cloud forests, 500– 1800 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region). Aragua, Mérida. ◆Fig. 513.

Fig. 509. Stelis effusa

588

O RCHIDACEAE

Fig. 510. Stelis latisepala

Fig. 511. Stelis pygmaea

1 cm

1 mm

2 cm

Fig. 512. Stelis ophioglossoides

Fig. 513. Stelis zonata

Stellilabium 589

140. STELLILABIUM Schltr., Orchideen 530. 1914. [Subtribe Telipogoninae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Twig epiphytes, inconspicuous, cespitose, usually sun-loving. Roots relatively thick, sometimes photosynthetic. Rhizome very abbreviate. Stem very abbreviate, completely enveloped by the leaf sheaths. Leaves conduplicate, articulate, sometimes almost absent during anthesis, imbricate; blades linear to oblong-elliptic. Inflorescences usually terminal, less often lateral, axillary from the leaf sheaths, solitary or geminate, racemose or rarely branched, erect or spreading, long-pedunculate; peduncle usually flattened, remotely several-sheathed, sheaths cup-like, fleshy; rachis spiral or subdistichous, successively flowered, often ± zigzag, flattened, sometimes ± congested; floral bracts similar to the sheaths of the peduncle but smaller; pedicellate ovary ± equal to or longer than the floral bracts. Flowers resupinate or not, fleshy, inconspicuous, short-lived, often cleistogamous; perianth segments widely spreading, greenish or yellowish, sometimes with dark maroon spots, bands or blotches, or totally dark purple to maroon; sepals free, subequal, the laterals ± oblique; petals subequal to the sepals but narrower and often ± attenuated basally. Lip firmly attached to column base, larger than sepals, spreading, ecallose, 3-lobed at the very base; lateral lobes much smaller than central lobe, linear-oblong to linear-ovate, falcate, flat or erect; central lobe oblong-elliptic, acute or obtuse, margin often finely retrorse-ciliate. Column short, thick, erect, sometimes basally flanked by apically furcate hairs; anther dorsal, erect, operculate, imperfectly 2 or 4locular; pollinia 4, in 2 superposed, subequal pairs, dorsoventrally compressed, waxy; tegula elongate, apically dilated and 2-parted, viscidium elongate, hooked; clinandrium shallow; rostellum linear, elongate, erect; stigma entire, subapical. Capsules ellipsoid. Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, Bolivia, most diverse in the Andean areas; ca. 10 species, 2 or 3 in Venezuela, 1 of these in the flora area. In those species of Stellilabium that are ± leafless at anthesis, the flattened peduncle and the green, thick roots assume the photosynthetic role. The hairs or bristles on the column have been reported to attract tachinid flies that pseudocopulate with the flowers, effecting pollination. Stellilabium alticolum Dodson & R. Escobar, Orquideología 20(1): 48.1998. Twig epiphyte, inconspicuous, sun-loving; leaves 5–50 mm long; inflorescences 20–80 mm long; flowers greenish with sepals 2.5–4 mm long. Cloud forests, usually on small trees, 1200–1400 m; Bolívar (La Escalera to Cerro Venamo region). Aragua, Miranda, Portuguesa, Yaracuy; Andean Colombia and Ecuador. ◆Fig. 514.

Fig. 514. Stellilabium alticolum

590

O RCHIDACEAE

141. STENIA Lindl., Edwards’s Bot. Reg. 23: sub t. 1991. 1837. [Subtribe Zygopetalinae]. Stenopolon Raf., Fl. Tellur. 4: 49. 1836 [1838]. by Gustavo A. Romero-González Cespitose epiphytes. Stem abbreviate, not modified into pseudobulbs. Leaves 1–3, distichous, coriaceous, oblong to oblong-obovate, acute, plicate, articulate. Inflorescences lateral, arising from the axil of the leaf sheaths, pendent, 1-flowered, shorter than the leaves. Flowers resupinate or not, showy; floral bracts concave, appressed, ovate, acute, shorter than the pedicellate ovary. Sepals and petals similar, membranous, spreading, narrowly ovate; lateral sepals adnate to the column foot, sometimes slightly oblique. Lip sessile, adnate to the apex of the column foot, spreading, fleshy, helmet-shaped, deeply concave, triangular-ovate, ± 3-lobed; disk with a conspicuous, thickened, antrorse, transverse callus. Column short, slightly arched, trigonous, ventrally and basally pubescent, produced into a short foot; anther terminal; pollinia 4, yellow, cartilaginous, dorsoventrally flattened, linear, attached in 2 pairs to a short, subquadrate tegula, each pair with 1 pollinium much longer than the other. Colombia, Venezuela, Trinidad-Tobago, Guyana, Ecuador, Peru, Brazil, Bolivia; 9 species, 1 in Venezuela. Stenia is closely related to Chaubardiella Garay but is distinguished by the lip not articulate to the column. Stenia pallida Lindl., Edwards’s Bot. Reg. 23: sub t. 1991. 1837. Epiphyte; flowers large, white to pale yellowish green; lip with a toothed callus. Wet forests, 100–500 m; Delta Amacuro (Serranía de Imataca), Bolívar (Río Paragua). Coastal Cordillera of Distrito Federal and Miranda, Sucre (Cerro Patao); West Indies, Colombia, Trinidad-Tobago, Guyana, Ecuador, Peru, Brazil (Amazonas). ◆Fig. 515.

Fig. 515. Stenia pallida

142. STICTOPHYLLORCHIS Carnevali & Dodson, Lindleyana 8: 101. 1993. —Stictophyllum Dodson & M.W. Chase, Icon. Pl. Trop. ser. 2: t. 584. 1989, non Edgew. 1845. [Subtribe Oncidiinae]. Quekettia Lindl., Edwards’s Bot. Reg. 25: misc. 3. 1839, pro parte. by Germán Carnevali and Ivón M. Ramírez-Morillo Twig or branch epiphytes, cespitose, erect to pendulous. Pseudobulbs orbicular, flattened, apically 1-foliate, clothed by imbricating sheaths, the upper 2 or 3 with foliar blades, lower sheaths scarious. Leaves laterally compressed, articulate, straight

Teuscheria 591

or oblique, spotted with red or purple. Inflorescences lateral from pseudobulb base, thin, erect, an amply compound panicle ± equal or longer than the leaves, several flowers simultaneously open per branch. Flowers nonresupinate, campanulate. Dorsal sepal free, oblong-lanceolate, laterals fused to above middle; petals ovate, broader than sepals. Lip free, ecallose, ± lobed, the lateral lobes triangular, the central lobe acute. Column terete, foot well developed, each side of the stigma with a truncate wing; anther terminal, operculate; pollinia 2, obpear-shaped, tegula very small, viscidium elliptic; stigma ventral. Venezuela, Trinidad-Tobago, Ecuador, Amazonian Brazil; 1 species. Stictophyllorchis pygmaea (Cogn.) Carnevali & Dodson, Lindleyana 8: 101. 1993. —Ionopsis pygmaea Cogn. in Urb., Symb. Antill. 6: 624. 1910. —Trizeuxis pygmaea (Cogn.) Schltr. in Urb., Symb. Antill. 7: 498. 1913. —Quekettia pygmaea (Cogn.) Garay & R.E. Schult., Rhodora 59: 288. 1957. —Stictophyllum pygmaeum (Cogn.) Dodson & M.W. Chase, Icon. Pl. Trop. ser 2: 584. 1990. Plant to 6 cm, including the inflorescence; perianth segments white or pale purple; lip pale yellow; dorsal sepal 2 × 0.6 mm. Rain forests, often high in trees or lower when close to water courses, 50–500 m; Bolívar (basin of the Río Uaiparú, Río Cuyuní and Río Parguaza), northern Amazonas. Monagas; distribution as in genus. ◆Fig. 516.

Fig. 516. Stictophyllorchis pygmaea

143. TEUSCHERIA Garay, Amer. Orchid Soc. Bull. 27: 820. 1958. [Subtribe Lycastinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, inconspicuous, cespitose to creeping. Rhizome short to elongate, partially or totally clothed by imbricate, papery sheaths, eventually evanescent. Pseudobulbs aggregate or distanced on the rhizome, relatively small, 1-foliate, ovoid to pear-shaped, clothed by several evanescent sheaths, the largest of which surrounds the petiole of the leaf. Leaves plicate, narrow-oblong to linear-elliptic, acute to subacuminate, long-petiolate, not articulate. Inflorescences lateral, originating from pseudobulb base, 1-flowered, erect to pendulous, long to short pedunculate; peduncle clothed with 1–several distanced sheaths; floral bracts tubular; pedicellate ovary terete. Flowers relatively showy, resupinate or not, campanulate or with widely spreading perianth segments. Sepals and petals fleshy; sepals elliptic to oblong-elliptic, obtuse to subacute; dorsal sepal free; lateral sepals free or shortly connate, forming with the column foot a variously developed spur-like mentum; petals similar to dorsal sepal but narrower. Lip hinged to apex of column foot, 3-lobed or simple, basal portion included in the mentum, ending in a retrorse, spur-like appendage, apical portion surrounds column. Column erect, basally produced into a prominent foot, wingless; anther apical, operculate, incumbent; pollinia 4, compressed, superposed, cartilaginous, a small tegula present, viscidium small; rostellum transverse; stigma ventral, entire. Capsule ellipsoid.

592

O RCHIDACEAE

Southern Mexico, Central America, Colombia, Venezuela, Ecuador; 6 species, 1 in Venezuela. Teuscheria wageneri (Rchb. f.) Garay, Bot. Mus. Leafl. 21: 256. 1967. —Bifrenaria wageneri Rchb. f., Bonplandia (Hanover) 2: 17. 1854. Teuscheria venezuelana Garay, Rhodora 61: 39, fig. p. 40. 1959. Epiphyte; inflorescences pendulous; flowers not resupinate, opening widely; petals and sepals bronze with light maroon tinged; lip white flushed with pink toward margins and inner central lobe, the callus with golden-colored farinose material. Cloud forests, 1200–1400 m; Bolívar (La Escalera to Cerro Venamo region). Aragua, Distrito Federal. ◆Fig. 517.

Fig. 517. Teuscheria wageneri

144. TRICHOCENTRUM Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 11, pl. 115. 1836. [Subtribe Oncidiinae]. Acoidium Lindl., Edwards's Bot. Reg. 23: sub. t. 1951. 1837. Lophiaris Raf., Fl. Tellur. 4: 40. 1836 [1838]. Oncidium sect. Cebolletae Lindl., Bot. Reg. 28: sub t. 4. 1842. Cohniella Pfitzer in Engl. & Prantl, Nat. Pflanzenfam. 2(6): 194. 1889. Stilifolium Koniger & Pongr., Arcula 7: 186. 1997. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, small to medium-small, cespitose or very shortly creeping. Rhizome short, almost absent. Stems thickened into small, oblong, ellipsoid or ovoid pseudobulbs (or the pseudobulbs larger, to 4–6 cm in three Central American species here referred to as the Trichocentrum splendidum-complex), apically 1-foliate, clothed by 1 or 2 non-leaf-bearing sheaths. Leaves conduplicate to terete (in the Cohniella complex), usually very fleshy, concolorous, often variegated, more rarely totally purplish or reddish, linear to narrowly fusiform when terete, when flat elliptic, oblong, or narrowly obovate, articulate at the apex of the pseudobulbs, erect, arched or pendent. Inflorescences lateral, originating from pseudobulb base, 1-flowered to racemose or paniculate, few- to multiflowered, very short to much surpassing the leaves. Flowers medium-small to relatively large and showy, resupinate, short- to long-lasting, all opening ± simultaneously or few open at a time and successive (in a few species of the Lophiaris complex and in most taxa of the Trichocentrum sensu stricto complex), widely opening to campanulate, often fragrant (most common in the Trichocentrum sensu stricto complex), lip yellow, pink, brown, or white, petals and sepals usually the same base color but spotted or barred in brown, red, or purple, rarely totally white, cream, or pink; sepals subsimilar, free; petals usually similar to the sepals; lip usually larger and fleshier than other perianth segments, adnate to the base of the column, erect to suberect, simple, pandurate to shallowly or sharply 3-lobed, when 3-lobed, the apical, central lobe often much larger and showier than the laterals, frequently transverse; the lip blade concave, flat, or convex, basally pro-

Trichocentrum 593

duced into a variously developed spur (in the Trichocentrum sensu stricto complex) or the spur totally lacking; disk basally callose or with longitudinal keels or lamellae; column erect, relatively short and thick, apically with a pair of variously developed wings or auricles, footless; anther terminal, operculate, incumbent, sometimes pubescent or papillose (often in the Trichocentrum sensu stricto complex); pollinia 2, cartilaginous, notched to complanate, tegula broad, ± triangular, relatively short, viscidium small; rostellum short; stigma ventral. Capsule ellipsoid or 3-edged. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 20–25 species, ca. 10 in Venezuela, 5 of these in the flora area. The genus Trichocentrum is here taken to encompass several groups that share reduced pseudobulbs topped by a single, fleshy, flat or terete leaf. The sheaths that envelope the pseudobulbs lack foliar blades (or, if present, these are teratological). One clade within the genus (Trichocentrum sensu stricto) features well-developed spurs derived from the base of the lip while the other have spurless lips. Another complex features terete leaves: the former sect. Cebolleteae of Oncidium (or the genus Cohniella), commonly called the “rat-tail orchids.” A third complex displays relatively large, elliptic or oblong-elliptic, flat to subconduplicate, fleshy leaves (thus popularly called the “mule-ear” orchids) on top of reduced pseudobulbs, and racemose or paniculate inflorescences (the Lophiaris complex). Still another group of taxa, all from montane environments in Mexico and Guatemala, feature larger pseudobulbs and much fleshier but rigid leaves. Flowers within the genus are variable in size, shape, and color and while some apparently mimic Malpighiaceous flowers, still other resemble the flowers of the high-elevation Andean genus Cyrtochilum Kunth, with a relatively small, reflexed lip. All members of the genus display lower chromosome numbers (2n = 21, 22) than most members of the Oncidiinae, and certainly lower than true members of Oncidium. Key to the Species of Trichocentrum 1. 1.

2(1). 2.

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

4.

Plants with terete leaves; lip bright yellow .................................. T. cebolleta Plants with mule-ear leaves (flat, conduplicate, fleshy, single on top of pseudobulbs); lip cream, whitish, or pink with darker pink or purple spots ........................................................................................................ 2 Inflorescences successively 1–4-flowered; lip white, basally produced into a conspicuous spur; leaves spotless ............................................. T. fuscum Inflorescences usually simultaneously few- to multiflowered; lip usually yellowish, pink- or purple-spotted, not produced basally into a spur; leaves variously spotted with purple or reddish ................................... 3 Flowers large; lip 25–36 mm long ............................................. T. lanceanum Flowers smaller; lip to 12 mm long ........................................................... 4 Mature plants relatively small; leaves 5–15(–23) cm long; central lobe of lip at least twice as broad as basal blade; plants of rain forests ....................................................................................................... T. nanum Mature plants larger; leaves usually 20 cm or longer (to 60 cm long); central lobe of lip ± equal to width of the basal lobe, when rarely broader, not > 1.3 times broader; plants of dry or deciduous forests, very rarely from rain forests ............................................................... T. carthagenense

594

O RCHIDACEAE

Trichocentrum carthagenense (Jacq.) M.W. Chase & N.H. Williams, Lindleyana 16(2): 137. 2001. —Epidendrum carthagenense Sw., Enum. Syst. Pl. 30. 1760. —Oncidium carthagenense (Jacq.) Sw., Kongl. Vetensk. Acad. Nya Handl. 21: 240. 1800. —Lophiaris carthagenensis (Jacq.) Braem, Schlechteriana 4: 17. 1993. Usually epiphyte, sometimes muscicolous, sun- or shade-loving; leaves (9–)20– 60 cm long; flowers whitish, greenish, or pinkish, mottled or variegated with pink, purple, or maroon, lip 8–16 × 7–14 mm. Rare in deciduous forests (rarer in rain forests), ca. 50–200 m; Amazonas (Isla La Hormiga). Widespread in northern Venezuela in areas with a pronounced dry season, rarely in rain forests; Colombia, possibly Panama and Costa Rica. ◆Fig. 520. The name Oncidium carthagenense has been used to refer to several valid entities ranging from southern Mexico into northern South America. Quite a few of these are still undescribed. Here the name Trichocentrum carthagenense is restricted to populations of this complex with a prominent, tumor-like callus on a lip with the basal lobe about as broad as the width between the basal lobes while the column wings are dolabrate. Trichocentrum cebolleta (Jacq.) M. W. Chase & N. H. Williams, Lindleyana 16(2): 137. 2001. —Dendrobium cebolleta Jacq., Enum. Syst. Pl. 30. 1760. —Oncidium cebolleta (Jacq.) Sw., Kongl. Vetensk. Acad. Nya Handl. 21: 240. 1800. —Cohniella cebolleta (Jacq.) Christenson, Lindleyana 14(4): 177. 1999. Epiphyte, very rarely on rocks or muscicolous on the soil, sun-loving; leaves (7–)20–70 cm long, sometimes purplish when exposed; flowers bright yellow with brown or maroon blotches on sepals, petals, and callus; lip 7–25 × 7–27 mm, bright yellow. Dry or deciduous forests, near sea level to 300 m; Delta Amacuro (Isla Curiapo, Río Grande), Bolívar (Altiplanicie de Nuria, Cerro Cuchivero, Danto Manchado, near La Paragua, lower Río Orinoco, Río Toro), Amazonas (granitic outcrops near Puerto Ayacucho). Widespread in northern Venezuela in areas with a

pronounced dry season, more common in the Coastal Range; apparently widespread in the northern Neotropics. A member of the rat-tail orchids (Cohniella sensu stricto), this species is rare in the flora area but common elsewhere in Venezuela. Several entities may be encompassed within the broad concept of Trichocentrum cebolleta adopted here. Oncidium sprucei, described from Amazonian Brazil, is commonly referred to as a synonym of T. cebolleta. This taxon has relatively similar flowers, but leaves are thinner and longer, the isthmus of the lip is triangular, and the callus is differently shaped, and is here treated as a distinct species. It may eventually be found in Amazonas State. Some populations from eastern Venezuela (Sucre and Monagas states) have larger flowers than usual and require further study. Trichocentrum fuscum Lindl., Edwards's Bot. Reg. 23: sub. t. 1951. 1837. —Acoidium fuscum Lindl., Edwards's Bot. Reg. 23: sub. t. 1951. 1837, nom. nud. Trichocentrum cornucopiae Linden & Rchb. f., Gard. Chron. 1866: 266. 1866. Trichocentrum hartii Rolfe; Bull. Misc. Inform. Kew 1894: 395. 1894. Epiphyte, cespitose, fleshy-leaved, shadeloving; leaves often flushed with purple; flowers medium-sized, subcampanulate, sepals and petals greenish brown flushed with pink; lip white with yellow in middle and dark red in basal callus. Gallery forests in areas of strong dry season and in rain forests, 50–700 m; Bolívar (Altiplanicie de Nuria, Río Cuyuní basin, near Salto Angel, near Tumeremo). Monagas; Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil, Bolivia. ◆Fig. 518. The related Trichocentrum albo-coccineum Linden (= T. orthoplectrum Rchb. f.) has been reported in the literature as native from Venezuela and the type of the species was collected in Amazonas (epiphytic in forests along Río Negro). The species, however, has never been collected again. Since T. albococcineum has been collected in northern Brazil and Amazonian Colombia close to the Venezuelan border, and seems to be widespread in the Amazon Basin, it probably will be found eventually in the Venezuelan

Trichocentrum 595

Fig. 518. Trichocentrum fuscum

Fig. 519. Trichocentrum nanum

Fig. 520. Trichocentrum carthagenense

596

O RCHIDACEAE

Guayana. Trichocentrum albo-coccineum would be easy to recognize by its lanceolate to strictly triangular column wings. These structures are obtuse to truncate at apex. Trichocentrum purpureum Lindl. ex Rchb. f., known only from its Guyanan type, has an obovate-oblong lip and may also eventually be collected in the flora area. Trichocentrum lanceanum (Lindl.) M.W. Chase & N.H. Williams, Lindleyana 16(2): 137. 2001. —Oncidium lanceanum Lindl., Trans. Hort. Soc. London. 2: 100. 1836. —Lophiaris lanceana (Lindl.) Braem, Schlechteriana 1: 17. 1993. Lophiaris fragrans Raf., Fl. Tellur. 4: 41. 1836 [1838]. Epiphyte, erect or rarely pendulous; leaves 12–50 × 4–12 cm, spotted with purple or maroon; flowers on relatively short and stout inflorescences, featuring widely spreading perianth segments, sepals and petals yellowish, densely covered with large red-brown spots; lip pale purple, rarely almost pink or whitish, the callus and isthmus darker purple, 25–36 × 18–25 mm. Rain forests, very high on trees, sometimes close to rivers or streams, near sea level to 200(–300); Delta Amacuro (Caño Simoina, Río Acure, Río Cuyubini), Bolívar (near Icabarú, basin of lower Río Caura, Río Cuyuní), Amazonas (Río Cataniapo). Monagas, Sucre; Trinidad-

Tobago, Guyana, Suriname, French Guiana, northern Brazil. This spectacular species is much sought after by horticulturists, though not really easy to grow. It is widespread in nature and occurs in many inaccessible localities, but it has been almost stripped out close to roads and human dwellings. Fortunately, it has been propagated from seeds and it is readily available in the trade for orchid fanciers. Trichocentrum lanceanum is the type species of the genus Lophiaris (L. fragrans Raf.). Trichocentrum nanum (Lindl.) M.W. Chase & N.H. Williams, Lindleyana 16(2): 138. 2001. —Lophiaris nana (Lindl.) Braem, Schlechteriana 1: 19. 1993. —Oncidium nanum Lindl., Bot. Reg. 29: misc. 37. 1842. Epiphyte, erect or rarely pendulous; leaves 7–23 cm long; sepals and petals yellow or dull orange with purple, brown, or maroon spots; lip dull yellow with brownish on the basal 1/2, 6–12 × 6–12 mm. Rare or locally common in rain forests, 100–200 (–400) m; Bolívar (Cerro Guaiquinima, Río Icabarú), Amazonas (Río Cataniapo, upper Río Orinoco, Tamatama). Yaracuy; Amazonian Brazil, Guyana, Ecuador, Peru, Bolivia. ◆Fig. 519. This interesting species has been poorly collected.

145. TRICHOPILIA Lindl., Intr. Nat. Syst. Bot. ed. 2, 446. 1836. [Subtribe Oncidiinae]. Pilumna Lindl., Edwards’s Bot. Reg. 30: misc. 73. 1844. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes or rarely lithophytes, inconspicuous to large and showy, cespitose. Pseudobulbs heteroblastic (one internode), apically 1-leaved, usually oblong but frequently ellipsoid or ovoid, usually strongly flattened and ancipital (flattened and with sharp edges), partially covered with nonleaf-bearing sheaths. Leaves conduplicate, erect, subcoriaceous to rarely somewhat fleshy, sessile to short pseudopetiolate, usually oblong or elliptic. Inflorescences lateral, originating from pseudobulbs base, usually pendulous, sometimes erect to arching, 1-flowered; peduncle terete to somewhat flattened, remotely few-bracted. Flowers medium-small to very large and showy, resupinate, opening well, often with submembranous segments and sparkling texture, short to rather long-lived; floral bracts usually conspicuous. Sepals narrow, often twisted, mostly similar and free, or rarely the lateral ones somewhat basally connate; petals similar to dorsal sepal. Lip usually larger and wider than other perianth segments, simple, 3- or 4-lobed, clawed, connate to

Trichopilia 597

connivent to column base or basal 1/2, basal lobes or margins convolute, forming a tube which surrounds the column; disk ecallose, carinate or rarely callose; column semiterete, elongate, erect; anther terminal, operculate, clinandrium prominent with smooth to lacerate-fimbriate margins, pollinia 2, hard, stigma ventral, rostellum short. Capsules ellipsoid. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia; ca. 25 species; 4 or 5 in Venezuela, 2 of these in the flora area. Leucohyle is closely related to Trichopilia, differing only in the narrower, fleshier leaves. Some authors treat both Leucohyle and Helcia Lindl. as congeneric with Trichopilia. Key to the Species of Trichopilia 1.

1.

Inflorescences 1–4-flowered; margins of petals and sepals not undulate, smooth; lip 3–4 cm long, 2 cm wide, basal margins parallel to column, rounded or truncate at apex; disk shallowly 3-carinate on throat .................................................................................................... T. aenigma Inflorescences 1- or 2-flowered; margins of petals and sepals undulate; lip 4–5 cm long, 3.3–4 cm wide, basally funnel-form and completely enfolding the column, deeply emarginate at apex; disk thickly 3-carinate on throat . ..................................................................................... T. oicophylax

Trichopilia aenigma Garay, Lindleyana 11: 233. 1996. Trichopilia albida auct. non H. Wendl. 1851: sensu Dunst. & Garay, Venez. Orchid. Ill. 3: 316. 1965; Dunst. & Garay, Orchids Venez. Ill. Field Guide 1023. 1979. Trichopilia fragrans auct. non (Lindl.) Reich. f. 1858: sensu Foldats in Lasser, Fl. Venez. 15(5): 111. 1970. Epiphyte 30–40 cm tall; perianth segments greenish, sepals 40 mm long, 7 mm wide; lip white with yellow throat, apical part with erose-undulate margin. Rain or cloud forests, ca. 1300 m; Bolívar (Auyántepui, northeastern Luepa, Río Cuyuní, Santa Elena de Uairén). Aragua, Distrito Federal, Mérida; Colombia. Trichopilia oicophylax Rchb. f., Allg. Gartenzeitung 24: 97. 1856. Epiphyte; perianth segments greenish, translucent, lip white, basally and internally yellow, sepals 40–50 mm long, 7 mm wide. Rain or cloud forests, ca. 1300 m; Bolívar (La Escalera to Cerro Venamo region). Aragua, Distrito Federal, Lara. ◆Fig. 521.

Fig. 521. Trichopilia oicophylax

598

O RCHIDACEAE

146. TRICHOSALPINX Luer, Phytologia 54: 393. 1983. [Subtribe Pleurothallidinae]. Pleurothallis sect. Brachystachyae subsect. Lepanthiformes Lindl., Fol. Orchid., Pleurothallis 25. 1859. —Pleurothallis sect. Lepanthiformes (Lindl.) Cogn. in Mart., Fl. Bras. 3(4): 591. 1896. Pleurothallis sect. Bipaleolatae Pabst, Orch. Brasilienses 162. 1975. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, lithophytes, less often muscicolous herbs, mostly inconspicuous, usually cespitose but sometimes long-creeping, erect to pendulous, sun- or shadeloving. Rhizome abbreviate to long and creeping; ramicauls ascending to erect or descending, terete, much shorter to several times longer than the leaves, thin and wiry to relatively stout, nonpseudobulbous, 1-leaved at apex, sometimes apically prolific (producing another ramicaul from the apex) or superposed; enveloped by tubular to funnelform, ribbed, ± imbricate sheaths (lepanthiform sheaths) with oblique, dilated openings (ostia), the ribs and margins of the ostia grossly or microscopically ciliate to scabrous, rarely glabrous. Leaves erect or suberect in relation to the ramicaul, conduplicate, articulate, submembranous to coriaceous or fleshy-coriaceous, linear-elliptic to orbicular, usually flat, sometimes concave or convex, basally sessile to subpetiolate, often purple- or maroon-tinged, the apex acute to obtuse, notched with an apiculum in the sinus. Inflorescences originating laterally with an annulus at or near the apex of ramicaul, or abscission layer, 1-flowered (rarely) to racemose, lax to dense, much shorter to several times longer than subtending leaves, erect to pendulous; distichous or secund, simultaneously to successively few- to many-flowered, peduncle remotely sheathed; floral bracts usually inconspicuous; pedicel short and ± stout to filiform and much longer than ovary; ovary terete to, more often, obclavate, costate or usually smooth. Flowers usually inconspicuous, sometimes large for plant size, usually resupinate, widely opening to campanulate, sometimes cleistogamous, usually petals parallel to column. Perianth segments hyaline, white, yellowish, or variously streaked on red or purple, to completely red or dark purple, usually membranous or fleshy; sepals ± ovate, with or without tails, carinate or subcarinate, glabrous or pubescent, often ciliate, subsimilar and free, or the laterals variously connate into a flat, convex to concave synsepal; petals free, small, usually smaller than sepals, membranous, entire, denticulate or ciliate. Lip hinged to column foot or base, simple or 3-lobed, basally often with 2 small, retrorse lobes; disk callose or ecallose, usually keeled; margins entire to denticulate or fimbriate to ciliate. Column short to long, erect, straight to somewhat arched, usually with a well-developed foot, sometimes footless, apically often winged or cucullate; anther apical, subapical or ventral, operculate, incumbent, 1locular; pollinia 2, free, subspherical to obovoid, with a granular caudicle; clinandrium entire, lobed, fimbriate to ciliate; rostellum, ventral to subapical; stigma ventral to subapical. Capsules variable. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 150 species, 20 species in Venezuela, 11 of these in the flora area. Trichosalpinx includes 4 subgenera, 2 of these present in the flora area. Subgenus Trichosalpinx includes plants with petals not entire and the lip with basal lobes, and the inflorescences are normally shorter than the subtending leaf; subgenus Tubella includes plants with petals entire and the lip without basal lobes, and inflorescences that are usually longer than the leaves.

Trichosalpinx 599

Key to the Species of Trichosalpinx

1.

1.

2(1).

2.

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

4.

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

6. 7(6).

7.

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

Inflorescences much shorter to rarely as long as the leaves; flowers purple, red, maroon, or wine; dorsal sepal obtuse to acute, never caudate-acuminate; lip simple to subpandurate; ramicauls not superposed or prolific ................................................................................................ 2 Inflorescence longer than leaves; flowers mostly white, cream, or yellowish, sometimes with red or purple streaks or dots; dorsal sepal ± caudate-acuminate; lip ± 3-lobed; plants small to minute; ramicauls often proliferous or superposed ...................................................................... 6 Lip longer to or as long as synsepal, narrow-oblong, 5–6 times as long as broad, its margin long ciliate; peduncle of the inflorescence surpassing 1/2 the length of the subtending leaf ............................................... T. egleri Lip much shorter than synsepal, oblong to obovate or narrow-triangular, 2–4 times as long as wide (when to 5 times as long as wide, then narrow-triangular); peduncle of the inflorescence 1/6–1/3 the length of subtending leaf ............................................................................................ 3 Lip narrow-triangular or oblong-ovate, acute at apex; inflorescences 2(4)-flowered .......................................................................................... 4 Lip oblong-pandurate, obtuse or rounded at apex; inflorescences usually with > 5 flowers ...................................................................................... 5 Lateral sepals connate for 1/4–1/2 of their length, spreading at the apical half; leaves usually suborbicular to broad-elliptic, sessile or inconspicuously pseudopetiolate ............................................................ T. orbicularis Lateral sepals connate for 3/4 of their length, subparallel to dorsal sepal; leaves narrow-elliptic to elliptic, shortly but conspicuously pseudopetiolate ..................................................................................... T. oxychilos Lip with a basal longitudinal callus extending up to the middle of its length; leaves 2–3 times longer than wide; clinandrium ciliate .............. T. memor Lip with a short, transversal or conic callus extending < 1/3 of its length; leaves 4–5 times longer than wide; clinandrium glabrous .......... T. patula Stems inconspicuous, much shorter than leaves; lepanthiform sheaths nearly devoid of cilia or scabrosities, then resembling those of a Pleurothallis ................................................................................. T. pumila Stems always conspicuous, longer than leaves: sheaths with well-developed cilia or scabrosities ........................................................................ 7 Leaves elliptic; ramicauls superposed sequentially, forming multifoliate stem-like units; lateral lobes of lip small, rounded or truncate; lip basally cuneate ...................................................................... T. cedralensis Leaves widely elliptic to suborbicular; ramicauls prolific or not, but not forming apparently multifoliate stems; lateral lobes of lip well developed or absent; falcate or retrorse; lip sessile or shortly clawed ................................................................................................................ 8 Lip subsimple or simple; leaves dorsally with 3 purple veins; petals 1/3 or less wider at dorsal sepal ................................................................. T. dura Lip sharply 3-lobed; leaves several-veined dorsally, when 3-veined, dorsally concolorous; petals at least 1/2 broad as dorsal sepal ................... 9 Petals much wider than lateral sepals; sepals long caudate-acuminate; lip bicallose at base of lateral lobes ................................................ T. intricata

600

O RCHIDACEAE

9.

Petals narrower to subequaling lateral sepals; sepals shortly acuminate; lip essentially ecallose ......................................................................... 10 10(9). Anterior part of the lip oblong, basal lobes on the basal 1/4–1/3 .................. ............................................................................................... T. roraimensis 10. Anterior part of the lip triangular, basal lobes filing the basal 1/2 of the lip .............................................................................................. T. steyermarkii Trichosalpinx cedralensis (Ames) Luer, Phytologia 54: 395. 1983. —Pleurothallis cedralensis Ames, Sched. Orchid. 4: 18. 1923. [Subgenus Tubella]. Epiphyte, erect or arching; leaves usually purple or black speckled, 0.8–2 cm long; inflorescences simultaneously 3–10-flowered, flowers inconspicuous, subcampanulate, relatively large for the plant; dorsal sepal 5.5–9.5 mm long, all perianth segments yellowish or whitish, with pink or brown blush. Locally common in cloud forests, (500–) 1000–1500(–1800) m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Jaua-Sarisariñama massif, La Escalera to Cerro Venamo region, Serranía Marutaní, Sierra Pakaraima), Amazonas (Cerro Asisa, Cerro Marahuaka). Costa Rica, Panama, Ecuador, Bolivia. Trichosalpinx dura (Lindl.) Luer, Phytologia 54: 395. 1983. —Pleurothallis dura Lindl., Fol. Orchid., Pleurothallis 32. 1859. —Humboldtia dura (Lindl.) Kuntze, Revis. Gen. Pl. 2: 667. 1891. [Subgenus Tubella]. Pleurothallis foliata Griseb., Fl. Brit. W. I. 610. 1864. —Humboldtia foliata (Griseb.) Kuntze, Revis. Gen. Pl. 2: 667. 1891. —Trichosalpinx foliata (Griseb.) Luer, Phytologia 54: 395. 1983. Pleurothallis corazonica F. Lehm. & Kraenzl., Bot. Jahrb. Syst. 26: 443. 1899. Pleurothallis broadwayi Ames, Orchidaceae 2: 267. 1908. Pleurothallis guadalupensis Cogn. in Urb., Symb. Antill. 6: 432. 1909. Pleurothallis amygdalina Rolfe ex Schltr., Repert. Spec. Nov. Regni Veg. 14: 386. 1916, nom. nud. Pleurothallis lepanthopsis Schltr., Repert. Spec. Nov. Regni Veg. 14: 386. 1916. Pleurothallis williamsii Ames, Orchidaceae 7: 120. 1922. Pleurothallis anomala Hoehne, Arch. Inst. Biol. (São Paulo) 2: 43. 1929. —Pleurothallis broadwayi subsp. anomala (Hoehne) Garay, Arch. Jard. Bot. Rio de Janeiro 11: 53. 1951.

Epiphyte, erect or scandent; leaves 0.6– 1.9 cm long, usually with 1–3 longitudinal purple stripes; inflorescences 1–6-flowered; flower inconspicuous, usually cleistogamous, subcampanulate; perianth yellow or yellowwhite; dorsal sepal 3–6 mm long; lip suffused with red, purple or maroon. Cloud forests, 1400–1800 m; Bolívar (Auyán-tepui, La Escalera to Cerro Venamo region), Amazonas (Cerro Aracamuni, Cerro Yutajé). Nueva Esparta; Mexico, Central America, West Indies, Ecuador, Peru. ◆Fig. 522. Trichosalpinx egleri (Pabst) Luer, Phytologia 54: 395. 1983. —Pleurothallis egleri Pabst, Ann. XIV Congr. Bot. Bras. 14. 1964. [Subgenus Trichosalpinx]. Epiphyte, erect; leaves 3.5–4 cm long, the lower surface purple-tinged; inflorescences straight, 1/2–3/4 the length of the subtending leaf; flowers inconspicuous with dorsal sepal 2.8–3.3 mm long; sepals basally cream, dark magenta purple at the upper half; synsepal ± spreading apically; petals, lip, and ventral face of column dark magenta-purple. Rain forests; 200–700 m; Delta Amacuro (Río Cuyubiní), Bolívar (Altiplanicie de Nuria, Serranía de Imataca). Grenada, Guyana, Suriname, French Guiana, Peru, Bolivia, Amazonian Brazil. ◆Fig. 523. Trichosalpinx intricata (Lindl.) Luer, Phytologia 54: 396. 1983. —Pleurothallis intricata Lindl., Orchid. Linden. 1. 1846. Epiphytes; ramicauls erect, sometimes arched; leaves 1–2(–3) cm long; inflorescences with zigzag (fractiflex) rachis; flowers inconspicuous but large for the plant, dorsal sepal 6–13 mm long, all perianth segments yellow or yellow-green. Rare in cloud forests, 1700–2000 m; Bolívar (Auyán-tepui). Aragua, Mérida, Sucre, Táchira, Trujillo; Colombia. Trichosalpinx memor (Rchb. f.) Luer, Selbyana 7: 47. 1982. —Pleurothallis memor Rchb. f., Bonplandia (Hanover) 4:

Trichosalpinx 601

330. 1856. —Humboldtia memor (Rchb. f.) Kuntze, Revis. Gen. Pl. 2: 667. 1891. [Subgenus Trichosalpinx]. Pleurothallis brevis Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19: 183. 1923. Pleurothallis gnomonifera Ames, Sched. Orchid. 6: 61. 1923. Pleurothallis minutipetala Ames & Schweinf., Sched. Orchid. 10: 32. 1930. —Trichosalpinx minutipetala (Ames & Schweinf.) Luer, Phytologia 54: 396. 1983. Trichosalpinx greenwoodiana Soto Arenas, Orquídea (Mexico) 10: 257. 1987. Trichosalpinx nageliana Soto Arenas, Orquídea (Mexico) 10: 259. 1987. Pleurothallis ciliaris auct. non (Lindl.) L.O. Williams 1842: sensu Foldats in Lasser, Fl. Venez. 15(3): 239. 1970; Dunst. & Garay, Venez. Orchid. Ill. 2: 271. 1961. Epiphyte, rarely a lithophyte, mostly erect but sometimes subpendulous; ramicauls straight to slightly zigzag; leaves 1.7– 4.5(–7) cm long, often purplish- or reddishtinged; flowers campanulate, inconspicuous with dorsal sepal 2.7–4 mm long, sepals translucent heavily streaked or tinged with dark wine purple, petals translucent with few dark purple longitudinal streaks; lip dark wine purple. Locally common in rain or cloud forests, (200–)700–1400(–2000) m; Bolívar (Auyán-tepui, Gran Sabana, JauaSarisariñama massif, La Escalera to Cerro Venamo region, Macizo del Chimantá [Amurí-tepui], Río Cuyuní), Amazonas (Cerro Marahuaka, near San Fernando de Atabapo, Sierra Parima). Aragua, Distrito. Federal, Mérida, Portuguesa, Sucre, Yaracuy; southern Mexico, Guatemala, Honduras, Costa Rica, Granada, Cuba, Panama, Colombia, Guyana, Suriname, Trinidad-Tobago, Ecuador, Peru, Brazil, Bolivia. Material of Trichosalpinx memor from high elevations (1900–2000 m) in Macizo del Chimantá has larger leaves (5–7 cm long) and flowers (dorsal sepal 4.5–5.5 mm long) and might represent a different variety or subspecies. Trichosalpinx orbicularis (Lindl.) Luer, Phytologia 54: 396. 1983. —Specklinia orbicularis Lindl., Edwards’s Bot. Reg. 24: misc. 31. 1838. —Pleurothallis orbicularis (Lindl.) Lindl., Edwards’s Bot.

Reg. 28: misc. 79. 1842. —Humboldtia orbicularis (H. Focke) Kuntze, Revis. Gen. Pl. 2: 668. 1891. [Subgenus Trichosalpinx]. Pleurothallis biflora H. Focke, Tijdschr. Wis- Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 2: 197. 1849. —Humboldtia biflora (H. Focke) Kuntze, Revis. Gen. Pl. 2: 667. 1891. Lepanthes orbiculata Lindl. ex Rchb. f., Xenia Orchid. 1: 152. 1856. Pleurothallis trachytheca F. Lehm. & Kraenzl., Bot. Jahrb. Syst. 26: 444. 1899. Pleurothallis lancifera Schltr., Repert. Spec. Nov. Regni Veg. Beih. 27: 48. 1924. Epiphyte, erect; leaves 2.2–4.5 cm long; inflorescences much shorter than subtending leaves; flowers inconspicuous with dorsal sepal 3–6 mm long; sepals spreading, maroonreddish or yellowish with purple or maroon longitudinal streaks, petals translucent yellowish or cream with few maroon streaks, lip dark maroon. Locally common in rain forests, 100–700(–1800); Bolívar (Altiplanicie de Nuria, Auyán-tepui, Cerro Guaiquinima, Cerro Jaua, Cerro Uroi, Gran Sabana, La Escalera, Serranía Imataca), Amazonas (Río Ararí, Río Cataniapo, Río Coro Coro, Río Siapa, near San Carlos de Río Negro, Sierra de la Neblina). Miranda, Sucre; Costa Rica, Panama, Guyana, French Guiana, Suriname, West Indies, Colombia, Ecuador, Peru, Brazil. Trichosalpinx orbicularis is a common, widespread, variable species, restricted to low elevations, it is remarkably consistent in most of its features in the Guayanan area and remains distinct from the higher elevation, parapatric T. oxychilos. Trichosalpinx oxychilos Carnevali & G.A. Romero, Novon 3: 21. 1993. Epiphyte, erect; leaves 2.5–3.5 cm long; flowers inconspicuous, bilabiate; dorsal sepal lanceolate, 4.3–4.8 mm long; sepals pale brown with red tinges; petals translucent white, apically lacerate; lip red with white basal lobules, apically and medially yellow. Cloud forests, ca. 1800 m; Bolívar (Cerro Jaua). Endemic. Trichosalpinx oxychilos was recently included in a superspecies circumscription of T. orbicularis (Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 64: 29. 1997). However, this entity from higher elevations is distinct

602

O RCHIDACEAE

Fig. 522. Trichosalpinx dura

Fig. 523. Trichosalpinx egleri

2 mm 5 mm

2 mm

2 mm 3 cm Fig. 524. Trichosalpinx pumila

enough from the lower-elevation members of the T. orbicularis complex in the Guayana region to warrant its continued recognition as a species. Trichosalpinx patula Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 65: 82. 1998. —Trichosalpinx patula Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 64: 31. 1997, nom. inval. [Subgenus Trichosalpinx]. Epiphyte, pendent; ramicauls 5–15 cm long; leaves 5–8 cm long, underside sometimes purplish; inflorescences shorter than leaves; flowers inconspicuous; dorsal sepal 5–6.5 mm long; synsepal spreading to

Fig. 525. Trichosalpinx roraimensis

subreflexed; sepal hyaline white at base grading to dark maroon-purple in the apical half; petals hyaline, white; lip dark maroonpurple. Rain forests, ca. 400 m; Bolívar (Cerro Paraucaipa near Auyán-tepui). Ecuador, Bolivia. Trichosalpinx pumila (Luer) Luer, Phytologia 54: 397. 1983. Pleurothallis pumila Luer, Selbyana 1: 258. 1975. [Subgenus Tubella]. Epiphyte, 2.5–3 cm tall, growing on twigs or thin branches; leaves 8–13 mm long; flowers inconspicuous, apparently always cleistogamous, translucent white or yellowish; dorsal sepal 2.5 mm long; lip ca. 2 mm long.

Trigonidium 603

Cloud forests, 1200–1300 m; Bolívar (La Escalera to Cerro Venamo region). Ecuador. ◆Fig. 524. This is the first collection of Trichosalpinx pumila outside eastern Ecuador, where it grows at comparable elevations in rain and cloud forests. The present report suggests a disjunction, but it is more likely that this little species has been overlooked by previous collectors. Trichosalpinx roraimensis (Rolfe) Luer, Phytologia 54: 397. 1983. —Pleurothallis roraimensis Rolfe, Trans. Linn. Soc. London. ser. 2, 4: 58. 1901. [Subg. Tubella]. Epiphyte, lithophyte, or musciculous herb, erect, usually sun-loving; leaves often purple- or reddish-tinged, 0.4–1 cm long; inflorescences erect; flowers often cleistogamous, inconspicuous with dorsal sepal 2.5–5 mm long; perianth segments creamy, yellowish or whitish. Cloud forests, (1200–)1600–

2800 m; widespread and common in Bolívar and Amazonas. Guyana, Brazil (Amazonas: Serra da Neblina, Roraima). ◆Fig. 525. Recently included in a broad circumscription of the Andean Trichosalpinx pusilla, the Guayanan material appears sufficiently distinctive, particularly in the shape of the lip, to be treated as a species of its own. Trichosalpinx steyermarkii Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 64: 79. 1997. [Subgenus Tubella]. Epiphyte 13–16 mm tall (smallest of the genus); ramicauls erect, 3–4 mm long, leaf 3– 4 × 2–3 mm; inflorescence 10–15 mm long; flowers yellow or white. Cloud forests, ca. 1800 m; Bolívar (Auyán-tepui). Endemic. Material referred to Trichosalpinx steyermarkii was not seen for this treatment. More collections are required to ascertain whether this entity is different from T. roraimensis or just a diminutive form of it.

147. TRIGONIDIUM Lindl., Edwards’s Bot. Reg. 23: t. 1923. 1837. [Subtribe Maxillariinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Mostly epiphytes, rarely lithophytes or subterrestrials, erect. Rhizome abbreviate to more commonly creeping to subscandent, appressed to substrate to ascendent, frequently branching, terete. Pseudobulbs pear-shaped to ellipsoid or oblong, in some species strongly ancipital (flattened and with sharp edges), commonly distanced on the rhizome, 1–4(5)-foliate apically, clothed by 1–several imbricate sheaths of which the innermost 1–3 in some species may have foliar blades. Leaves conduplicate, articulate, flat, coriaceous, subsessile or basally attenuate into a short pseudopetiole, blades linear to oblanceolate. Inflorescences originating from the base of the mature pseudobulb or the new growth, solitary or in fascicles, 1-flowered, long-pedunculate, erect; peduncle terete or somewhat compressed, clothed by several conspicuous, loose bracts; floral bracts similar to the peduncle bracts, ± equal to the pedicellate ovary; pedicellate ovary elongate, terete. Flowers erect or horizontally spreading, campanulate, long-lasting. Sepals membranous, free, elliptic or oblanceolate, acute or acuminate, often basally clawed, yellow, greenish or cream with purple or reddish veins, subequal or the dorsal sharply different, subparallel to the column, forming a cup that encloses the petals, lip and column, apices spreading; petals much smaller than the sepals, parallel to column, similarly colored but with a blue “eye” at the apex. Lip articulate to column foot, parallel to column, shorter than petals, 3-lobed in the apical 1/3, central lobe larger than the laterals, frequently callose-thickened, lateral lobes flat or erect and embracing the column; disk with a longitudinal callus. Column erect, shorter than lip, semiterete, straight, wingless, basally with a short foot or the foot almost absent; anther terminal, operculate, incumbent, very convex, 1-locular; pollinia 4, waxy, dorsoventrally compressed, 1 pair

604

O RCHIDACEAE

larger than the other, tegula transversely semilunate, the viscidium is the basal face of the tegula; clinandrium shallow, entire; rostellum transverse; stigma ventral. Capsules ellipsoid. Southern Mexico, Central America, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 12 species, 3 in Venezuela, 2 of these in the flora area.

Fig. 526. Trigonidium obtusum

Fig. 527. Trigonidium acuminatum

Triphora 605

Key to the Species of Trigonidium 1.

1.

Pseudobulbs pear-shaped, little or not flattened, 1-foliate; leaves 5– 13 mm wide; dorsal sepal 20–28 mm long; anther apically smooth ............................................................................................. T. acuminatum Pseudobulbs flattened, 2-foliate; leaves 2–6 cm wide; dorsal sepal 3– 4.2 cm long; anther apically glandular-pilose ........................... T. obtusum

Trigonidium acuminatum Bateman ex Lindl., Edwards’s Bot. Reg. 23: misc. 74. 1838. Trigonidium tenue Lodd. ex Lindl., Edwards’s Bot. Reg. 25: misc. 44. 1839. Epiphyte, rarely a lithophyte, mostly sunloving; leaves 9–27 cm long; petals and sepals greenish, yellowish, orange, or brownish, with brown or purple veins; lip with purple veins. Rain forests, 50–800(–1200) m; widespread in the flora area. Apure, Monagas, Zulia; Colombia, Suriname, Guyana, Ecuador, Peru, Brazil. ◆Fig. 527.

Trigonidium obtusum Lindl., Edwards’s Bot. Reg. 23: t. 1923. 1837. Epiphyte, rarely a lithophyte, mostly shade-loving; leaves 20–60 cm long; flowers greenish yellow to brown. Rain to rarely cloud forests, 100–800(–1200) m; Bolívar (throughout), Amazonas (basin of Río Casiquiare, Río Negro, Río Sipapo). Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 526.

148. TRIPHORA Nutt., Gen. N. Amer. Pl. 2: 192. 1818. [Tribe Triphoreae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrials, humicolous on the ground or rarely on low branches or rotting logs, sometimes saprophytes, 5–30 cm tall, shade-loving, delicate. Roots fleshy or with nodular tuberoids, basal or from the lower part of the erect stem. Rhizome abbreviate or lacking. Stems fleshy, terete, slender, erect or short-creeping at base, the basal part enveloped by tubular sheaths. Leaves not articulate, convolute, subdistichous or solitary at midstem to almost absent in near-saprophytes, sessile, remote in the apical part of the stem, sometimes originating tubers in the foliar axils; blades usually broad-ovate to broad-elliptic, often purplish. Inflorescences terminal, 1-flowered, or racemes of few spiral flowers; floral bracts similar to leaves so flowers can be interpreted as axillary from upper internodes. Flowers resupinate, fugacious, successive, ± showy; perianth segments membranous, pink, purple, or yellow, glabrous, narrowly lanceolate to narrowly oblanceolate, ± spreading; sepals free, subsimilar; dorsal sepal concave or cucullate; lateral sepals oblique; petals free, similar to dorsal sepal. Lip erect, articulate, usually 3-lobed, parallel to column, sessile, with a cuneate base, central lobe usually larger than laterals; lateral lobes erect, often embracing the column; disk ecallose or variously callose. Column erect, semiterete, apically thickened, subclavate, footless; anther terminal, erect, fleshy, firmly connate to the column apex; the thecae nearly as long as the anther; pollinia 4, powdery-granular, without viscidium; clinandrium with entire, dentate, or lobed margin; rostellum transverse, plate-like, convex, rigidly erect with forward-facing cells; stigma ventral, entire. Capsule erect or pendulous, ellipsoid. U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, Paraguay, northern Argentina; ca. 16 species, 3 in Venezuela, 1 of these in the flora area.

606

O RCHIDACEAE

Most of the species of Triphora live in small, localized colonies that, unlike most orchids, flower gregariously. Triphora surinamensis (Lindl. ex Benth.) Britton in Britton & P. Wilson, Bot. Porto Rico 1: 184. 1924. —Pogonia surinamensis Lindl., London J. Bot. 2: 674. 1843. Muscicolous herb 5–25 cm tall; sepals and petals pink to light purple, ca. 1.5 cm long, with 3 conspicuous veins; lip apically 3lobed, pink, ca. 1 cm long, with 3 longitudinal rows of papillose projections in middle; column winged. Rain forests, 600–1100 m; Bolívar (La Escalera to Cerro Venamo region), Amazonas (Cerro Duida, Cerro Huachamacari, basins of Río Cataniapo and Río Cuao). Mexico, Panama, West Indies, Colombia, Trinidad-Tobago, Guyana, Suriname, Brazil. ◆Fig. 528.

1 cm 1 cm

Fig. 528. Triphora surinamensis

4 cm

1 cm

149. TRISETELLA Luer, Phytologia 47: 57. 1980. —Masdevallia sect. Triaristellae Rchb. f., Gard. Chron. n.s. 6: 226. 1876. —Triaristella Brieger, Orchideen 449. 1976, nom. inval. —Triaristella Brieger ex Luer, Selbyana 2: 205. 1978, non Malyavk. 1949. —Triaristellina Rauschert, Repert. Spec. Nov. Regni. Veg. 94: 459. 1983, nom. illeg. [Subtribe Pleurothallidinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Small, cespitose, erect epiphytes. Stems short, subterete, clothed by scarious sheaths, apically 1-foliate. Leaves conduplicate, flat to subterete, erect, usually narrowly elliptic to narrowly obovate. Inflorescence originating from the apex of ramicaul with an annulus, erect, slender, long-pedunculate, peduncule sometimes scabrous to puberulent, usually surpassing the leaves, bearing a short successively few-flowered raceme; rachis straight to zigzag. Flowers small to medium-sized for the subtribe, large for the plant, resupinate. Dorsal sepal dorsally connate to the cup of the synsepal, variously adnate; lateral sepals connate into a synsepal concave to ± flat with tails originating below the apices of lateral sepals or synsepal, or sometimes subapically. Petals small, membranous, parallel to the column in natural position, usually oblong, apically truncate and variously dentate. Lip hinged to apex of column foot, simple to ± 3-lobed, longitudinally callose, the base cordate, cleft, with the basal lobes flanking the column, hinged to the column foot in the cleft. Column elongate, erect, clinandrium entire, hooded, basally produced into a laterally compressed column foot; anther ventral, incumbent, operculate. Pollinia 2, pear-shaped; stigma ventral, relatively large. Costa Rica, Panama, Colombia, Venezuela, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 20 species; 1 in Venezuela.

Trizeuxis 607

Trisetella is closely related and recently segregated from Masdevallia Ruiz & Pav. from which it differs by its laterally compressed column foot and ecallose petals. The only Guayanan species is easy to recognize among other pleurothallids of the area by the tails originating subapically from the synsepal. Trisetella triglochin (Rchb. f.) Luer, Phytologia 47: 58. 1980. —Masdevallia triglochin Rchb. f., Gard. Chron. n.s. 2: 648. 1877. —Triaristella triglochin (Rchb. f.) Luer, Selbyana 2: 206. 1978. —Triaristellina triglochin (Rchb. f.) Rauschert, Repert. Spec. Nov. Regni. Veg. 94: 459. 1983. Trisetella huebneri (Schltr.) Luer, Phytologia 47: 58. 1980. —Masdevallia huebneri Schltr., Beih. Bot. Centralb. 42(2): 88. 1925. Masdevallia triaristella auct. non Rchb. f. 1876: sensu Foldats in Lasser, Fl. Venez. 15(3): 56. 1970. Epiphyte, cespitose, erect, shade-loving; leaves 2–7 cm long, inflorescence erect; flowers yellowish to purplish with orange sepal tails, petals translucent green, lip purple; dorsal sepal 9–21 mm long. Rain forests, 100–600 m; Amazonas (Río Arari, Río Pasimoni). Costa Rica, Panama, Colombia, Ecuador, Peru, Amazonian Brazil, Bolivia. ◆Fig. 529.

Fig. 529. Trisetella triglochin

150. TRIZEUXIS Lindl., Coll. Bot. t. 2. 1823. [Subtribe Oncidiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Twig epiphytes, usually sun-loving, inconspicuous, cespitose, erect to subpendulous, sometimes the whole plant yellowish. Roots many, long and wiry. Rhizome abbreviate. Pseudobulbs relatively small, oblong-cylindric or ellipsoid, strongly compressed, apically 1-foliate, enveloped by several imbricate sheaths of which the innermost 2–5 have foliar blades which, along with the pseudobulb leaf, make a fan-like plant. Leaves laterally compressed, ventrally sulcate, dorsally rounded to ancipitous, articulate, fleshy-coriaceous, falcate to rarely straight, linear to linear-lanceolate, acute. Inflorescences lateral, originating from the base of the pseudobulb or among the leaf sheaths, usually longer than the leaves, paniculate with the flowers produced successively in crowded terminal racemes; peduncle remotely bracted, terete; rachis abbreviate, lengthening with age; floral bracts inconspicuous, fleshy, narrowly triangular. Flowers minute, inconspicuous, campanulate, resupinate or not; pedicellate ovary about the same length as the sepals, terete; perianth segments subfleshy. Sepals very pale translucent green; dorsal sepal connate in the basal 1/2 with the petals, and about the same length as the petals, elliptic, acute to obtuse, concave and forming a hood over the lip; lateral sepals connate for their basal 1/2–2/3 into a flat synsepal lying beneath the lip, oblanceolate, obtuse to rounded; petals same color as sepals, broader than dorsal sepal, overlapping it and

608

O RCHIDACEAE

enclosing lip and column, ovate or ovate-elliptic, rounded to subacute. Lip very pale greenish yellow at base grading to orange at apex, fleshy, articulate at the base of the column, concave, 3-lobed in the apical 1/3, basal blade of lip parallel to column, oblong-oblanceolate; apical lobe lanceolate to ovate, deflexed in natural position; disk inconspicuously bicallose. Column erect, short, subterete, basally attenuate, footless, wingless; anther terminal, operculate, incumbent, 1-locular, conspicuous for column size; clinandrium shallow; pollinia 2, waxy, narrowly obpyriform, tegula about 2 times longer than pollinia, oblong, viscidium articulate, elliptic; rostellum transverse; stigma ventral. Capsules subspheric. Costa Rica, Panama, Colombia, Venezuela, Trinidad-Tobago, Guyana, Ecuador, Peru, Brazil, Bolivia; 1 species. Trizeuxis falcata Lindl., Coll. Bot. t. 2. 1823. Twig epiphyte; leaves 1.2–3 cm long; inflorescences 3–23 cm long. Rain forests, 50–600 m; Delta Amacuro (Caño Atoiba), Bolívar (basins of Río Caroní and Río Cuyuní). Widespread in northern Venezuela; Costa Rica, Panama, Colombia, Venezuela, Trinidad-Tobago, Guyana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 530.

Fig. 530. Trizeuxis falcata

151. ULEIORCHIS Hoehne, Arq. Bot. Estado São Paulo n.s. formato maior 1: 129, t. 144. 1944. [Subtribe Gastrodiinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Saprophytes, erect, shade-loving, originating from a subterranean tuber; tuber ellipsoid, horizontal in natural position. Roots few, thin. Stems simple or basally 1branched, white, pinkish, purplish, or tan-colored, semi-translucent, terete, clothed by short, membranous sheaths at nodes. Leaves reduced to linear, basal filaments or totally absent, colored as the stem. Inflorescence terminal, racemose, lax, 1–4-flowered. Flowers resupinate, spreading, opening simultaneously; floral bracts similar to the sheaths of the stem but longer, loosely fitting the pedicellate ovary; pedicellate ovary much shorter than the sepals, obconic, subtrigonous to cylindric. Sepals and petals fused into a ± tubular or obconic cup or hood (perianth cup), longitudinally opened at the ventral face, this opening totally or partially filled by the lip; perianth cup gibbous at base, 13–16 mm long, whitish, cream, pinkish, or tan-colored; free apices of the sepals and petals triangular to ovate, 1.5–3.5 mm long, the apices of the sepals somewhat longer. Lip hinged to column foot, simple, fleshy; blade somewhat concave, ovate to narrow-oblong or narrow-elliptic, margins sometimes undulate or crenate; disk smooth or black-verruculose. Column elongate, subcylindric, parallel to lip, basally produced into a well-developed, oblique foot; anther terminal, incumbent, operculate, 1-locular, pollinia 2, sectile; rostellum transverse, flat, horizontal; stigma ventral, emergent, longitudinally elongate. Capsules obovoid or oblong.

Vanilla 609

Panama, Venezuela, Guyana, Suriname, Peru, southeastern and Amazonian Brazil; 2 species, both in the flora area. Plants of Uleiorchis elongate considerably when in fruit, presumably to aid in seed dispersal. Key to the Species of Uleiorchis 1.

1.

Lip narrowly oblong-elliptic, 5 times or more longer than wide, 15–16 mm long, longer than or ± equaling the perianth cup; growing above 1000 m ..................................................................................................... U. liesneri Lip ovate-elliptic or ovate, 2.5–3 times longer than wide, 10–12 mm long, shorter than the perianth cup; growing between 100–300 m ....... U. ulaei

Uleiorchis liesneri Carnevali & I. Ramírez, Novon 3: 105, fig. 3. 1993. Saprophyte; flowers pinkish tan, apex of lip yellow; inflorescences 1-flowered. Forests on tepui slopes, 1000–1200 m; Amazonas (Cerro Yutajé). Endemic. ◆Fig. 531. Uleiorchis ulaei (Cogn.) Handro, Arq. Bot. Estado São Paulo n.s. formato maior 3: 144. 1958. —Wullschlaegelia ulaei Cogn. in Mart., Fl. Bras. 3(4): 144. 1895. Uleiorchis cogniauxiana f. maior Hoehne, Arq. Bot. Estado São Paulo n.s. formato maior 1: 129. 1944. Saprophyte; flowers brownish white, sometimes with a lilac or pink tinge; lip yellow; inflorescences 1–4-flowered. Rain forests, 100–300 m; Bolívar (lower Río Caura basin), Amazonas (Sierra de la Neblina). Zulia; Panama, Ecuador, Peru, southeastern and Amazonian Brazil. ◆Fig. 532.

Fig. 531. Uleiorchis liesneri

Fig. 532. Uleiorchis ulaei

152. VANILLA Mill., Gard. Dict. Abr. ed. 4, 3. 1754. [Tribe Vanilleae, Subtribe Vanillinae, Subfamily Vanilloideae]. Vanillophora Neck., Elem. Bot. 3: 134. 1790. Myrobroma Salisb., Parad. Lond. 1: pl. 82. 1805. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes, generally climbing, rarely erect and not climbing. Leaves coriaceous, fleshy, sessile or attenuate into a pseudopetiole, rarely reduced to scales or absent. Inflorescences short racemes, axillary, rarely subterminal. Flowers large and showy. Sepals similar, spreading or, less frequently, reflexed; petals similar to sepals. Lip simple or 3-lobed, the basal margins adnate to the column. Column elongate, footless and wingless or with inconspicuous wings; anther joined to the clinandrium margins, incumbent, convex, semiglobose or somewhat conic; pollinia powdered or granulate. Capsule elongated, fleshy, ± fragrant or not.

610

O RCHIDACEAE

Pantropics, U.S.A. (Florida), southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina; ca. 50 species, 8 in Venezuela, all in the flora area. Some species of this genus are cultivated for the fruits, which is the source of vanilla flavoring. Key to the Species of Vanilla 1. 1. 2(1).

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

4.

5(3). 5. 6(5). 6. 7(6). 7.

Plants leafless or nearly so, leaves when present, reduced and narrowly elliptic to ovate-elliptic, (2.8–)3–7 cm long, 3–3.4 mm wide ..... V. penicillata Plants with well-developed leaves ............................................................. 2 Plants often purplish or grayish purple; internodes much longer than leaves; leaves elliptic, 2.3–2.8(–3.1) times wider than longer; plants growing at 600–1500 m .............................................................. V. wrightii Plants greenish; internodes equaling or shorter than the leaves; plants growing usually below 1100 m .............................................................. 3 Floral bracts 2–4 cm long ........................................................................... 4 Floral bracts > 1.2 cm long ........................................................................ 5 Leaves membranaceous, acuminate, with few parallel veins and reticulate veins between them; floral bracts widely ovate; lip 3-lobed; plants growing in rain forests .............................................................. V. mexicana Leaves smooth-coriaceous, acute; veins parallel without reticulate veins between them; basal floral bracts widely elliptic to suborbicular, the apical ones narrrowly ovate; lip simple; plants growing in Mauritia palm swamps .......................................................................... V. palmarum Leaves linear or lanceolate, acute, 7.5–20.5 × 0.5–2.2 cm .............. V. odorata Leaves elliptic, ovate, or ovate-oblong, 7–23.7 × 2.8–6.3 cm .................... 6 Leaves elliptic, acuminate, basally attenuate; lip rhombic, acute, with an apical callus ................................................................................ V. gardneri Leaves oblong to ovate-oblong, entire or ± 3-lobed ................................... 7 Petals dorsally carinate; lip ± 3-lobed, the central lobe emarginate .................................................................................................. V. planifolia Petals dorsally ecarinate; lip simple, the central lobe not emarginate ................................................................................................... V. pompona

Vanilla gardneri Rolfe, Bull. Misc. Inform. Kew 1895: 177. 1895. Climber to 10 m long; perianth segments white, lip basally cream or yellow, apically orange-yellow. Rain forests, 100–900 m; Bolívar (near Icabarú), Amazonas (Río Cunucunuma, Río Capibara). Monagas; Amazonian Brazil. Vanilla mexicana Mill., Gard. Dict. ed. 8, no. 1. 1768. Vanilla inodora Schiede, Linnaea 4: 574. 1829.

Climber; flowers fragrant, olive-green, lip white with a yellow callus. Rain forests, 50– 1000 m; Delta Amacuro (Río Amacuro, Río Cuyubini, Vega del Guayo), Bolívar (Amaruay-tepui, La Escalera to Cerro Venamo region, Mata Cuchilla), Amazonas (Río Baría, near San Carlos de Río Negro, Yavita-Maroa). Anzoátegui, Aragua, Miranda, Sucre; Mexico, West Indies, northern South America. Vanilla odorata C. Presl, Reliq. Haenk. 1: 101. 1830.

Vanilla 611

Fig. 533. Vanilla odorata

Fig. 534. Vanilla planifolia

Vanilla ensifolia Rolfe, Kew Bull. 1892: 141. Climber; leaves 7–20 cm long, variously oblique; flowers slightly fragrant, sepals and petals greenish white, translucent, lip whitish green, with a pale yellow-striped throat; dor-

Fig. 535. Vanilla penicillata

sal sepal 4.5–5 cm long. Rare in rain forests, 50–300 m; Bolívar (lower Rio Caroní, mid and lower Rio Caura), Amazonas (Cerro Marahuaka, near Puerto Ayacucho). Zulia; Southern Mexico to Panama, Colombia, Ecuador, Peru, and Bolivia, possibly Brazil. ◆Fig. 533.

612

O RCHIDACEAE

Vanilla palmarum (Salzm. ex Lindl.) Lindl., Gen. Sp. Orchid. Pl. 436. 1840. —Epidendrum palmarum Salzm. ex Lindl., Gen. Sp. Orchid. Pl. 436. 1832. Epiphyte, pendent, growing in Mauritia palm swamps; flowers greenish yellow, lip white with a yellow callus. Rain forests, 50– 400 m; Bolívar (Altiplanicie de Nuria, Gran Sabana, El Potrero, La Paragua, near Represa Guri), Amazonas (San Juan de Manapiare). Anzoátegui, Portuguesa; Ecuador, Brazil. Vanilla penicillata Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 3: 324. 1965. Terrestrial-epiphyte; flowers pale green without, greenish cream-colored within. Rain forests, 100–800 m; Bolívar (Cerro Guaquinima), Amazonas (Río Guayapo, San Carlos de Río Negro). Amazonian Brazil. ◆Fig. 535. Vanilla planifolia Andrews, Bot. Repos. t. 538. 1808. —Vainilla. Terrestrial-epiphyte, climber; flowers greenish yellow. Rain forests, 50–400 m; Bolívar (La Paragua, La Vigía, Río Caroní

basin, middle Río Caura, San Pedro de las Dos Bocas, Tumeremo), Amazonas (near Puerto Ayacucho, Raudal Guaharibos, near San Carlos de Río Negro). Cojedes, Falcón, Lara, Mérida, Miranda, Monagas, Táchira, Zulia; U.S.A. (Florida), Mexico, Central America, West Indies, south to Brazil and Paraguay. ◆Fig. 534. Vanilla pompona Schiede, Linnaea, 4: 573. 1829. —Vainillón. Terrestrial-epiphyte, climber; flowers yellow, lip yellow-orange. Rain forests, 100–200 m; Bolívar (near Tumeremo), Amazonas (upper Río Orinoco, San Carlos de Río Negro). Distrito Federal, Miranda, common in the Coastal Cordillera; southern Mexico, West Indies, south to northern Argentina. Vanilla wrightii Rchb. f., Flora 48: 273. 1865. Terrestrial-epiphyte, climber; flowers dark tan, lip cream or creamish white. Lowland and tepui-summit rain forests, 100–1200 m; Bolívar (Altiplanicie de Nuria, La Escalera to Cerro Venamo region, Serranía Marutaní), Amazonas (Cerro Aracamuni). West Indies, Guyana, Suriname, French Guiana.

153. VARGASIELLA C. Schweinf., Bot. Mus. Leafl. 15: 150. 1952. [Subtribe Vargasiellinae]. by Gustavo A. Romero-González Terrestrial herbs, sometimes epiphytes, slender or robust, short or tall, rhizomatous. Stem elongate, decumbent, not modified into pseudobulbs, of apparently indefinite apical growth, entirely concealed by appressed, imbricate, tubular, leafbearing sheaths, the lower ones scarious and disintegrating, the upper ones leafbearing. Leaves membranous to subcoriaceous, congested when young and scattered at maturity, distichous, elliptic to oblong or ovate, acuminate, attenuate toward the base, plicate, articulate. Inflorescences lateral, arising from the axil of the leaf sheaths, erect, racemose, few- to many-flowered, longer than the leaf. Flowers subfleshy, resupinate; floral bracts concave, oblong or narrowly ovate, acute or acuminate, shorter than to nearly 1/2 as long as the pedicellate ovary. Sepals concave and, at least in species in the flora area, strongly reflexed; dorsal sepal narrowly oblong-ovate or ovate-oblong, obtuse, acute, or mucronate, lateral sepals similar, slightly oblique, sometimes dorsally keeled with the keel produced into a conspicuous mucro; petals erect, concave, shorter and broader than the sepals, oval to oblong-elliptic, obtuse to subacute. Lip sessile to clawed, sometimes articulate with the column foot, simple, when spread triangular-ovate to ovate-oblong, basally cordate, sometimes excavate, apically obtuse or abruptly apiculate, the margins involute, undulate, the basal 1/2 sometimes with 2 axial, fleshy, callose thickenings. Column

Wullschlaegelia 613

short, erect, slightly arched, semiterete, winged, produced into a short, fleshy foot; anther terminal; pollinia 4, yellow, cartilaginous, dorsoventrally flattened, ovate to obovate, attached in 2 subequal pairs to a short, subquadrate tegula, the tegula bifid, hooked. Venezuela, Peru; 2 species, 1 in Venezuela. Vargasiella venezuelana C. Schweinf., Bot. Mus. Leafl. 18: 219. 1958. Terrestrial to 2 m tall, decumbent, rhizomatous; pseudobulbs lacking; inflorescence erect, racemose; flowers purple. Low vegetation on tepuis, 1900–2300 m; Bolívar (Auyán-tepui, Macizo del Chimantá [Chimantá-tepui]). Endemic. ◆Fig. 536.

Fig. 536. Vargasiella venezuelana

154. WULLSCHLAEGELIA Rchb. f., Bot. Zeitung (Berlin) 21: 131. 1863. [Subtribe Wullschlaegeliinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial saprophytes, erect, slender, shade-loving, 10–60 cm long; the whole plant whitish, pinkish, flesh-colored, or pale brown, covered by substellate pubescence composed of T- or Y-shaped, 2(3)-furcate trichomes with short, 1-cellular, basal stalks, and 2-cellular longer branches; terminal segment of hairs ellipsoid, acute, or glandular-globose; pubescence denser on the inflorescence, pedicellate ovary, and outer surface of sepals. Roots dimorphic, narrow-ellipsoid, fasciculate, and tuberose, or thin and fibrous, both types present simultaneously. Stem terete, thinner toward apex, remotely, few-sheathed. Inflorescence terminal, racemose, many-flowered; rachis nodding when young, dense, lax when old; floral bracts 1/4–1/3 the length of the pedicellate ovary; pedicellate ovary obclavate. Flowers inconspicuous, resupinate or not, successive, fugacious, cleistogamous, spiraled on rachis; perianth segments hyaline-white within at anthesis, membranous, subcampanulate or

614

O RCHIDACEAE

not spreading at all; sepals and petals connivent; dorsal sepal free, much smaller than laterals; lateral sepals decurrent on prominent column foot, basally connivent to form a conspicuous, sack-like mentum or spur; petals free, subsimilar to dorsal sepal. Lip erect, not articulate, sessile, simple, obovate or oblong-obovate, concave, ± saccate basally, ecallose; disk pubescent. Column erect, short, thick, basally extended into a long, distinct foot; anther erect, terminal and embedded in the apex of the column, very fleshy, 2-locular with curved thecae; pollinia 2, sectile, consisting of many slender massulae, viscidium conspicuous, ventral with respect to the anther; clinandrium, large, fleshy, entire or ± dentate; rostellum vertical, ovoid, stipitate, 2-lobed; stigma entire, subterminal, located vertically below the anther and enclosed in a pit with a protruding, free, ciliate margin. Capsules obovoid. Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Paraguay, northern Argentina; 2 species, 1 in Venezuela. Wullschlaegelia calcarata Benth., J. Linn. Soc., Bot. 18: 342. 1881. Saprophyte 10–60 cm tall; pubescence pale or dark brown, very dense, terminal segment of trichomes elongate, ellipsoid, acute; flowers resupinate; dorsal sepal 13–26 mm long. Rain forests, ca. 50–500(–1000) m; Bolívar (Gran Sabana, near La Escalera, La Paragua, Macizo del Chimantá, Río Bonita, Río Parguaza), Amazonas (Cerro Yapacana, near Puerto Ayacucho, lower Río Ventuari, near San Carlos de Río Negro, Sierra de la Neblina, Yavita to Maroa road). Apure, Distrito Federal, Sucre, Zulia; West Indies, Panama, northwestern and Amazonian Colombia, Guyana, French Guiana, Ecuador, Peru, Amazonian Brazil. ◆Fig. 537.

Fig. 537. Wullschlaegelia calcarata

155. XERORCHIS Schltr., Repert. Spec. Nov. Regni Veg. 11: 44. 1912. [Subtribe Xerorchidinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Terrestrial herbs, medium-sized. Rhizome branched, underground; roots filiform, wiry, elongate, pubescent; stems simple or branched, in the apical part zigzag, basally covered with scarious sheaths, gradually developing leaf blades and then bracts toward the tip. Leaves not articulate with their sheaths, folding at the base, distichous, grass-like, lanceolate, spreading. Inflorescences in the stem apex or

Xerorchis 615

branches, few- to several-flowered; rachis strongly zigzag. Flowers small; floral bracts similar to the leaves but smaller. Perianth segments subsimilar, ligulate, narrow. Lip parallel to the column, pandurate, slightly 3-lobed due to a medial constriction, subspatulate, central lobe longer than the lateral ones. Column slender, semiterete, footless, with a subovoid tubercle at each side of the stigmatic surface, curved downward; anther terminal, 2-locular; pollinia 8 in 2 groups, pear-shaped or obovate, without stipe, jointed with a disciform viscidium. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia; 2 species, both in the flora area. Key to the Species of Xerorchis 1. 1.

Floral bracts linear; central lobe of lip orbicular; disk ecallose ................................................................................................ X. amazonica Floral bracts ovate-lanceolate; central lobe of lip cuneate-ovate; disk with 2 obliquely inserted calli in the middle ................................. X. trichorhiza

Xerorchis amazonica Schltr., Repert. Spec. Nov. Regni Veg. 11: 45. 1912. Terrestrial to 50 cm tall; flowers yellow. Lowland rain forests, 100–900 m; Bolívar (Cerro Guaiquinima, El Paují), Amazonas (Caño Ucata southeast of Síquita, Cerro Huachamacari, San Carlos de Río Negro, Sierra de la Neblina, Yavita to Maroa road). Peru, Amazonian Brazil. ◆Fig. 538. Xerorchis trichorhiza (Kraenzl.) Garay, Canad. J. Bot. 34: 241. 1956. —Epidendrum trichorhizum Kraenzl., Repert. Spec. Nov. Regni Veg. 25: 21. 1928. Terrestrial to 50 cm tall; flowers pale green to cream. Lowland rain forests, 700– 900 m; Bolívar (Río Churún near Guarimba). Colombia, Peru, Brazil, Bolivia.

Fig. 538. Xerorchis amazonica

616

O RCHIDACEAE

156. XYLOBIUM Lindl., Bot. Reg. 11: sub t. 897. 1825. [Subtribe Lycastinae]. by Germán Carnevali and Ivón M. Ramírez-Morillo Epiphytes or terrestrial herbs, usually shade-loving. Pseudobulbs usually fleshy, oblong, oblong-pear-shaped, sometimes not thickened and then, stem-like and narrowly cylindric, when young covered with scarious sheaths, then sulcate and naked, apically 1–3-foliate. Leaves subcoriaceous, oblong, lanceolate, elliptic, or oblanceolate, acute to acuminate, base attenuate into a short or elongate pseudopetiole, blade with 3–5 veins very prominent, vernation of the blade convolute. Inflorescences racemes, few- to many-flowered, short or elongated, straight or curved, originating from the base of the pseudobulb; peduncle covered with several sheaths. Flowers from relatively small and inconspicuous to medium-sized and ± showy, mostly white, cream-colored, yellowish, or dull maroon to brownish, lasting 4–8 days, shortly pedicellate; floral bracts as long as or slightly longer than the pedicellate ovary, rarely shorter. Sepals membranaceous, subequal; dorsal sepal free; lateral sepals oblique, wider than the dorsal sepal, inserted on the column foot and forming a conspicuous mentum; petals similar to dorsal sepal, slightly shorter and usually slightly oblique. Lip articulate to the apex of the column foot, simple to deeply 3-lobed; lateral lobes erect and folding the column; central lobe short, wide; disk with a ± elongate central callus and usually with many fleshy papillae or verruculose keels. Column short, erect, slightly curved, semiterete, with a prominent basal foot; clinandrium obliquely truncate; anther terminal, operculate, incumbent, 1-locular or imperfectly 2-locular; pollinia 4, cartilaginous, in 2 pairs, the members of each pair frequently ± connate. Capsule ellipsoid, erect, relatively large. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 25 species, 10 in Venezuela, 2 of these in the flora area. Key to the Species of Xylobium 1.

1.

Pseudobulbs broadly ovoid or almost spheric, 2–4 cm long; leaves 4– 4.5 times longer than wide, rounded to obtuse; inflorescences pendulous, shortly pedunculate, < 3 cm long; flowers brown or purplish, perianth segments obtuse; lip simple, pandurate ....................... X. colleyi Pseudobulbs narrowly cylindric-fusiform, 6–15 cm long; leaves approximately 7 times longer than wide, acute to acuminate; inflorescences erect, with peduncle 10–27 cm long; flowers yellow or greenish, perianth segments acute; lip ± 3-lobed ............................. X. pallidiflorum

Xylobium colleyi (Bateman ex Lindl.) Rolfe, Gard. Chron. ser. 3, 7: 288. 1890. —Maxillaria colleyi Bateman ex. Lindl., Edwards’s Bot. Reg. 24: misc. 87. 1838. Epiphyte; leaves 20–70 cm long; flowers chestnut brown with purple spots. Rain forests, 600–700 m; Bolívar (Altiplanicie de Nuria, Río Uairén). Distrito Federal, Miranda; Trinidad-Tobago, Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil. ◆Fig. 539.

Xylobium pallidiflorum (Hook.) G. Nicholson, Ill. Dict. Gard. 4: 225. 1887. —Maxillaria pallidiflora Hook., Bot. Mag. 55: t. 2806. 1828. Epiphyte; flowers pale yellow or pale green, with the base of lip and apex of the column orange. Rain or cloud forests, ca. 1600 m; Bolívar (Cerro Uaipán). Aragua, Distrito Federal, Monagas, Sucre; West Indies, Ecuador, Peru, Bolivia.

Zygosepalum 617

Fig. 539. Xylobium colleyi

157. ZYGOSEPALUM Rchb. f., Ned. Kruidk. Arch. 4: 330. 1859. [Subtribe Zygopetalinae]. by Gustavo A. Romero-González Epiphytes, sometimes terrestrial herbs, creeping to climbing, cespitose or rhizomatous. Stem modified into oblong-ovate or cylindrical pseudobulbs, sometimes laterally compressed, aggregate or distant on the rhizome, when young entirely concealed by distichous, scarious sheaths. Leaves 1 or 2, rarely 3 or 4, membranous to coriaceous, narrowly ovate or narrowly oblong-ovate to elliptic-obovate, acute to acuminate, plicate, attenuate toward the base, subpetiolate, articulate. Inflorescences lateral, arising from the base of the pseudobulbs, arching or erect, racemose, few- to many-flowered, shorter to longer than the leaves. Flowers membranous, resupinate, showy; floral bracts concave, narrowly ovate to ovate, shorter to longer than the pedicellate ovary. Sepals erect, narrowly ovate or ovate, acuminate, the lateral sepals sometimes slightly oblique; petals erect, narrowly ovate to ovate, acuminate, generally narrower than the sepals, sometimes slightly oblique. Lip subsessile to clawed, spreading to slightly reflexed, ovate or very broadly ovate to subcircular, simple to apically ± 3-lobed, basally cordate to slightly cuneate, apically obtuse or acute to abruptly apiculate; disk with a horseshoe-shaped or flabellate callus, radially costate, the margins crenate to irregularly dentate. Column erect, slightly arched, semiterete, apically auriculate, produced into a short, slender foot; anther terminal, apically hooked; pollinia 4, yellow, cartilaginous, dorsoventrally flattened, subequal, attached in 2 pairs to a short, subquadrate to rhomboid tegula. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 7 species, 4 in Venezuela, all in the flora area. Key to the Species of Zygosepalum 1. 1.

Plants found on high plateaus and forested tepui slopes or summits, generally above 1000 m; rhizome with aggregate pseudobulbs ................. 2 Plants found in wet lowland forests, below 500 m; rhizome with distant pseudobulbs ............................................................................................ 3

618

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

O RCHIDACEAE

Pseudobulb to 3 cm long; inflorescence ≤ 15 cm long, with 1 or 2 flowers; lip narrowly ovate and abruptly apiculate ..................... Z. angustilabium Pseudobulbs 3.5–6 cm long; inflorescence ≥ 35 cm long, with 2–6 flowers; lip ovate or rhombic-ovate and acuminate ....................................... Z. tatei Bract as long as or longer than the pedicellate ovary; lip white with some violet markings ......................................................................... Z. labiosum Bract 1/2 as long as the pedicellate ovary; lip flushed with red, with strongly marked red veins .......................................................... Z. lindenii

Zygosepalum angustilabium (C. Schweinf.) Garay, Orquideología 8: 34. 1973. —Zygosepalum tatei var. angustilabium C. Schweinf., Fieldiana, Bot. 28: 194. 1951. Epiphyte; pseudobulbs aggregate; inflorescence short, erect, with 1 or 2 flowers; sepals and petals pale yellow or olive green with pale brown transverse markings; lip white or creamy yellow with purple or dull brown at the base. Wet forests on high plateaus, Bonnetia forests on tepui summits, 1300–2500 m; Bolívar (Auyán-tepui, Carraotepui, Kamarkawarai-tepui, Kilómetro 122 on road to Santa Elena de Uairén, Macizo del Chimantá [Chimantá-tepui]). Endemic. Zygosepalum labiosum (Rich.) Garay, Orquideología 1: 2. 1967. —Epidendrum labiosum Rich., Actes Soc. Hist. Nat. Paris 1: 112. 1792. —Medadenium labiosum (Rich.) Cogn. in Mart., Fl. Bras. 3(5): 582, t. 108. 1902. Epiphyte, climbing; pseudobulbs ovate, compressed, distant on the rhizome; inflorescence with 1–3 white to greenish or yellowish white flowers with purple markings at the base of the lip. Wet forests, 50–400 m; Delta Amacuro (Río Amacuro), Bolívar (Canaima, upper Río Paragua, Ucaima), Amazonas (Río Cataniapo, Río Sipapo, Río Yatúa). Guyana, Suriname, French Guiana, Brazil (Amazonas, Pará). ◆Fig. 540. Zygosepalum lindeniae (Rolfe) Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 3: 336. 1965. —Zygopetalum lindeniae Rolfe, Lindenia 6: 73, t. 275. 1890. —Medadenium lindeniae (Rolfe) Cogn. in Mart., Fl. Bras. 3(5): 584. 1902. Epiphyte, climbing; pseudobulbs ovate, compressed, distant on the rhizome; inflorescence with 1–3 white to yellowish white flowers with clearly marked red veins on the lip.

Wet forests, 50–200 m; Amazonas (Río Orinoco near San Fernando de Atabapo, Río Siapa, road between Yavita and Maroa). Colombia, Brazil (Amazonas). ◆Fig. 541. Zygosepalum tatei (Ames & C. Schweinf.) Garay & Dunst. in Dunst. & Garay, Venez. Orchid. Ill. 5: 318. 1972. —Zygopetalum tatei Ames & C. Schweinf., Bull. Torrey Bot. Club 58: 351. 1931. Lithophyte, rarely an epiphyte; pseudobulbs small, cylindrical, tapering slightly toward the apex, of 1 or 2 leaves; raceme erect, with 2–6 showy flowers; sepals and petals greenish to greenish yellow with brown to purple spots; lip white with purple at the base. Boggy vegetation and scrub forests on tepui summits, 1300–2500 m; Bolívar (Cerro Jaua), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú, Cerro Sarisariñama, Cerro Sipapo, Sierra de la Neblina). Brazil (Amazonas). ◆Fig. 542.

Fig. 540. Zygosepalum labiosum

O X A L I D A C E A E 619

Fig. 541. Zygosepalum lindeniae

Fig. 542. Zygosepalum tatei

OXALIDACEAE by Gerardo A. Aymard C. and Paul E. Berry Perennial or annual herbs, subshrubs, sometimes larger shrubs or trees. Leaves alternate, often forming basal rosettes or apical clusters; blades pinnately or palmately compound (then usually 3-foliolate), rarely 1-foliolate, usually exstipulate; leaflets entire, usually emarginate at the apex, pulvinulate. Inflorescence cymose and often appearing umbellate or racemose, axillary or appearing terminal, sometimes the flowers solitary. Flowers actinomorphic, bisexual, hypogonyous, sometimes cleistogamous. Sepals 5, imbricate; petals 5, free or shortly connate at the base, contorted. Stamens 10, biseriate, sometimes the outer antepetalous whorl staminodial; filaments free to basally connate, those of the outer whorl with basal nectariferous glands; anthers dorsifixed, versatile, longitudinally and introrsely dehiscent. Ovary superior, 3–5-locular, generally 5-lobed, each carpel with (1)2–15 pendulous ovules, placentation axile; styles 5, distinct, persistent; stigmas capitate or shortly bifid. Fruit a 5-lobed, loculicidal capsule or a berry. Seeds frequently arillate, with fleshy endosperm. Cosmopolitan; 6 genera and ca. 900 species, 2 genera and 8 species in the flora area.

620

O XALIDACEAE

Averrhoa carambola L. (Carambola, Tamarindo chino, Star fruit) is a small tree that is cultivated in some towns in northern Bolívar state for its fleshy, starshaped, acidic fruits. Key to the Genera of Oxalidaceae 1. 1.

Leaves pinnately many-foliolate, tightly clustered at the ends of the stems ...................................................................................... 1. Biophytum Leaves palmately 1- or 3-foliolate, rarely 4-foliolate, loosely distributed along the length of the stems ........................................................ 2. Oxalis

1. BIOPHYTUM DC., Prodr. 1: 140. 1824. Subshrubs with erect to decumbent stems terminating in a congested cluster of leaves and inflorescences, usually bearing a ring of stiff retrorse hairs at the tip. Leaves elongate, short-petiolate, paripinnate (the terminal leaflet reduced to a bristle); leaflets opposite, shortly petiolulate and articulate at the rachis, usually inequilateral. Inflorescences axillary, cymose or few-flowered, pseudo-umbellate, sessile to long-pedunculate, the pedicels subtended by persistent, imbricate bracteoles. Sepals scarious, the inner ones narrower than the outer ones; petals 5, coherent near the middle, white, yellow, orange, pink, or red-purple. Stamens basally connate, all fertile or sometimes the shorter whorl sterile. Ovary 5-lobed, each locule with 2–6 ovules; styles each with an enlarged subcapitate or papillate stigma. Fruit capsular, ovoid to subglobose or ellipsoid, the valves opening upward and remaining attached at their base, the sepals persistent. Seeds 1–6 per carpel, arillate. Pantropics; ca. 80 species, 5 or 6 in Venezuela, 5 of these in the flora area. Biophytum somnians (Mart. & Zucc. ex Zucc.) R. Knuth from Amazonian Colombia, Ecuador, Peru, and Brazil is likely to be found in southern Venezuela. It has pedunculate inflorescences, but fewer and much larger leaflets than B. calophyllum. Key to the Species of Biophytum 1. 1. 2(1). 2. 3(2).

3.

4(3).

Leaf rachis 9–25 cm long; penduncles 5–15 cm long ............. B. calophyllum Leaf rachis 2–10 cm long; inflorescences sessile or subsessile (peduncles < 1 cm long) ............................................................................................ 2 Leaflets elongate, ca. 5 × 1.25 mm, the apex acute ....................B. cardonaei Leaflets obovate-obdeltoid to squatly rhomboid, ca. 5 mm wide, the apex broadly rounded or obliquely truncate .................................................. 3 Leaves 2–6 cm long; leaflets in 5–9 pairs, obovate-obdeltate to broadly oblong, rigid-coriaceous, the venation prominent on the upper surface and the margins thickened, the apex broadly rounded; petals light purple; tepui summit above 1500 m ................................................ B. sp. B Leaves 4–10 cm long; leaflets mostly in 10–18 pairs, broadly and obliquely rhomboid, chartaceous to coriaceous, venation prominent to obscure on the upper surface, the margins not thickened; the apex broadly acute to obliquely truncate; petals white; lowland forests and riversides below 1000 m .................................................................................................... 4 Densely branched subshrub 0.5–1.5 m tall; leaves 10–18 per apical rosette, 6–10 cm long; leaflets chartaceous to subcoriaceous, 8–12-veined,

Biophytum 621

4.

the venation obscure on the upper surface; peduncle 2–7 mm long; sepals 6–8-veined; petals obovate; capsules appressed-pubescent at the apex .................................................................................................. B. sp. A Branched subshrub 0.3–0.5 m tall; leaves 8–13 per apical rosette, 4–6 cm long; leaflets coriaceous, 12–14-veined, the venation reticulate on the upper surface; peduncle ca. 2 mm long; sepals 12–14-veined; petals oblong; capsules glabrous ....................................................................B. sp. C

Fig. 543. Biophytum calophyllum

Biophytum calophyllum (Progel) Guillaumin, Bull. Mus. Hist. Nat. (Paris) 15: 125. 1909. —Oxalis calophylla Progel in Mart., Fl. Brasil. 12(2): 517. 187. —Poko mën (Panare). Biophytum casiquiarense R. Kunth, Notizbl. Bot. Gart. Berlin-Dahlem 7: 317. 1919. Biophytum passargei R. Kunth, Notizbl. Bot. Gart. Berlin-Dahlem 7: 318. 1919. Mostly single-stemmed shrub 40–80 cm tall; leaves often drooping; inflorescences erect, petals white with a yellow base. Understory of lowland evergreen and semideciduous forests, 100–400 m; Bolívar (20 km east of Túriba, 6 km from Maniapure to Caicara), Amazonas (Culebra, Galipero ca. 30 km north of Puerto Ayacucho, Puerto Ayacucho, Río Casiquiare basin, Río Cuao, Río Cunucunuma, Río Guainía, Río Ocamo, Yutajé). Carabobo, Táchira; Amazonian Colombia (Vaúpes) and Brazil. ◆Fig. 543.

622

O XALIDACEAE

4 mm

5 cm

Fig. 544. Biophytum sp. A

Biophytum 623

2 mm

2 mm

5 mm 5 cm

Fig. 545. Biophytum sp. C

Fig. 546. Biophytum sp. B

624

O XALIDACEAE

Biophytum cardonaei Pittier, Bol. Soc. Venez. Ci. Nat. 11: 13. 1947. Small shrub 40–80 cm tall; petals white. Tepui meadows, shady stream banks, 300– 1700 m; Bolívar (Cerro Guiaquinima, Río Carapo). Guyana? Biophytum sp. A Branched subshrub 0.4–1.5 m tall; petals white. Stream banks, river gravel bars, low forest understories, 100–300(–1000) m; Amazonas (Caño Surumoni, Culebra, between Cerro Duida and Cerro Marahuaka near base of Cerro Duida, 15 km west of La

Esmeralda, upper Río Cunucunuma basin). Endemic. ◆Fig. 544. Biophytum sp. B Branched subshrub ca. 20–35 cm tall, petals light purple. Along stream in tepui scrub, ca. 1700 m; Bolívar (Cerro Guanay). Endemic. ◆Fig. 546. Biophytum sp. C Branched subshrub to 40 cm tall, petals white. Understory of riparian forests, 50– 200 m; Amazonas (Caño Cabeza de Manteco in Río Autana basin, Río Cuao, Río Mawarinuma). Endemic. ◆Fig. 545.

2. OXALIS L., Sp. Pl. 433. 1753. Herbs or subshrubs (rarely vines); stems rhizomatous, bulbous, or aerial. Leaves alternate or subopposite to pseudoverticillate, basal or cauline, pinnately or palmately 3-foliolate (rarely 1-foliolate or 4-foliolate), petiolate, stipules sometimes present and connate; leaflets often obcordate. Inflorescences axillary, umbelliform cymes or solitary flowers; bracts small, bracteoles 2. Flowers usually showy, often dior tristylous (rarely cleistogamous and reduced). Sepals free or united near the base, imbricate, persisting in fruit; petals free, narrowed at the base, contorted in bud, caducous. Filaments basally connate. Ovary 5-locular, with 1–15 ovules per locule; stigmas capitate or rarely ellipsoid. Fruits capsular, cylindrical to oblong or globose, loculicidally dehiscent, each locule with (1)2–15 seeds, the valves remaining attached to the central axis of the fruit. Seeds usually ovoid, flattened on the sides, oblong to apiculate; testa chartaceous, longitudinally ribbed to transversally striate or sculptured and densely verrucose, external integument fleshy and arillate, breaking elastically and expelling the ripe seed; endosperm fleshy. Cosmopolitan; ca. 800 species, ca. 15–20 in Venezuela, 3 of these in the flora area. Key to the Species of Oxalis 1. 1. 2(1). 2.

Petals with a pinkish white blade and a white to yellow base; sepals subglabrous; upper surface of leaflets glabrous ..................... O. barrelieri Petals yellow throughout; sepals generally pubescent; upper surface of leaflets glabrescent to densely pubescent ............................................. 2 Upper surface of leaflets usually glabrescent; sepals hirsute-pilose to glabrous; flowers and capsules nodding ................................. O. frutescens Leaflets densely pubescent on both surfaces; sepals densely appressedpubescent; flowers and capsules erect .................................... O. juruensis

Oxalis barrelieri L., Sp. Pl. ed. 2, 624. 1762. —Trébol, Trébol sabanero. Herb or subshrub; flowers and fruits nodding, petals pink-white with yellowish white base. Deciduous forests, edges of savannas and brush, 50–300 m; Bolívar (near El

Manteco, Represa del Guri, Río Botanamo, Río Orinoco at El Torno, San Martín de Turumbán). Widespread in northern Venezuela; Central America, West Indies, tropical South America, introduced in Africa and Polynesia. ◆Fig. 547.

P A S S I F L O R A C E A E 625

Oxalis frutescens L., Sp. Pl. 435. 1753. Oxalis pentantha Jacq., Oxalis 23. 1794. Herb or subshrub; flowers and fruits nodding, petals yellow. U.S.A. (Texas), Mexico, Central America, West Indies, south to northwestern Argentina; 2 subspecies, both in the flora area. Key to the Subspecies of O. frutescens 1. Subshrub; leaflets suborbiculate or oblong, glaucous, shortly pilose to glabrescent; fruit glabrous ........... subsp. borjensis 1. Herb or subshrub; leaflets obovate, ovate or oblong, pubescent to tomentose; fruit pilose ....................... subsp. frutescens

cucho, Raudal de Atures, Río Cataniapo). Apure; Colombia (Vichada). Oxalis juruensis Diels, Verh. Bot. Vereins Prov. Brandenburg 48: 173. 1907. Branched shrub to 2 m tall; flowers and fruits erect, petals yellow. Along river rapids, 100–200 m; Amazonas (upper Río Orinoco at Raudal de los Guaharibos and Raudal Peñascal). Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia.

O. frutescens subsp. frutescens. —Tripa de pollo. Savannas, secondary vegetation, Mauritia palm swamps, 50–800 m; Bolívar (northeast of Canaima, Ciudad Bolívar, northwest of El Manteco, west of Kamarata, Puerto Ordaz, Santa Elena de Uairén). Widespread in northern Venezuela; U.S.A. (Texas), Mexico, Central America, West Indies, south to northwestern Argentina. O. frutescens subsp. borjensis (H.B.K.) Lourteig, Phytologia 29: 463. 1975. —Oxalis borjensis H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 249. 1821 [1822]. Open areas, secondary vegetation, 50–200 m; Bolívar (Río Orinoco at Raudal San Borja), Amazonas (Isla Ratón, Puerto Aya-

Fig. 547. Oxalis barrelieri

PASSIFLORACEAE by Stephen S. Tillett Trees, shrubs, herbs, or most frequently vines or woody lianas climbing by means of usually axillary tendrils. Leaves alternate and spirally arranged; stipules small to large, frequently deciduous; petiole normally present; blade simple, variously lobed, divided, or compound; margin entire to variously toothed. Extrafloral glands frequently present, either on the petiole, blade margin, or lower surface, and on bracts or sepals; glandular hairs less frequent. Basic inflorescence compound-cymose with peduncle of the central flowers converted into a tendril, in most taxa the inflorescence reduced to an axillary tendril beside one or more inflorescences or accessory branch initials that may or may not develop; branched tendrils infrequently present.

626

P ASSIFLORACEAE

Peduncle normally axillary to a leaf, supporting a pedicel, each of varying size, and enlarging and/or elongating in fruit, usually with an articulation between, and subtended frequently with 3 minute to large bracts on these supporting structures, which, as with some tendrils, may also have small subtending bracts, all a result of derivation from a compound inflorescence. Flowers normally bisexual (in all neotropical species), a few Old World species monoecious or rarely dioecious; a simple receptacle or torus may be present, but this generally enlarged and greatly modified into a disk, cup-like or tubular structure, with the addition of tissues derived from perianth and androecium. Perianth normally actinomorphic, the calyx and corolla usually 5-merous (but 3–8-merous in some genera); sepals free or basally connate, often petaloid; petals sometimes absent. Between perianth and androecium, there may be 1–several verticils (series) of accessory structures derived variously from perianth tissue, the androecium (staminodial), or from axial tissue; these are involved in the attraction, control, and compensation (nectar) of pollinators (normally insects, birds, and bats). Androecium of (3–)5(–10) stamens, free or slightly united basally, frequently borne upon a short or quite long androgynophore; filaments usually long; anthers versatile and opening by longitudinal slits. Gynoecium superior, frequently borne upon a short gynophore, this above the insertion of the stamens on the androgynophore when present; ovary syncarpic, carpels 3(–5), 1-locular with 3 (–5) parietal placentas and an equal number of styles, these free or united; stigmas large, capitate, reniform, or discoid. Fruit a berry or capsule. Seeds enveloped by a usually fleshy aril produced from the apex of the relatively long funiculus; testa thick, hard, and sculptured in rather diagnostic patterns; embryo with large, flat cotyledons, in the center of an ample, oily, fleshy endosperm. Tropical and warm-temperate regions of both hemispheres in all vegetation types, from tropical rain forests to deserts and páramos, with Passiflora native as far north as Virginia and Missouri in the U.S.A., and south to central Argentina and Uruguay; 18 genera and ca. 700 species; 3 genera and 53 species in the flora area. Many members of the family produce cyanogenic glucosides; alkaloids and flavonoids are also present. Basic Passifloraceae References for the Venezuelan Guayana Escobar, L. K. 1994. Two new species and a key to Passiflora subg. Astrophea. Syst. Bot. 19: 203–210. Gentry, A. H. 1981. Distributional patterns and an additional species of the Passiflora vitifolia complex: Amazonian species diversity due to edaphically differentiated communities. Pl. Syst. Evol. 137: 95–105. Killip, E. P. 1938. The American species of Passifloraceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19: 1–613. Tillett, S. S. 1988. Passionis passifloris II. Terminología. Ernstia 48: 1–40. Key to the Genera of Passifloraceae 1.

Normally herbaceous vines to large woody lianas, but also subscandent shrubs, or trees; leaves simple to variously lobed or divided, membranaceous to coriaceous, tendrils simple; ‘hypanthium’ formed of floral cup and floral tube supporting 1–several verticils (series) of acces-

Ancistrothyrsus 627

1.

2(1).

2.

sory structures; perianth 5-merous; stamens 5; carpels, styles, and stigmas 3 .................................................................................. 3. Passiflora Woody lianas or small trees; leaves simple, stiff-coriaceous, tendrils apically very shortly trifid or in the form of thickened hooks; receptacle not expanded, this or perianth supporting 1 or 2 verticils of accessory structures; perianth 4-merous; stamens 8; carpels, styles, and stigmas 4 ................................................................................................ 2 Tendrils simple, inflorescence with a short tendril, grossly thickened and hooked; corona in 1 verticil, tubular or filamentous, included; fruit a capsule; plants pubescent and with pellucid glandular scales .........................................................................................1. Ancistrothyrsus Tendrils (when present) trifid at very apex, inflorescence lacking a tendril; corona in 2 exserted verticils, the inner with a mass of fine crisped threads; fruit a coriacous or hard-shelled berry; plants glabrous ........................................................................................................ 2. Dilkea

1. ANCISTROTHYRSUS Harms, Notizbl. Bot. Gart. Berlin-Dahlem 11: 146, fig. 4. 1931. Large canopy lianas, pubescent or hirtellous. Branches terete. Leaves alternate, petiolate; blades simple, entire, scabrellous, pilose, or hirtellous, with reddish patelliform (saucer-shaped) glandular scales; stipules apparently early deciduous. Inflorescences axillary, supported on a long branch terminating in 2 simple dichasial cymes (occasionally once-compound) on each side of a thickened, hooked or coiled tendril. Flowers of moderate size, showy, white, borne on short peduncles and pedicels; receptacle simple, not formed of a floral cup and tube. Sepals and petals similar in size and form. Corona a single cylindric series of filaments, fused basally or almost completely. Androgynophore very short. Stamens 8; filaments free or very shortly connate at base; anthers versatile. Gynophore short; ovary narrow, 4-carpellate; ovules 2–5 on each parietal placenta; styles 4, slender; stigmas large, capitate or reniform-capitate. Fruit a large, 4-valved, globose or ovoid, relatively thin-walled capsule. Seeds large, transversely shallowly costate-rugulose, arillate, not filling the locule. Mainly lowland tropical forests of northeastern Amazonia, in Colombia, Venezuela, Ecuador, Peru, and Brazil; 2 species, 1 in Venezuela. Ancistrothyrsus is rarely collected, but capsules or their valves may be found on the forest floor. Ancistrothyrsus hirtellus A.H. Gentry, Novon 2: 333, fig. 1. 1992. Large canopy liana, branchlets hirtellous with somewhat crisped trichomes; tendril stout, helicoidal on young growth; petiole 8– 15 mm long, grooved adaxially, densely rufous-strigose; blade obovate to oblong-elliptic, 7–24 × 3.5–12 cm, base broadly obtuse to rounded, apex rounded, entire, upper surface with conspicuous reddish patelliform-glandular scales and sparsely hirtellous, lower surface densely hirtellous and equally patel-

liform-glandular; inflorescence on an axillary branch, 4–13 cm long; sepals lanceolate or lance-elliptic, ca. 30 × 7–8 mm, acute, densely tan-puberulous without, persistent in fruit; petals lanceolate, ca. 25 mm long; corona forming a tube around the staminal filaments and ovary, densely white-pilose; staminal filaments 7–9 mm long, arising from base of ovary at the apex of a 4–5 mm long androgynophore; ovary ovoid, densely villous; fruiting capsule yellow-brown, 6–9 × 3.5–4 cm, widely ovoid, densely tan-tomen-

628

P ASSIFLORACEAE

Fig. 548. Ancistrothyrsus hirtellus

tose and -velutinous (the 2 trichome types intermixed), and with reddish patelliformglandular scales, base rounded and confluent with the 1–2 cm long conically enlarged gynophore, apex bluntly acute, 4 valves woody but appearing somewhat fragile, ca. 3 mm thick; seeds flat, obovate, 12–14 mm long,

transversely shallowly costate-rugulose, the aril completely enveloping the seed, but thin, fragile, and fragmented at maturity. Evergreen lowland, nonflooded forests, 100–300 m; Amazonas (base of Sierra de la Neblina). Amazon basin of Ecuador, Peru, and Brazil. ◆Fig. 548.

2. DILKEA Mast., Trans. Linn. Soc. London 27: 627. 1871. Lianas, subscandent shrubs (in open areas), or small trees, glabrous. Leaves alternate; stipules subulate-triangular, early deciduous; petiole stout, normally glandless; blade simple, penniveined, coriaceous, lustrous, quite variable in form, even on the same plant. Tendrils, when present, axillary, lateral to the inflorescence primordium, helically coiled, apically shortly (1–4 mm) and unequally trifid. Inflorescence axillary or terminal, flowers solitary, glomerate, or short-racemose; peduncles short to long; bracts basal, triangular. Flowers pedicellate, small to large, white, showy, fragrant; bracts basal. Sepals 4, initially coalescent below to form a tube, free and deciduous at the base when mature, 2 sepals wider and exterior to the other 2 and touching or separated laterally above. Petals 4 (or rarely 5?), free to base where adnate to sepals, more slender and thinner than sepals, deciduous. Corona adnate to base of perianth, in 2 series, the outer of erect, somewhat ligulate filaments, free or slightly united at the base, the inner tubular basally, divided above in segments margined and divided within into many fine, crisped threads, both series deciduous. Androecium of 8 hypogynous stamens; filaments erect, slightly united at the base into a tube surrounding the gynophore, white; anthers versatile, linear,

Dilkea 629

base slightly saggitate, introrse, yellow. Gynoecium borne on a short gynophore, 4merous, rarely (aberrantly?) 5-merous; ovary unilocular, placentae parietal; styles united to middle, exserted; stigmas large, reniform-capitate. Fruit a coriaceous- or hard-walled berry, globose or ellipsoid, yellowish. Seeds few, large, covered with a normally fleshy aril, the testa thin. Panama, Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, Brazil; 7–9 species, 1 in Venezuela.

Fig. 549. Dilkea acuminata

Dilkea acuminata Mast., Trans. Linn. Soc. London 27: 628. 1871. Dilkea johannesii var. parvifolia Hoehne, Comiss. Linhas Telegr. Estratég. Mato Grosso Amazonas, anexo 5, Bot. 5: 73, pl. 111. 1915. Dilkea parviflora Killip, Publ. Field Mus. Nat. Hist., Bot. Ser. 19: 575. 1938. Dilkea magnifica Steyerm., Acta Bot. Venez. 3: 186, fig. 12. 1968. High-climbing liana, with stems attaining 4 cm diameter or more, or, as regrowth in opened spaces, a shrub producing long wiry stems with tendrils above scars of very early deciduous reduced leaves; stipules 1–2 mm long, linear, thin, very early deciduous; petiole stout, 1–4 cm long, very rarely with 1 or 2 scar-like glands near apex; blade extremely variable even on the same plant, 6–30 × 4–14 cm, broadly to narrowly ovate, elliptic, oblong, oblanceolate, or obovate, the base acute, obtuse, or rounded, the apex acuminate, acute, obtuse, rounded, emarginate, or retuse with a short apiculate tip; margin entire, coriaceous, normally lustrous; peduncles axillary, usually solitary on young growth, or in clusters on the older stems, infrequently shortracemose; bracts basal on the axis of clusters

and peduncle, subulate, ca. 1 mm long; flowers to ca. 7 cm wide, white, perianth reflexed; sepals 4, oblong-oblanceolate, 15–65 × 5–10 mm, the inner narrower, apex rounded to subacute; petals subequal and thinner; the outer corona filamentous-ligulate, 2/3 as long to almost as long as sepals, the inner slightly shorter; free portion of staminal filaments to 30 mm long, anthers narrow, to 9 mm long, slightly saggitate at base; ovary fusiform, to 5 mm long, styles and stigmas 4, the styles united shortly or nearly to middle, erect, extending ± to the top of the corona, the stigmas capitate to reniform, ca. 3 mm diameter; fruit to 8 cm long, apex rounded to sharply apiculate, pericarp coriaceous, locule not filled by the few, large oblong-ellipsoid seeds, the (edible) aril normally fleshy, sweet. Evergreen lowland to lower montane forests, 100–600 m; Delta Amacuro (Sierra Imataca),

630

P ASSIFLORACEAE

Bolívar (apparently generally distributed to the south and west), Amazonas (over most of the area, particularly in the south). Panama, Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 549. This is a variable species, but one that is recognized here as distinct from Dilkea retusa Mast., which is considered limited to

the area around Manaus and farther downriver. Both Dilkea acuminata and D. retusa show a bewildering variability in leaf shape. They can be differentiated in late bud, when the two outer sepals of D. retusa are essentially valvate, whereas in D. acuminata the margins do not touch, revealing the midrib area of the two inner sepals.

3. PASSIFLORA L., Sp. Pl. 955. 1753. Mainly suffrutescent or woody vines climbing by means of tendrils, some subscandent or erect shrubs, and several trees; most herbaceous species perennial and slightly or definitely woody near the base. Pubescence, when present on the plant, very variable, though not with branched hairs; in 2 subgenera large, multicellular, stipitate, vascularized glandular hairs present. Leaves alternate and of extreme variability in the genus as to form and size, and equally so in some species or individuals (with variation in size sometimes greatly affected by soil fertility); juvenile leaf, or at times those of regrowth, may be completely different from the adult in form and texture (all evidently an evolutionary strategem to evade predators). Stipules lateral to the petiole, varying from mere bristles to large blades, the latter almost always asymmetrical with the narrow side toward the petiole; a few species have pinnatisect stipules, a condition absent in the leaf blade; stipules may be early deciduous. Petiole always present, although may be very short, frequently bears one or more pairs of petiolar glands, of variable form, size, and position, from scars to stipitate glands, easily distinguished from glandular hairs. Leaf blade varies in size from 1 cm to over 1 m long; secondary veins pinnate or mostly palmately 3–5-veined; in form varying from unlobed to 2- or 3-lobed, or to palmately or pedately lobed or compound; margin entire or with varying size and form of toothing, frequently with either marginal glands (very infrequently submarginal on the underside) or laminar glands (ocelli) on the lower surface, the latter normally between the midvein and the 2 basal lateral veins. Inflorescence basically an axillary cyme, as in the tree species and some Asian vines and other genera; in most of the American vine species, flowers normally occur singly or in pairs, but a few present cymes, pseudoracemes, or true racemes. Tendril, in the great majority, springing directly from the leaf axil; tree species lack tendrils, and those of shrubs often convert to woody spines; a few American species have a tendril in the inflorescence; the tendrils coil helicoidally and are normally unbranched. Peduncles most frequently 1 or 2 per axil, each normally supporting a pedicel, almost always separated by an articulation; the base of the flower usually invaginated (umbilicate) at the insertion of the pedicel. Bracts normally 3, minute to very large, usually verticillate and involucrate, free or united, at or near the articulation between peduncle and pedicel, but may be dispersed and in positions other than the articulation (or sometimes absent). Flowers small (1 cm) and inconspicuous to large or long and very showy; in species with 1 or 2 per axil, flowers develop continually along the stem, normally with only 1 or 2 axils each with one open flower, relatively short-lived; a very few species are mast-flowered. Floral cup (derived from axillary tissue) supporting the floral tube (derived from the perianth), formerly considered together as a hypanthium (and so referred to in the key and descriptions, for brevity, when considered together); floral cup varies from a flat disk to a tall cup, floral tube arising from the floral cup, frequently tubular or

Passiflora 631

obconic, but in many species reduced to a mere ring of tissue surmounting the floral cup. Sepals and petals 5(–8 in some Asian species), quincuncially imbricate; petals normally springing from apex of floral tube, rarely absent; sepals normally free, may be corniculate without near the apex. Corona, between the perianth and the androecium, formed of various structures derived from perianth, staminodial, and axillary tissues (and making the structure of the flower difficult to describe, but of major taxonomic value, particularly since these parts determine which animal is the effective pollinator (see Fig. 550 and Tillett 1988), composed of (from the outermost to innermost): radii, formed of one or more verticils (series) of relatively long filaments; pali, of one or more series of normally rather short filaments, sometimes united into a membrane; operculum, generally a delicate membrane, delimiting the inner base of the floral tube where it arises from the floral cup; annulus, a raised ring of tissue; nectary, a band of secretory tissue which may form a raised ring or be fused to the annulus; limen (derived from a former ring of staminodia) generally at the base of the androgynophore or upon it, or somewhat out on the floral cup, serving as a seat for the rim of the operculum, and closing off the nectar chamber (and by their structure determining which visitors may obtain nectar), it may be fused with the annulus. Androgynophore (gynandrophore) from less than 1 mm to more than 10 cm long; may produce a bulbous swelling near the base, the trochlea, to which the limen may be fused. Androecium formed of 5(–8 in some Asiatic species) stamens inserted at apex of the androgynophore; filaments may be united at the base into a short, narrow tube; anthers are relatively large, dorsifixed, introrse in bud, extrorse in the open flower. Gynoecium may be borne on a short gynophore; normally 3(–5 in some Asiatic species) carpels; ovary unilocular with parietal placentation; ovules anatropous with relatively large funiculi, borne in 2–4 rows on each placenta; styles, equal in number to carpels, arise almost united at their base or individually separated away from the common center at the apex of the ovary, frequently claviform; stigmas large, normally capitate or reniform. Fruit most commonly a berry (in a few an irregularly dehiscent ‘capsule’), from less than 1 cm diameter to the size of a melon; exocarp membranous, coriaceous, or stiff and brittle, mesocarp spongy or fleshy, endocarp a thin membrane; dehiscence in some of the capsular forms is by a rather violent irregular rupture; (the androgynophore immediately supporting the fruit should not be confused with the pedicel). Seeds usually numerous, of medium size in proportion to that of the fruit, mostly flattened, testa thick, hard, normally sculptured in a characteristic, frequently species-diagnostic pattern, enclosed in a fleshy, translucid, usually colorless aril, which may be sweet or acid. Tropical and subtropical North America and South America, Southeast Asia to Australia; ca. 500 species, all but 20 in the Americas, 85–100 in Venezuela, 51 of these in the flora area. In addition to the 3 species cited as Passiflora sp. A, B, and C, there are several more obviously different taxa still without sufficient material for their characterization. Almost all species are heliophilic, requiring full sunlight on the leaves for flowering, even those normally of forests, which must reach the canopy before flowering (as Passiflora pyrrhantha and P. quadriglandulosa, which usually flower on old wood near the ground). Many form large populations in secondary growth. The seeds have evolved to pass though the digestive tract of animals, and those with palatable fruits have been widely spread by man beyond their natural ecologic or geo-

632

P ASSIFLORACEAE

graphic ranges. This has contributed to hybridization and introgression among species, which makes identification in some species groups very problematic. In some large-flowered species, a scent is produced by the radii, which also serve as a landing platform for insect pollinators. The vegetative portions of the plant may contain high concentrations of cyanogenic glucosides, and the immature fruits may contain enough to cause death in animals and man. The aril is the major edible part of the fruit; probably most species are edible when fully ripe, though many are not very palatable. Some Asian species of Passiflora present well-developed cymes in which the development of the tendril from parts of the inflorescence can be seen; a few American species have a tendril in the inflorescence, but in the great majority the tendril springs directly from the leaf axil and lateral to the flower bud primordium, with the branch primordium above these. The tendrils coil helicoidally (as a normal coiled spring, not in a flat watch-spring as in the Sapindaceae), and are normally unbranched (not to be confused with Cucurbitaceae, with mainly branched tendrils arising at the node a quarter turn from the petiole base, or Vitaceae, with tendrils arising at the node, but opposite the leaf). To facilitate identification, collections should include, when possible, at least one flower sectioned longitudinally, leaving the pedicel and androgynoecium complete on one of the halves.

Fig. 550. Diagrammatic longitudinal half-sections of representative Passiflora species. Left to right: P. vespertilio L., P. nitida H.B.K., P. quadriglandulosa Rodschied, P. sclerophylla Harms, P. securiclata Mast. The dotted line indicates the limit of the floral cup and the beginning of the floral tube (in solid black). The androgynophore (unlabeled) is the central axis bearing the stamens and the ovary. pu = peduncle, ar = articulation, pi = pedicel, br = bract, C = calyx + corolla, r = radii, p = pali, l = limen, t = trochlea, o = operculum, a = annulus. See Passiflora genus description; drawings not to scale.

Passiflora 633

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

Leaf pedately parted or compound ............................................................ 2 Leaf not parted or compound ..................................................................... 3 Plant densely to sparsely short-tomentose; leaf blades pedately 3-parted, the 2 lateral segments 3-parted to base or nearly compound; petiolar glands < 0.5 mm diameter, short-stipitate, near or below middle of petiole ............................................................................................ P. pedata 2. Plant glabrous to pubescent, glaucous; leaf blades pedately compound, 5or 7-foliolate, the leaflets markedly aristate-dentate at base on one or both sides; petiolar glands sessile, 2 mm diameter, plate- or scar-like, at or near base of petiole ............................................................. P. cirrhiflora 3(1). Leaves lobed ............................................................................................... 4 3. Leaves unlobed ......................................................................................... 23 4(3). Leaves symmetrically 2-lobed, 3-veined from base but the central lobe much reduced if present; petiolar glands absent .................................. 5 4. Leaves symmetrically 3-lobed (occasionally entire or asymmetrically 2-lobed on the same plant) or 5-lobed, the central lobe much the more prominent; petiolar glands or glandular hairs, present ............................... 10 5(4). Ocellae (laminar glands) absent; floral bracts absent; base of leaf blade deeply cordate to truncate, lobes mainly acute ............................. P. rubra 5. Ocellae present; floral bracts present; base of leaf blade cuneate, truncate, or shallowly cordate, lobes obtuse to acute ........................................... 6 6(5). Leaf longer than broad, the lobes ascendent ............................................ 7 6. Leaf broader than long to transversely rectangular, the lobes divergent to ascendent ............................................................................................... 8 7(6). Basal pair of ocellae larger than the others and close to but not completely occupying the angles between midvein and the principal lateral veins, usually a second pair of major lateral veins from the midvein just below the sinus ..................................................................................... P. tuberosa 7. Ocellae small, frequently 1 or 2 at the base outside the principal lateral veins, but not occupying the angles between these and midvein, without second pair of lateral veins ....................................................... P. sp. A 8(6). Leaf blade transversely rectangular, lateral lobes narrowly oblong to ovate or lanceolate, apex acute to rounded, obtuse, or retuse, mucronate, the central lobe normally absent or very short, truncate or rounded to amply retuse; 1(2) flower per node ............................ P. misera 8. Leaf blade broader than long, but not transversely rectangular, central lobe more frequently absent .................................................................. 9 9(8). Stipules narrowly linear-subulate to setaceous or falcate; ocellae evident; peduncles paired or solitary; floral bracts subulate or setaceous, 2– 3 mm long, greenish ................................................................. P. vespertilio 9. Stipules linear-falcate to curled; ocellae usually minute; peduncles stout, rarely paired; floral bracts ovate to elliptic, obovate, or suborbicular, verticillate, 10–22 × 4–10 mm, brownish to purplish red ....... P. pulchella 10(4). Petiolar glands absent, but plant in general covered with short, malodorous glandular-viscid hairs, plant otherwise glabrous to variously or densely hairy ................................................................................. P. foetida

634

P ASSIFLORACEAE

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

12. 13(12).

13. 14(13).

14. 15(14).

15.

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

18.

19(16).

19.

Petiolar glands present, plant without glandular-viscid hairs, otherwise glabrous or hairy .................................................................................. 11 Stipules broad, cordate or oblique (the half adjacent to petiole much narrower than the outer) .......................................................................... 12 Stipules setaceous to linear-lanceolate ................................................... 16 Stipules with a filiform apical awn ≥ 7 mm long, the awn normally oblanceolate-spatulate, swollen and yellow to black near tip (see also couplet 25) .............................................................................................. P. retipetala Stipules at most aristate .......................................................................... 13 Lateral lobes of blade almost at right angles to middle lobe; petiole minutely stipitate-biglandular at middle, stipules cordate or inequally cordate to semiovate ........................................................................ P. sp. B Lateral lobes of blade ascending; petiolar glands (2–)4–8, scattered; stipules semiovate or subreniform ...................................................... 14 Floral bracts scattered on lower half of peduncle; petiole biglandular, 1– 1.5 cm below blade, the glands ca. 2 mm diameter, normally long-stipitate (1–8 mm) .......................................................................... P. adenopoda Floral bracts verticillate at articulation of peduncle; petiolar glands 2– 8, minute to ca. 1 mm diameter, sessile to substipitate ..................... 15 Floral bracts sessile, subcordate, about as long as pedicel; leaf blade thickmembranous to -coriaceous, the lower surface whitish-glaucous; petiolar glands 4–8, sessile to substipitate, ca. 1 mm diameter ........P. garckei Floral bracts shortly petiolate, much longer than pedicel; leaf blade thinmembranous; petiolar glands 2–6, very thin-filiform to normal, 1– 1.5 mm long (see also couplet 25) ............................................. P. picturata Leaf margins entire (or at most with a single large tooth on each side, representing reduced lateral lobes) .......................................................... 17 Leaf margins toothed ............................................................................... 19 Petals present, delicate; petiolar glands large, sessile, auriculate, near base (see also couplet 28) ........................................................ P. auriculata Petals absent; petiolar glands slender, stipitate, capitate ...................... 18 Glabrous to densely pubescent perennial from rootstock, or older stems with thick, sharply angled corky ridges; glands near middle of petiole, large-capitate, cupped, sessile to stipitate; lobes of blade acute or obtuse, extremely variable, even on same plant, base rounded or peltate; fruit spherical, dark purple or black, 5–15 mm diameter; aril translucent, juicy (see also couplet 27) ................................................. P. suberosa Glabrous annual, stems without corky ridges; glands below middle of petiole, cupped, sessile; lobes of blade rounded-obtuse, base truncatecordate; fruit ellipsoid, scarlet or orange, 2–5 cm long; aril coral-red, mealy and oily .............................................................................. P. gracilis Petiolar glands basal; flowers red; at least outer series of radii red; hypanthium short-cylindric, to 2 cm long; plants glabrous or glabrescent, velutinous or densely tomentulose; floral bracts linear to linear-lanceolate, < 5 mm wide, margin glandular-serrulate (see also couplet 39) ..................................................................................... P. quadriglandulosa Petiolar glands apical or near middle; flowers white; radii white, banded with blue or violet-purple; floral cup and tube campanulate to funnelform ...................................................................................................... 20

Passiflora 635

20(19). Flowers erect; floral bracts < 2.5 cm long, not enclosing the bud, serrate to lacerate, at times glandular-toothed, thick-membranous to subcoriaceous, medium to dark green, free; leaves lustrous dark green, thick membranous (cultivated and escaped; see also couplet 45) ..........P. edulis 20. Flowers pendulous; floral bracts large, completely enclosing the bud, 2– 6 cm long, entire, thin-membranous, white to greenish white, at base partially united, above appressed by a marginal band of dense tomentum; leaves medium green, membranous to coriaceous ..................... 21 21(20). Petiolar glands normally one pair, near apex, short-stipitate, slightly cupped or auriculate (see also couplet 28) .............................. P. seemannii 21. Petiolar glands 1 or 2 pairs, near middle of petiole or below, sessile, not auriculate ............................................................................................. 22 22(21). Stipules narrowly linear-oblong to -oblanceolate, to 6 × 1.5 mm, minutely glandular-denticulate above, coriaceous; middle lobe of leaf blade obtuse or acute; floral bracts obovate, subobtuse to acute, united at base for ca. 5 mm (see also couplet 33) ............................................. P. serrulata 22. Stipules linear-attenuate, to 20 × 3 mm, shallowly glandular-dentate; middle lobe of leaf blade acuminate, the tip 1–2 cm long; floral bracts ovate, long acuminate-apiculate, > 1 cm long, united at base for ca. 1 cm (see also couplet 33) .............................................................. P. multiformis 23(3). Stipules ovate or semi-ovate, not early deciduous .................................. 24 23. Stipules linear, subulate, or setaceous, frequently early deciduous ...... 26 24(23). Stem with 4 prominently winged angles; petiole stout, caniculate above, with 3 pair of subsessile glands ...................................... P. quadrangularis 24. Stem terete or only slightly angled; stipules with apical arista or awn; petioles 2–8-glandular ......................................................................... 25 25(24). Stipules to 4 × 2 cm, with an apical awn ≥ 7 mm long, this normally oblanceolate-spatulate, widened and yellow or black near tip; floral bracts sessile (see also couplet 12) ...................................................... P. retipetala 25. Stipules to 2.5 × 1 cm, apically aristate; floral bracts petiolate (see also couplet 15) ................................................................................. P. picturata 26(23). Petiolar glands stipitate or auriculate, distinctly raised above surface .... 27 26. Petiolar glands wart-like and raised but not stipitate, or flat and platelike or scar-like .................................................................................... 29 27(26). Petiolar glands small, sessile to distinctly stipitate, at or above middle, (blades normally 3-lobed; see also couplet 18) .......................... P. suberosa 27. Petiolar glands auriculate or cupped ...................................................... 28 28(27). Petiolar glands lateral near base, large, sessile, auriculate; entire blades normally stiff-coriaceous (blade may have incipient lateral lobes); stipules filiform, curled (see also couplet 17) ......................... P. auriculata 28. Petiole with a pair of apical, short-stipitate, slightly cupped or auriculate glands; stipules oblanceolate to ligulate, medially serrulate (see also couplet 21) ................................................................................ P. seemannii 29(26). Petiolar glands wart-like, distinctly raised above surface (when basal may be low, or partially hidden by pubescence) ......................................... 30 29. Petiolar glands scar-like or patelliform, never basal .............................. 51 30(29). Petiolar glands near middle to near to base, but not basal .................... 31 30. Petiolar glands apical or basal ................................................................. 37 31(30). Leaves 3-veined from base ....................................................................... 32

636

P ASSIFLORACEAE

31. Leaves penniveined .................................................................................. 34 32(31). Blades normally unlobed; ovary globose, pericarp of fruit normally very hard (only one collection reported, probably cultivated) ...... P. maliformis 32. Blades normally 3-lobed; ovary narrowly ovoid or ellipsoid; pericarp of fruit coriaceous..................................................................................... 33 33(32). Stipules narrowly linear-oblong to -lanceolate, to 6 × 1.5 mm, denticulate above; base of leaf blade truncate or slightly cordate, tip obtuse to acuminate, rarely rounded; floral bracts obovate, subobtuse to acute, united at base for ca. 5 mm, margin serrulate (see also couplet 22) ................................................................................................... P. serrulata 33. Stipules linear-attenuate, to 20 × 3 mm, glandular-dentate; base of leaf blade cordate or subtruncate, tip acuminate; floral bracts ovate, longacuminate (> 1 cm), united at base for 1–2 cm, lower margin glandulartoothed (see also couplet 22) ................................................. P. multiformis 34(31). Hypanthium cylindric, longer than wide; flowers red or reddish; radii not banded; margin of leaf blade entire or undulate .................. P. glandulosa 34. Hypanthium broadly campanulate, equal or wider than long; flowers white to pinkish within; radii white banded with blue or violet; margin of leaf blade entire or minutely denticulate or serrulate ................... 35 35(34). Petiolar glands large, wart-like; flowers borne singly in the leaf axils; floral bracts 1.5–3.5 cm long, glandular-margined; sepals 2.5–3 cm long, relatively thin .................................................................. P. guazumaefolia 35. Petiolar glands low wart-like, nearly patelliform; flowers borne in racemes, pseudoracemes, or singly; floral bracts 3–6 cm long; sepals 4– 5 cm long, thick .................................................................................... 36 36(35). Flowers normally borne on short axillary branches, racemose or pseudoracemose; floral bracts obovate, glandular-margined; perianth pinkish white; sepals thick-fleshy; both series of radii thick and of equal length (see also couplet 51) ................................................. P. riparia 36. Flowers solitary in the axils or in a very short raceme; floral bracts ovate, glandless; perianth flushed with white or spotted with pink or purplish; sepals fleshy; outer series of radii more slender and half as long as inner ....................................................................................... P. ambigua 37(30). Petiolar glands basal or nearly so ........................................................... 38 37. Petiolar glands apical ............................................................................... 42 38(37). Leaf blade 3-veined at base, margin obviously toothed; flowers red ..... 39 38. Leaf blade penniveined, margin entire or shallowly toothed; flowers red or white ................................................................................................ 40 39(38). Floral bracts large, ovate, gland-toothed, brilliant red, 1–3 cm wide; blade simply or doubly serrate or doubly crenate, margin slightly revolute, with small glands on the teeth; plant densely rufo-tomentose ........ P. coccinea 39. Floral bracts linear to linear-lanceolate, < 5 mm wide, 2 or more large glands near base, finely glandular-serrate or nearly pectinate; blade (normally 3-lobed) uniformly or irregularly repand-serrate, the teeth minutely gland-tipped; plant glabrous, glabrescent or velutinous to densely whitish- to ferruginous-tomentulose (see also couplet 19) ..................................................................................... P. quadriglandulosa

Passiflora 637

40(38). Hypanthium cylindric, longer than wide; flower scarlet red or reddish; petiolar glands below middle to near base; margin of leaf blade entire or undulate (see also couplet 34) ........................................... P. glandulosa 40. Hypanthium shorter than, to as long as, wide; flower red or whitish; petiolar glands basal; leaf blade margin entire or undulate-dentate ...... 41 41(40). Leaf blade membranous to coriaceous, entire; sepals with 2–5 blackish green marginal glands; peduncles stout, 2–8 cm long; flower red; radii 8–12 mm high, filamentous in upper 1/4–1/2 .............................. P. variolata 41. Leaf blade ± stiff-coriaceous, undulate-dentate, the teeth shallow and distant; peduncles very stout, 10–18 cm long; flower whitish; radii 8– 14 mm long, membranous in basal 2/3 ..................................... P. amicorum 42(37). Floral bracts large; radii in 2 series of ligulate, subulate, linear, or capillary filaments, 2–5 cm long, mostly with concentric bands of red to blue; hypanthium short-campanulate; flowers large (5–11 cm diameter) ...................................................................................................... 43 42. Floral bracts mostly minute; radii in 1 or 2 series of usually compressed, nonlinear filaments, mostly < 2 cm long; hypanthium cylindric to cylindric-campanulate; flowers smaller, less showy ................................... 48 43(42). Leaf blade toothed .................................................................................... 44 43. Leaf blade entire ...................................................................................... 46 44(43). Stipules setaceous, < 1 mm wide; leaf blade subentire to shallowly glandular-serrulate-dentate; radii capillary; pali in 1–3 series; ovary finely ferruginous-velutinous .............................................................. P. gleasonii 44. Stipules linear-subulate, ≥ 1 mm wide; leaf blade serrate; radii thick; pali in several series; ovary glabrous ......................................................... 45 45(44). Flowers pendulous; leaf blade ovate-oblong to elliptic, or broadly ovate to obovate, 7–17 × 3–10 cm, base rounded, apex acute to acuminate, margin variously serrate to subentire, usually drying blackish or dark silvery gray; floral bracts oblong-ovate, 3.5–4.7 × 1.5–2.5 cm, margin large-glandular near base; fruit ovoid-ellipsoid, 3–9 × 3–7 cm, bright yellow to orange-yellow; native to riversides, occasionally cultivated ........................................................................................................ P. nitida 45. Flowers erect; leaf blade (normally 3-lobed) ovate; stipules linear-subulate, ca. 1 cm long, entire or minutely glandular-serrate; floral bracts 10–25 × 7–15 mm, glandular-serrate to pectinate; fruits globose to oblong-ellipsoid, 4–10 cm long, light yellow to purplish; cultivated and escaped (see also couplet 20) ..........................................................P. edulis 46(43). Leaf blade basically lanceolate, apex acute to acuminate; radii in 2 ± equal series, narrow-linear with filiform, crisped tips, banded with lavender; perianth white ............................................................. P. acuminata 46. Leaf blade ovate to oblong, apex subacute to rounded; radii in 2 unequal series, banded dark red to purple or blue; perianth reddish or white spotted with red ................................................................................... 47 47(46). Leaf blade ovate to oblong, length normally < 2 times the width, apex subacute to obtuse, blade dark green ............................................ P. laurifolia 47. Leaf blade narrowly oblong, length normally > 2 times the width, apex rounded, often emarginate, margin slightly revolute, entire or minutely

638

P ASSIFLORACEAE

indented-glandular, blade medium to light green ........... P. capparidifolia 48(42). Flowers solitary or paired in leaf axils; petiolar glands shallowly wartlike; perianth light green (see also couplet 56) .............................. P. ovata 48. Inflorescence fasciculate or racemose ..................................................... 49 49(48). Inflorescence of horizontal racemes 8–60 cm long; leaf blade lance-oblong, ovate-lanceolate, ovate, or rarely ovate-orbicular, thick-coriaceous, the lower surface densely glaucous ........................................ P. longiracemosa 49. Inflorescence fasciculate, or racemose on a short rachis to 2.5 cm long; leaf blade various ........................................................................................ 50 50(49). Petiolar glands low, wart-like, nearly patelliform, under small subpeltate base of blade; flowers white to light yellowish green; sepals 20–25 mm long; operculum 10–13 mm long, reaching to throat of floral tube; leaf blade ovate to elliptic (see also couplet 56) ............................. P. cauliflora 50. Petiolar glands wart-like, at apex of petiole and extending under blade margin on midrib; flowers orange-yellow to deep pink; sepals 10– 20 mm long; operculum at base of floral tube, 2.5 mm long; leaf blade ovate to suborbicular, the lower surface glaucous ............... P. fuchsiiflora 51(29). Petiolar glands at middle of petiole, low, wart-like, biglandular, nearly patelliform; radii 4–5 cm long, banded dark red; flowers large, pendent; floral bracts large; perianth white to purplish pink (see also couplet 36) ....................................................................................................... P. riparia 51. Petiolar glands at apex of petiole or base of midvein; radii shorter, at most bicolored ............................................................................................... 52 52(51) Petiolar glands at apex of petiole ............................................................ 53 52. Petiolar glands at very base of midvein or under a slightly decurrent margin of leaf blade .................................................................................... 57 53(52). Leaf blade stiffly thick-coriaceous, the lower surface with strongly elevated veins ......................................................................................... 54 53. Leaf blade coriaceous to subcoriaceous ................................................... 56 54(53). Petiolar glands hidden by subpeltate base of blade; blade broadly elliptic, obovate, or narrowly oblong-ovate, 5–25 × 4–16 cm, base cordulate or rounded, lower surface conspicuously reticulate and finely olive-puberulent; radii in one series 15–30 mm long; pali in 5 or 6 series ...... P. costata 54. Petiolar glands visible at apex as continuation of slightly decurrent margin of blade ........................................................................................... 55 55(54). Leaf blade suborbicular, broadly ovate, obovate, oblong, elliptic, or lanceolate, 3–11 × 2–11 cm, base rounded or obtuse, very prominently reticulate to the smallest veins, the lower surface densely to sparsely yellow to yellow-tan villous-scabrous; perianth white, spotted with pale lavender; radii in one series 10–20 mm long; pali in one series; 600– 2200 m elevation ................................................................... P. sclerophylla 55. Leaf blade lanceolate to elliptic-ovate, stiff- to thick-coriaceous, 3.5–11 × 2–6 cm, veins elevated and prominent on both surfaces, at most minutely puberulous on the lower surface; flowers light green and cream to salmon; hypanthium 16–18 × 6–7 mm; radii in one series 8–11 mm long; pali in one scanty series; 100–200 m elevation, in white-sand scrub (bana) ........................................................................... P. phaeocaula 56(53). Petiolar glands shallowly wart-like, apical; flowers solitary or paired in leaf axils; perianth light green; sepals to 20 mm long; radii in one series

Passiflora 639

56.

57(52). 57. 58(57). 58. 59(58).

59.

60(58).

60.

61(57).

61. 62(61).

62. 63(62).

ca. 10 mm long; pali in 1 series; leaves oblong (see also couplet 48) ......................................................................................................... P. ovata Petiolar glands low, wart-like, nearly patelliform, under small subpeltate base of blade; flowers white to light yellowish green; sepals 20–25 mm long; radii in 2 series, the outer 12–28 mm long; pali in 1–3 series, the inner decurved into throat of floral tube; leaf blade ovate to elliptic (see also couplet 50) ........................................................................ P. cauliflora Glands at petiole apex under decurrent margin of blade ....................... 58 Glands on base of midrib, extending onto blade base and/or under a slightly decurrent margin .................................................................... 61 Margin of leaf blade normally shallowly crenate or dentate .................. 59 Margin of leaf blade entire ...................................................................... 60 Petiolar glands under slightly decurrent margin of leaf blade; blade normally shallowly rounded-dentate, at least in upper half, the teeth tipped with a minute marginal gland, infrequently undulate; 50–600 m elevation, in open forests .......................................................P. pyrrhantha Petiolar glands wide, the border a continuation of the leaf blade margin, appearing as a small basal lobe; blade coarsely and shallowly crenatedentate, with 0.5 mm marginal glands at the apices; 100–200 m elevation, in white-sand scrub (bana) ..................................................... P. sp. C Leaf blade lanceolate-oblong to lanceolate, base rounded or broadly obtuse, apex obtuse to rounded or subacute; peduncles 1–1.5 mm long, axillary or in a raceme, paired in the axils of recurved, subulate bracts ca. 6 mm long, biglandular at middle, on a few- to many-flowered rachis 1.5–8 cm long; hypanthium longer than perianth segments; perianth reddish orange .................................................................. P. nuriensis Leaf blade ovate to elliptic, base obtuse to subcordate, apex acuminateattenuate to shortly acute; peduncles 2–3 mm long, densely fasciculate on a rachis ca. 1 cm long; hypanthium shorter than perianth segments; perianth greenish white to cream ............................................. P. maguirei Flowers white to greenish white, solitary or in pairs; leaf blade coriaceous, oblong-lanceolate, oblong, or slightly obovate, 7–10 × 3–5.5 cm, apex rounded or rounded-acute; hypanthium ca. 10 mm long, tube cylindric to cylindric-campanulate ........................................... P. cardonae Flowers red to orange, in racemes or pseudoracemes ............................ 62 Leaf blade usually with a thick, dark margin, normally oblong, but also ovate to lanceolate, base rounded, infrequently subcordate, apex most often emarginate, but also truncate, rounded, obtuse, or acute; peduncles paired, 1–5 mm long, in a pseudoraceme 5–50 cm long, with reduced leaves, or petioles reduced to spines with 2 basal glands, or on each side of a tendril reduced to a spine, infrequently in axils of normal leaves with tendrils; inflorescence many-flowered ................ P. securiclata Leaf blade lacking dark margin, mainly ovate to elliptic, apex acuminate .............................................................................................................. 63 Leaf blade coriaceous, entire, barely very shallowly crenate, with minute marginal glands in the sinuses, oblong, ovate, broadly elliptic, or obovate; peduncles solitary or in pairs in slender racemes or pseudoracemes to 25 cm long; hypanthium 30–40 mm long, tube cylindricconical; sepals 10 × 3 mm ............................................................ P. spinosa

640

63.

P ASSIFLORACEAE

Leaf blade subcoriaceous, entire, slightly revolute, ovate to ovate-lanceolate, reticulations prominent, 12–15 × 6–8 cm, apex caudate-acuminate; peduncles in pairs in apical half of a pseudoraceme 8–30 cm long; hypanthium 28–50 mm long, tube cylindric; sepals 15–20 × 3–5 mm .......................................................................................................... P. holtii

Passiflora acuminata DC., Prodr. 3: 328. 1828. Plant essentially glabrous; stipules narrow-linear, 4–8 mm long, falcate, broadened above, deciduous; petiole 1–1.5 cm long, prominently wart-like/biglandular at apex, glands ca. 2 mm diameter; blade lanceolate, lanceolate-ovate, or oblong-lanceolate, 7–16 × 2–8 cm, base acute, rounded, or subcordate, entire, sublustrous, membranous, or subcoriaceous; peduncle 1–5 cm long; pedicel 3–7 mm long; flowers pendent, 5–6 cm wide, pale blue, lavender, or pinkish; floral cup urceolate, 5 mm high, umbilicate, bracts narrowly elliptic to oblong, 1.5–4 × 0.5–1.5 cm, entire or glandular, floral tube short-campanulate, 5–8 mm long; sepals narrow-lanceolate, 20–30 × 6–10 mm, whitish green with purple spots, corniculate just below apex; petals similar, ca. 15 mm long, white or lavender; radii in 2 series 2.5–4 cm long, banded; pali in 3 or 4 series, the 2 or 3 outer few, setaceous < 1–2 mm long, innermost subulate, 5–7 mm long, banded violet on white; operculum incurved, 4–5 mm high, fimbrillate; ovary ovoid, minutely puberulent; fruit not seen. Evergreen lowland to lower montane forests, 100–400 m; Amazonas (Río Mawarinuma, San Carlos de Río Negro). Apure, Táchira; Colombia, Guyana, French Guiana, Brazil. Passiflora adenopoda DC., Prodr. 3: 330. 1828. —Parchita negra. Plant moderately velutinous and strigose, stem becoming large and woody; stipules 10– 15 × 7–15 mm, obliquely semi-ovate, entire to deeply palmatisect and cleft; petiole 3–13 cm long, the glands 1–1.5 cm below apex; blade 3-lobed to the middle or more, with lateral lobes slightly lobed, to 5-lobed, 7–17 × 8–18 cm, base deeply cordate, lobes ovate, narrowed at the base, the middle longer, apex acuminate, margin remotely shallowserrate; peduncles solitary or in pairs, 20–30 mm long; bracts deeply lacerate, 5–10 × 2–6

mm; pedicel 2 mm long; flowers 2–7 cm wide; floral cup saucered, 3–4 mm high, 10–14 mm diameter; floral tube a mere rim; sepals oblong-lanceolate, 15–40 × 6–10 mm, pale green or yellowish, apex cucullate, with a corniculus ca. 1 cm long; petals linear-lanceolate, shorter and narrower, white; radii in one series, filiform, 5–18 mm long, white basally with 2 purple bands, apically yellow; pali absent; operculum plicate, incurved; ovary subglobose to oblong, densely brownvelutinous; fruit globose to ovoid, 2–7 × 2–4.5 cm. Mainly in disturbed areas, 1500–1800 m; Amazonas (slope of peak near Cerro Yutajé). Aragua, Distrito Federal, Táchira, Vargas, Zulia; Mexico, Central America, Colombia, Ecuador, Peru. Passiflora ambigua Hemsl., Bot. Mag. 128: t. 7822. 1902. Plant essentially glabrous; stipules filiform, 5–8 mm long, deciduous; petiole 2–4 cm long, glands ± wart-like, at or below middle; blade oblong, ovate-oblong, or oblong-lanceolate, 10–20 × 5–10 cm, base rounded or cuneate, apex abruptly acuminate and mucronulate, lustrous; peduncles 4–7 cm long; bracts 3–6 × 3–4 cm, concave, entire, glabrous or short-puberulent without; pedicels 2–3 mm long; flowers 8–12 cm wide, pendent; floral cup ca. 3.5 mm high, urceolate, umbilicate; floral tube 8–10 mm high, cylindric-campanulate; sepals linearoblong to ovate, 4–5.5 × 1.2–2 cm, obtuse or rounded, short corniculate below apex, white externally, glabrous or finely short-puberulent; petals linear-lanceolate to lance-ovate, 3–4.8 × 0.6–1.3 cm, apex rounded to acute, membranous, white dotted with rose purple; radii in 2 series, outer slender, 1–1.5 cm long, banded white on red, inner coarser, 3.5–5 cm long, obtuse, banded white on violet; pali in ca. 3 indefinite series, capillary, simple, or capitellate, 0.5–2 mm long; operculum horizontal, margin recurved, crenulate; ovary narrowly ovoid, densely brown- to olive-to-

Passiflora 641

mentose; fruit globose to ovoid, 7–12 × 4–7 cm, greenish yellow, pericarp ca. 1 cm thick. Montane forests, ca. 800–900 m; Amazonas (Simarawochi). Aragua, Distrito Federal, Falcón, Lara, Miranda, Táchira, Vargas; Panama, Ecuador, Peru, Brazil. ◆Fig. 551. Passiflora amicorum Wurdack, Bol. Soc. Venez. Ci. Nat. 26: 429. 1966. Woody vine, densely puberulous; stipules 5–7 × 0.5 mm, linear-acute, early deciduous; petiole 0.5–1.5 cm long, wart-like-biglandular at the base, glands ca. 1 mm diameter; blade 4–10 × 2–6 cm, oblong-elliptic to broadly elliptic, base obtuse or rounded, apex broadly acute or obtuse, aristulate, slightly revolute, with few, minute, submarginal glands; peduncles solitary; pedicels 1.2–2 cm long; bracts subverticillate, 7–10 × ca. 1.5 mm, oblong-subulate, with 1 or 2 pair of large glands; flowers to 16 cm wide, whitish; floral cup urceolate, umbilicate, ca. 5 × 12 mm; floral tube reduced to a rim; sepals ca. 8 × 1 cm, narrow-oblong, corniculus 6–10 mm long, margin with several glands 0.5–1 mm diameter; petals slightly shorter, narrower, and thinner; radii in 1 series, 8–14 mm long, basal 6–10 mm membranous, remainder filamentous, salmon-brown to purplish; pali in 1 series of laciniate filaments to 4 mm long, lower 1/2 united into a membrane; operculum dependent, ca. 2 mm long, laciniate; ovary ovoid, densely light-puberulous; fruit broadly ellipsoid, 5–6 × ca. 3.5 cm, pericarp ca. 2 mm thick, hard, brittle. Open and disturbed areas, forest edges, 1200–1400 m; Bolívar (Kavanayén, Luepa). Endemic. Collectors of this species described it as “anthesis nocturnal, with fragrance like gin.” Considering this and the very stout peduncles, it is probably pollinated by bats. The fruits are edible. Passiflora auriculata H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 131. 1817. Young branches puberulent, mature plant glabrous to densely pubescent; stipules filiform, 0.5–4 mm long, curled, early deciduous; petiole 0.5–30 mm long, puberulent, glands with dark center and light rim; blade lanceolate, ovate-lanceolate, oblong-lanceolate, elliptic, ovate, or nearly orbicular, 2.5–16 × 0.5–15 cm, (1)3(5)-veined from base,

entire to slightly or amply 3-lobed and variable on same plant, rounded or subcordate to barely subpeltate (juvenile) at base, apex acuminate to rounded, apiculate, coriaceous to membranous, ocellae small, scattered, normally with dark center and raised light rim; peduncles paired, 5–25 mm long; bracts deltoid to subulate, nearly verticillate, basal to near middle of peduncle, 1–2 mm long, deciduous, rarely absent; pedicel 1–3.5 mm long; flowers 2–3.5 cm diameter, white to yellow-green to purplish, normally mast-flowering; sepals oblong-lanceolate, 9–15 × 2–4 mm, acute; petals narrow-linear, very delicate, 7 × 1 mm; radii in 1 series, filiform, radiate, 8–13 mm long, purplish at base, above yellowish green, tips crisped; pali in 1 series, capillary, < 3 mm long, capitellate, white; operculum plicate; ovary ovoid, pilosulous to tomentose; fruit globose, 10–17 mm diameter, glabrous to densely pilosulous, greenish yellow to purplish black, the peduncles crozierform. Savannas, shrublands, gallery forests, evergreen lowland to montane forests, secondary vegetation, 50–1300 m (to 1800 m on slopes of some tepuis); Delta Amacuro (Caño Capurito in Caño Capure basin, Río Amacuro, Río Cuyubini), Bolívar (widespread), Amazonas (widespread at low elevations). Apure, Barinas, Falcón, Lara, Mérida, Portuguesa, Táchira, Zulia; Nicaragua to Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 553. The size and shape of the leaf blade can be extremely variable. In forested areas, blades are typically large and wide, and membranous to subcoriaceous. In full sun, as in some Gran Sabana plants, blades are narrow and entire, coriaceous, with few ocellae, and revolute margins. Passiflora capparidifolia Killip, J. Wash. Acad. Sci. 14: 113. 1924. —Parcha, Parcha de culebra. Plant glabrous to minutely puberulous, the hairs transparent; stipules 4–8 × 0.5–1 mm, narrowly linear-ligulate, widened and frequently minutely glandular-margined above, then acute, subcoriaceous; petiole 7– 20 mm long, canaliculate above, with 2 large wart-like glands at apex or to 1 mm below base of blade; blade narrowly oblong, rarely

642

P ASSIFLORACEAE

elliptic or elliptic-obovoid, 7–14 × 2–6.5 cm, base rounded or subcuneate, very shortly decurrent, mucronulate, reticulate-veined, coriaceous; peduncles 2.5–8 cm long, slender; bracts 3–5 × 1.5–3 cm, broadly ovate, margin crenulate with large glands toward the obtuse apex; pedicel to 3–8 mm long; flowers 8– 10 cm diameter, floral cup urceolate, 3–6 mm high, deeply umbilicate; floral tube campanulate, 3–5 mm long; sepals lanceolate, 3.5–4 × 1 cm, apex obtuse, narrow keel with corniculus 2 mm long, without greenish white, within white or red-brown; petals to 3–3.2 × 1.5 cm, obtuse, white, pinkish red, or purplish; radii in 2 series, the outer filiform, 1.2–2 cm long, second series coarser, compressed and to 2 mm wide radially, 3.5–4 cm long, both series banded purple-red below, blue to bluish purple above; pali in 4 or more series of tubercles or threads 0.2–1 mm long, increasing down wall of floral tube; operculum membranous, horizontal to top of 1 mm high limen at base of narrow trochlea; staminal filaments purple-spotted; ovary ellipsoid, finely white- or tan-sericeous; fruit oblongovoid, finely pilose, glaucous dull lavender with dark green stripes. Forests, forest borders, secondary vegetation, 50–1200 m; Bolívar (Caicara, Isla Anacoco, La Escalera, Río Botanamo, Río Caroní, lower Río Caura, Río Chiguao, Río Cuyuní, Río Paragua, Río Parguaza, Río Suapure, Salto Pará, San Felix, Santa Elena de Uairén). Guyana, Suriname, northern Brazil. ◆Fig. 552. Passiflora cardonae Killip, Brittonia 3: 172. 1939. Plant suffrutescent, subscandent; stipules early deciduous, petiole 8–10 mm long, slender, glands scar-like at very base of midrib and part of blade, scarcely visible; blade entire, base rounded, lustrous, prominently finely reticulate on lower surface; peduncles solitary or in pairs, 1–5 mm long, occasionally on an extended branch with tendrils, but with the subtending leaves early deciduous; bracts triangular, ca. 0.5 mm long, pedicels 7–10 mm long; floral cup 3 mm high × ca. 3 mm diameter; floral tube ca. 6 mm long, 3–4 mm diameter; sepals linear, ca. 25 × 2–3 mm, obtuse; petals linear, 15 × 2 mm, obtuse; radii in 1 series, ligulate, 10–12 mm long, somewhat cultrate, yellowish orange; pali in

2 series, the outer filiform, ca. 3 mm long, the inner capillary, 1 mm long; operculum ca. 11 mm long, exserted, reddish; ovary ovoid, trigonous, purplish, densely gray-tomentulose, styles arising at the tops of the angles; fruit not seen. Montane shrublands, forest edges, 900–2300 m; Bolívar (Auyán-tepui, Cerro Roraima, Cerro Venado, Ptari-tepui). Endemic. Passiflora cauliflora Harms, Verh. Bot. Vereins Prov. Brandenburg 48: 185. 1907. Liana or scandent shrub, glabrous; stipules early deciduous; petiole 1–3 cm long; blade 13–30 × 5.5–10 cm, oblong, ovate, broadly ovate-elliptic, base rounded or slightly cordate, apex obtuse, acute, or slightly acuminate, entire, thick-coriaceous; peduncles 1–3 mm long, in pairs fasciculate on a rachis to 5–15 mm long and frequently subtended by a large-glanded bract 3–4 mm long, or in a raceme to 3 cm long, infrequently in pairs in axils of normal leaves on a short branch, bracts deltoid, 1 mm long, at middle of peduncle; pedicels 3–10 mm long; flowers with faint, unpleasant odor; floral cup 6–12 mm long, urceolate-tubular, 10ribbed externally; floral tube 7–12 mm long, 3–4 mm diameter; sepals oblong, 20–35 × 6– 8 mm, obtuse to acute, petals thinner and slightly shorter, both reflexed in anthesis; radii in 2 series, yellow, outer subdolabriform, 12–28 mm long, apex slightly attenuate, inner broadly linear to linear-subulate or slightly dolabriform, 3–6 mm long; pali in 1– 3 series, pink, linear, the outer erect or inclined horizontally inward, 2–3 mm long, the inner 1 or 2 decurved into throat, 2–4 mm long; operculum tubular, cleft above into 5 linear segments 2–5 mm long, ending above or at the throat of the floral tube; ovary narrowly obovoid, densely ferruginous-tomentulose; fruit narrowly ellipsoid, ca. 5–7 × 2.5–4 cm, sparsely tomentulose. Gallery forests, white-sand scrub (bana), 100–200 m; Bolívar (Río Paragua, Río Parguaza), Amazonas (Carmelitas, Cerro Yutajé, Coromoto, Galipero, Isla Ratón, Maroa, Río Atabapo, Río Cataniapo, Río Cuao, San Juan de Manapiare, Santa Barbara del Orinoco). Colombia, Peru, Brazil.

Passiflora 643

Passiflora cirrhiflora Juss., Ann. Mus. Natl. Hist. Nat. 6: 115, t. 41, fig. 2. 1805. Plant herbaceous; stipules setaceous to subulate, to 5 mm long; petioles to 10 cm long; blade to 10 × 12 cm, petiolules narrow, to 1.3 cm long, leaflets entire, oblong to broadly elliptic, the base rounded to acute, not attenuate, apex rounded to aristate; common peduncle solitary, 1–3 cm long, stout, 2flowered at apex, with a stout tendril between; peduncles 2–5 cm long; bracts linearsubulate, to 1 cm long; flowers to 8 cm diameter, erect; sepals oblong, ca. 30 × 8 mm, white; petals longer and narrower, yellowish or reddish; radii uniseriate, to 3 cm long, thick, curved upward, zigzag in upper 1/2 and verrucose along the margin, yellow, red at base; pali in 2 series, narrowly linear, to 1 cm long, capitate; ovary densely short-velutinous; fruit globose, pericarp coriaceous. Secondary vegetation at forest edges, 400– 500 m; Bolívar (Piedra de la Virgen at La Escalera). Guyana, Suriname, French Guiana.

ral cup urceolate, umbilicate, 6 × 8 mm; floral tube slightly campanulate, 2–8 × 8 mm; sepals linear-lanceolate, 30–50 × 8–10 mm, cucullate, carinate, with corniculus to 13 mm long; petals linear, 35–40 × 7–8 mm; radii in 2 series of subulate filaments, ca. 1 cm long, white to light pink in lower 1/2, deep red to purple above; pali in 1 series of filaments 6–8 mm long, white, united into a membrane basally; operculum dependent, recurved and short-filamentous at margin; ovary ovoid, densely yellow-tomentose or -sericeous; fruit ovoid or subglobose, 5–7 × 3–4 cm, orange or yellow, exocarp brittle, densely and finely tomentulose. Savannas, shrublands, low forests, open and disturbed areas in full sun, 50–1300 m; Delta Amacuro (Río Amacuro), Bolívar (widespread), Amazonas (widespread). Southern Colombia, Guyana, Suriname, French Guiana, Amazonian Peru, Brazil, Bolivia. This species has striking red flowers and is cultivated as an ornamental in many countries. The fruits are edible.

Passiflora coccinea Aubl., Hist. Pl. Guiane 828, t. 324. 1775. —Poró (Yanomami), Proyek (Taulipáng), Shakrakarimi (Yanomami). Subligneous vine, plant nearly glabrous to finely white- or yellow-puberulent, or more frequently rufo-puberulent to densely rufo-tomentose; stipules narrowly linear or linear-setaceous, 4–10 mm long, to lanceolate, 20 × 5 mm, entire or minutely glandular-serrulate; petiole to 3.5 cm long, sessile with two sessile wart-like glands at or near base, barely visible through the pubescence, rarely eglandular; blade ovate-oblong to rarely nearly orbicular, 6–20 × 3–11 cm, 3veined, base cordate to subcordate-rounded, apex acute and ± apiculate, to subobtuse or rounded, upper surface glabrescent and tomentulose to rufo-tomentose on veins, lower surface rufo- or cano-tomentose; peduncles stout, 5–12 cm long; bracts 3–8 × 1.5–5 cm, apex acute to rounded, crenate or sharply serrate, margin with dark glands (ca. 1 mm diameter), ferruginous-tomentose without, puberulent within, may have large, flat, pubescent glands at or very near base; pedicel 3–6 mm long; flowers 7–9 cm wide, brilliant red, scarlet, or reddish orange; flo-

Passiflora costata Mast. in Mart., Fl. Bras. 13(1): 573. 1872. —Parcha. Subshrubby vine or shrubby liana, tendrils stout; stipules early deciduous; petiole 1.5–4 cm long, with 2 dark scar-like glands at very apex, elliptic, 2–3+ mm long; blade olive green, apex rounded, emarginate to subacute; peduncles solitary or in pairs, 1– 3.5 cm long; bracts narrow-linear, 1–2 mm long, scattered ± at middle of peduncle; pedicel 5–12 mm long; flowers white, 6–8 cm wide, finely puberulous externally, fragrant; floral cup 1.5–5 mm tall, campanulate, not umbilicate; floral tube broadly funnelform, 3–4 mm long, 9–10 mm wide at throat, where it forms a narrow introrse horizontal ridge; sepals oblong-lanceolate, elliptic, narrowly obovate, or oblong-spatulate, 20–38 × 9–17 mm, slightly cupped at apex; petals oblongspatulate, 15–36 × 5–10 mm, obtuse; radii in 1 series, 15–30 mm long, subdolabriform, laterally compressed 2–3 mm wide and lobed along central margin, base orange-yellow, reddish at middle, tip long-attenuate, verrucose and curled, golden yellow; pali in ca. 5 or 6 series, minutely papillate, narrowly liguliform, decreasing from 3–5 to 0.3–1 mm long, the outermost spatulate-broadened at

644

P ASSIFLORACEAE

base, yellowish to wine-red; operculum a 2 mm horizontal membrane, the margin curved upward and faintly denticulate, appressed to androgynophore below the large trochlea at mouth of tube; ovary narrowly obovoid, sulcate, densely yellowish shortvelutinous, the stout styles arising from the top of the angles; fruit 3.5–16 × 2.5–6 cm, bluntly ellipsoid to obovoid, hard, thickwalled. Mostly in seasonally flooded forests, 100–300 m; Bolívar (Salto Pará), Amazonas (Río Cataniapo, Samariapo, San Carlos de Río Negro). Guárico; Guyana, French Guiana, Ecuador, Peru, Brazil. Passiflora edulis Sims, Bot. Mag. 45: pl. 1989. 1818. —Maracuyá, Parcha, Parchita. Mainly herbaceous, becoming woody in old stems; stipules linear-subulate, 4–14 × 1–1.5 mm, deciduous; petiole 1.5–4 cm long, glands apical at end of ± decurrent blade margin, wart-like, sessile or slightly raised, oblong, ca. 2 mm long; blade ovate, 6–15 × 6– 16 cm, glandular-serrulate, base cordate, lobes ovate-elliptic, the central much narrowed at base, apices acute (normally only the juvenile simple or mitten-shaped, but occasionally also on mature stems); peduncles solitary, stout, 4–6 cm long; bracts ovate, enveloping floral cup but not the entire bud; pedicel 1–10 mm long; flowers to 9 cm wide; floral cup 5–8 mm high, deeply umbilicate; floral tube 5–8 mm long, funnelform, blocked by the massive 5-lobed trochlea; sepals oblong, to 3.5 cm long, green without, slightly keeled and corniculate, the exterior margins frequently with a dark green gland at about the upper 1/3 similar to that of the petiole (the 5 glands easily seen in the bud), white within; petals smaller, obtuse, white; radii in 2 series, to 3.5 cm long, crisped at apex, white, atropurpureous in basal 1/3–2/3; pali atropurpureous, of 2 series to 2.5 mm long, at mouth of floral tube, the surface of the tube smooth or with numerous atropurpureous tubercles diminishing toward the base (or merely spotted), where there may be 1 or 2 series 0.2–4 mm long; operculum 1–5 mm tall, minutely plicate and atropurpureous tuberculate above, ascendent toward the cup-shaped limen inserted at the base of the trochlea; ovary ovoid or glo-

bose, glabrous, the fruit similar, greenish yellow to purple, 4–5 cm long (or to 10 cm in cultivated forms, mostly yellow), epicarp coriaceous, mesocarp spongy, to 1 cm thick. Usually in or near cultivated plots, sea level to 1200 m; grown throughout the flora area as a cultivar, or escaped and more or less naturalized. Widespread in cultivation throughout rest of Venezuela and tropical and subtropical countries in general, probably native to Brazil. ◆Fig. 556. Passiflora edulis Sims var. edulis, with purple to greenish yellow fruit (Parchita morada), generally sweet aril, and smaller flowers and fruits, is naturalized above 1000 m in the Andes and northern coastal ranges of Venezuela. Passiflora edulis f. flavicarpa Degener is the commercial variety, apparently originated in Brazil, originally with yellow fruit and acidic aril, now with numerous cultivars of different size and color of fruit, cultivated for juice, sherbet, ice cream, marmalades, and preserves. Passiflora foetida L., Sp. Pl. 959. 1753. Plant herbaceous; stipules deeply cleft into filiform divisions or even pinnatisect, the divisions tipped with glandular hairs, the base semicircling the stem; petiole to 6 cm long, glandular-hairy; blade very variable (but uniform on same plant), 1.5–15 × 1.3–12 cm, (subentire to) 3- or 5-lobed, base usually cordate; peduncles solitary, to 6 cm long; bracts involucrate, usually large and 2–4pinnatifid or -pinnatisect, the segments filiform and gland-tipped; pedicel 1–2 mm long; flowers 2–6 cm diameter, white, blue, purplish, or pink; sepals ovate-oblong, ovatelanceolate, or lanceolate-elliptic, with a corniculus below the apex; petals nearly as long as sepals, spatulate to oblanceolate; radii in 2 series, filiform, ca. 1 cm long, reflexed; pali in 3 or more series, 1–2 mm long; operculum a short erect membrane, limen a large somewhat funnelform cup; ovary globose, glabrous or pubescent; fruit globose to subglobose, 1.5–4 cm diameter, yellow to red, the pericarp thin, often brittle. Mostly in open vegetation, weedy and invasive in disturbed areas, near sea level to 2000 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Throughout the rest of Venezuela; southern U.S.A. throughout the

Passiflora 645

Neotropics to Chile and Argentina, introduced in the Paleotropics; 38 varieties, 5 of these in the flora area. This species comprises a large complex that is complicated by human disturbances and the transport of seeds. The fruits are edible, normally with a musky odor and taste, but much appreciated by children. Key to the Varieties of P. foetida 1. Ovary and fruit glabrous ........................ 2 1. Ovary pubescent, fruit sparingly to densely so .............................................. 3 2. Stem hispid, hairs 3–4 mm long; leaves hispid-hirsute; styles long-hispid; bracts very large, 3- or 4-pinnatisect, segments interwoven, glabrous ...... var. hispida 2. Plant glabrous, infrequently with short glandular hairs on lower surface of leaf blade and on petiole, bracts smaller, 2- or 3-pinnatisect, segments straight, sparse, not interwoven .......... var. orinocensis 3. Stem, petiole, and peduncle yellow to brownish hirsute, hairs > 1.5 mm long; bracts 2- or 3-pinnatisect ... var. foetida 3. Stem, petiole, and peduncle pilosulous, the hairs < 1.5 mm long ....................... 4 4. Bracts 2- or 3-pinnatisect, segments longer than rachis, hairs yellowish or brownish ................. var. gossypiifolia 4. Bracts very narrow, mostly 1-pinnatifid, the segments short; stem white-villosulous ............................ var. moritziana foetida var. foetida. —Chamuré (Piaroa). Bolívar (Altiplanicie de Nuria, slopes of Auyán-tepui, Caicara, La Tigrera, Las Nieves, Río Aro, lower Río Caroní, Río Grande, San Felix, Tumeremo, Túriba, Upata), Amazonas (Caño Mosquito, El Burro, Galipero, La Esmeralda, Puerto Ayacucho, Río Coromoto, Río Mavaca, Río Ventuari, San Antonio, San Fernando de Atabapo, San Juan de Manapiare, San Simón de Cocuy, Sierra Parima, Trapichote). Apure, Carabobo, Falcón, Portuguesa, Yaracuy, Zulia; Puerto Rico, Jamaica, Lesser Antilles, South America north of Argentina. ◆Fig. 561.

27: 631. 1871. —Passiflora gossypifolia Desv. ex Ham., Prodr. Pl. Ind. Occid. 48. 1825. Delta Amacuro (Tucupita), Bolívar (Guri). Anzoátegui, Apure, Guárico, Miranda, Portuguesa, Sucre, Zulia; U.S.A. (Texas) south to Chile. P. foetida var. moritziana (Planch.) Killip in Pulle, Fl. Suriname 3(1): 318. 1937. —Passiflora moritiziana Planch. in Triana & Planch., Ann. Sci. Nat. Bot. sér. 5, 17: 175. 1873. Bolívar (Altiplanicie de Nuria, Ciudad Bolívar, El Cristo, La Paragua). Apure, Aragua, Carabobo, Distrito Federal, Falcón, Lara, Miranda, Nueva Esparta, Portuguesa, Sucre, Zulia; Curaçao, Colombia, Suriname. P. foetida var. hispida (DC. ex Triana & Planch.) Killip ex Gleason, Bull. Torrey Bot. Club 58: 408. 1931. —Passiflora hispida DC. ex Triana & Planch., Ann. Sci. Nat. Bot. sér. 5, 17: 172. 1873. Delta Amacuro (Río Amacuro, Río Cuyubini, Tucupita), Bolívar (Casa Blanca, Río Cuyuni, San Martín de Turumbán, Tumeremo), Amazonas (El Mango, Mavaca, Piedra Culimacare, Río Cunucunuma, San Carlos de Río Negro, San Fernando de Atabapo, Yavita). Aragua, Distrito Federal, Lara, Mérida, Miranda, Portuguesa, Sucre, Trujillo, Zulia; West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil.

P.

P. foetida var. gossypiifolia (Desv. ex Ham.) Mast., Trans. Linn. Soc. London

P. foetida var. orinocensis Killip, Gentes Herb. 2: 205, fig. 107. 1930. Delta Amacuro (Tucupita), Bolívar (Caicara, Ciudad Bolívar), Amazonas (Puerto Ayacucho, Río Parhuaza). Apure, Barinas. Passiflora fuchsiiflora Hemsl. in Hook., Icon. Pl. 26: pl. 2553. 1898. — tuHaemaereka hu (Piaroa), Oroshomi thotho (Yanomami). Large liana, glabrous; stipules linear, 7–9 × 1.5 mm, acuminate, early deciduous; petiole to 8 cm long; blade 10–12 × 8–10 cm, base rounded or truncate, subpeltate, apex obtuse or emarginate, entire, thick-coriaceous; peduncles 10–30, 2–4 mm long, fasciculate or racemose on old wood, rachis to 15 cm long;

646

P ASSIFLORACEAE

bracts scattered on peduncle, 1 mm long; pedicels to 16–20 mm long; floral cup 3.5–4 mm high, urceolate, slightly umbilicate, 5lobed, internally the grooves as rounded septa 1–2 mm high joining the 5 angles of the base of the androgynophore; floral tube slightly conic-cylindric, 2.5–4.5 cm long, 6–9 mm diameter at throat, wider at middle; sepals lanceolate, 10–20 × 6 mm, obtuse; petals slightly smaller; radii in 1 series, ca. 10 mm long, cultrate, flattened tangentially, orange; pali in 1 or 2 series of tubercles 0.1–0.5 and 0.1–0.2 mm long; operculum tubular, horizontal then erect, margin fimbrillate for 1 mm, erect and appressed to androgynophore; ovary obovoid, narrow, glabrous; fruit narrowly fusiform or ellipsoid to ovoid, 7–10 cm long, violet-rose. Evergreen lowland to lower montane forests, disturbed margins, 100– 1000 m; Bolívar (San Ignacio de Yuruani), Amazonas (Río Ocamo, San Juan de Manapiare, Tencua). Guyana, Suriname, French Guiana. Passiflora garckei Mast., Trans. Linn. Soc. London 27: 639. 1871. —Kaso’kana (Panare). Stipules semi-ovate or subreniform, 1.5– 7.5 × 1–3 cm, mucronate 1–2 mm, remotely glandular or glandular-serrulate; petiole 3– 10 cm long, glands ± in pairs; blade 7–15 × 8– 25 cm, 5- or 7-veined, base subpeltate 2–5 mm, truncate or subcordate, 3-lobed to just below middle or less, lobes narrow to broadly ovate to lanceolate-ovate, acute or obtuse, mucronulate, the middle lobe at most slightly narrowed at base, glandular in sinuses, entire or rarely serrulate, often finely revolute, frequently with pink veins beneath; peduncle solitary, 1.5–6 cm long; bracts just below articulation, ovate to oblong-lanceolate, 6–15 × 4–6 mm, acuminate-apiculate; pedicel 6–15 mm long; flowers 7–9 cm diameter, perianth white to lilac or violet; floral cup 3–5 mm high, urceolate, umbilicate; floral tube 3–4 mm high, funnelform; sepals oblong, 28–40 × 10–12 mm, cucullate, corniculus 1–6 mm long, narrow-subulate; petals 25–40 × 10–12 mm, obtuse; radii in 2 or 3 series, filiform, normally as long as petals, basal 3/4 blue to violet, tips white or yellow and slightly crisped; pali in 5 or 6 dense, somewhat irregular series, 2–8 mm long,

capillary, frequently capitellate; operculum erect, base membranous 2 mm, above linearfilamentous 6–8 mm, the bases white, broadened radially, joined and plicate inward in a ruff ca. 1 mm high and wide, occasionally capitellate or broadened at blue apex, erect in a ring around the androgynophore; ovary ovoid to obovoid, glaucous; fruit globose to subellipsoid, 6 × 4 cm. Evergreen lowland to lower montane forests and forest borders, 50–1000 m; Bolívar (Altiplanicie de Nuria, Amaruay-tepui, Caicara, Campamento Río Grande, Cerro Uei, El Manteco, Isla Anacoco, La Escalera, Luepa, Río Maniapure–Hato El Nazareno, Río Uaiparú, Santa Elena de Uairén, Sierra de la Cerbatana, Tumeremo), Amazonas (Gavilán, Río Cataniapo, Río Ventuari). Anzoátegui, Carabobo, Guárico, Monagas, Yaracuy; Guyana, French Guiana. ◆Fig. 560. Plants of Passiflora garckei are semicultivated for the edible fruits. Passiflora glandulosa Cav., Diss. 10: 453, t. 281. 1790. Stipules setaceous or subulate, deciduous; petioles 1–1.5 cm long, with 2 sessile wartlike glands below middle to near base; blade coriaceous, ovate, broadly ovate-elliptic to oblong-lanceolate, 8–16 × 3.5–10.5 cm, penniveined, slightly cordate to acute at base, apex acute or acuminate to rounded, mucronulate, glabrous to densely and minutely tomentulose beneath, with minute submarginal glands, slightly revolute; peduncles solitary, stout, 2.5–9 cm long; bracts linear-lanceolate, 5–10 mm long, at or just below articulation, margin swollen-glandular in lower half; pedicel 3–5 mm long; floral cup 2–3 mm long, urceolate, umbilicate; floral tube 10–12 mm long, 8–10 mm diameter at throat; sepals becoming reflexed, linearoblong to oblong, 3–5 × 1–1.5 cm, with very short subapical corniculus, margin with few dark glands, petals shorter, becoming reflexed; radii in one series at throat of tube, 4–10 mm long, subulate with wide base, or united in a membrane in lower 1/3, white or pink; pali to 13–14 mm long, membranous in the lower 1/2–3/5, the upper portion filamentous; operculum 3–5 mm high, filamentous in upper 1/4–1/2; ovary ellipsoid, densely short white-sericeous or glabrous; fruit ovoid, 4–6

Passiflora 647

cm long, pericarp coriaceous. Evergreen lowland to lower montane forest borders and open areas, near sea level to 1200 m; Delta Amacuro (Caño Cariabo, Río Amacuro), Bolívar (Gran Sabana, Isla Anacoco, Tumeremo), Amazonas (San Carlos de Río Negro). Guyana, Suriname, French Guiana, Brazil. ◆Fig. 554. Passiflora gleasonii Killip, J. Wash. Acad. Sci. 14: 112. 1924. Plant glabrous to minutely puberulous; stipules to 8 mm long; petiole to 2 cm long, caniculate above, with 2 sessile or slightly stipitate wart-like glands 2–5 mm below apex; blade oblong-lanceolate to lanceolateovate, 9–16 × 4.4–9 cm, base truncate to shallowly cordate, apex obtuse, abruptly acuminate, subcoriaceous, grayish beneath; peduncles 2.5–5 cm long; bracts at articulation, oblong-elliptic, 20–35 × 4–14 mm, base narrowed, basal half with large marginal glands, apex rounded, finely serrate, abruptly cuspidate-acuminate, finely puberulent; pedicel 5–7 mm long; flowers white to yellowish, to 8 cm wide; floral cup 3–5 mm high, urceolate, umbilicate; floral tube broadly campanulate, 5–8 mm long; sepals 30–35 × 10 mm, lanceolate, obtuse, with somewhat foliaceous corniculus to 4 mm long; petals linear, to 20 × 5 mm; radii in 2 series, 4–5 cm long, banded; pali linear, 1–2 mm long; operculum 7–8 mm high, cleft to base in segments to 3 mm wide; ovary ovoid; fruit globose to ovoid, 7 × 3.5 cm. Sea level to 200 m; Delta Amacuro (Macareo, Caño Mariusa), Amazonas (base of Sierra de la Neblina). Apure, Cojedes; Guyana. Passiflora gracilis J. Jacq. ex Link, Enum. Hort. Berol. Alt. 2: 182. 1822. Stipules narrowly linear-subulate, 2–15 mm long; petiole 4–12 cm long, slender, glands one pair, small; blade consistently 3lobed to middle or less, the lobes ± equal, broadly ovate, 3–7(–17) × 7–10(–20) cm, thin-membranous, normally 3-veined from base, submarginal ocellae may be present, and infrequently 2 marginal glands near base of blade; peduncle solitary, filiform, 2–4 cm long; bracts setaceous, 1–1.5 mm long, dispersed near apex of peduncle; pedicel ca. 1 mm long; flowers white, 20–25 mm diameter;

sepals oblong, 8–12 mm long, obtuse, concave; radii in one series, 5–8 mm long, filiform, curved, with 2 bands of purple-red on upper side near middle; 1 series of capillary pali, 1–2 mm long, capitellate or swollen; ovary ovoid, glaucous; fruit 3–5 × 1.3–2.5 cm, pericarp thin and spongy, splitting irregularly, or drying papery. Upland forest margins, 800–900 m; Bolívar (Santa Elena de Uairén). Aragua, Lara, Mérida, Miranda, Táchira; Costa Rica, Guyana, Brazil, cultivated in other countries. Passiflora guazumaefolia Juss., Ann. Mus. Natl. Hist. Nat. 6: 112, t. 39, fig. 1. 1805. Plant glabrous or rarely minutely puberulous; stipules setaceous, ca. 6 mm long; petioles 1–3 cm long, with 2 wart-like glands at middle; blade membranous to subcoriaceous, 7–12 × 3–4 cm long, oblong-lanceolate, margin minutely revolute, base rounded or truncate, apex abruptly acuminate, the tip frequently rounded and cucullate, subentire or minutely glandular-denticulate; peduncles slender, 2–4 cm long; bracts ovate, yellowish green, marginal glands largest near the narrowed base, apex rounded; pedicel 5–8 mm long; flowers to 7 cm diameter, white to cream; floral cup ca. 3 mm high, urceolate, umbilicate; floral tube a 2– 2.5 mm horizontal rim; sepals 2.5–3 × 1–1.2 cm, narrowly oblong, obtuse, with a short corniculus 4 × 1 mm; petals shorter, linearlanceolate, apex obtuse; 2 series of subequal radii, 1–2 cm long, filiform, white, extended parallel to perianth; pali in ca. 6 or 7 dense series, 2–3 mm long; operculum erect, 2–4 mm high, membranous, exserted, the margin denticulate; ovary ovoid, glabrous; fruit globose, to 4 cm diameter. Riparian forests, forest edges, sea level to 300 m; Delta Amacuro (Tucupita), Bolívar (Ciudad Bolívar, Moitaco, banks of Río Orinoco), Amazonas (Gavilán, Puerto Ayacucho, San Carlos de Río Negro). Barinas, Táchira, Zulia; Colombia. Passiflora holtii Killip, Publ. Field Mus. Nat. Hist., Bot. Ser. 19: 560. 1938. Woody vine or subscandent shrub, stem and petiole minutely puberulous, tendrils often reduced to spines; stipules deciduous; petiole 3–10 mm long, glands scar-like, on

648

P ASSIFLORACEAE

midrib at very base; blade base rounded or suboblique, tip ca. 15 mm long; pseudoraceme with a few leaves reduced to < 5 mm, with large glands; peduncles 4–5 mm long, thick, erect; bracts deltoid, ca. 1 mm long, just below articulation; pedicels 2–3 mm long; flowers red, minutely puberulous without; floral cup 8–15 mm high, ca. 3.5 mm diameter at urceolate, umbilicate base, 5lobed and fused to the androgynophore by 5 septa ca. 8 mm high; floral tube 20–35 mm long, 3 mm diameter at base, 5 mm at apex, infrequently curved; sepals narrowly oblong, obtuse, petals slightly narrower; radii in 1 series, radially subulate, laterally compressed, 2.5–3.5 mm long; pali in 1 series, capillary, ca. 1 mm long, base with an enlarged tubercle toward inside; operculum erect, 3–5.5 mm long, apically filamentous 2–3.5 mm; ovary 4 mm long, narrowly oblong, yellowish tan-sericeous, styles 5 mm long, the bases well separated; fruit green. Evergreen lowland forests and forest edges, 100–200 m; Amazonas (Río Casiquiare, Río Guainía, Río Siapa, San Simón de Cucuy). Colombia, Brazil. Passiflora laurifolia L., Sp. Pl. 956. 1753. Plant essentially glabrous; stipules narrowly linear-subulate, 3–5 mm long, coriaceous; petiole 5–15 mm long, stout, with 2 wart-like glands at apex or just below; blade 6–12.5 × 3.5–8 cm, base rounded or slightly emarginate, apex mucronulate, reticulate veined, entire, thick-coriaceous, lustrous; peduncles 2–3(–8) cm; bracts ovate-oblong, 2.5–4 × 1.7–2.5 cm, obtuse, glandular-serrate near apex, puberulent; pedicel 6 mm long; flowers pendent, 5–7 cm wide; floral cup urceolate, 5 mm high, deeply umbilicate; floral tube cylindric-campanulate, 3–10 mm long; sepals oblong, 2–2.5 × 1 cm, subulatecorniculate apically, externally green with red spots, internally red or purplish red; petals similar, smaller; radii in 2 series, outer ligulate, slightly attenuate, ca. 2 cm long, slightly declined from horizontal, inner 3–4 cm × 2 mm, ligulate, with slightly crisped tips, pendent in form of a bell around the androgynoecium, both series banded red, blue, or violet-purple, on white; pali in 4 series, the first 3 ca. 1 mm long, the innermost ca. 1 mm down floral tube, ca. 1.5 mm long,

filamentous or capillary; operculum horizontal, margin denticulate; ovary ovoid, densely short-velutinous; fruit ovoid, 5–8 cm long. Evergreen lowland to lower montane forests, gallery forests, sea level to 1000 m; Delta Amacuro (Río Orocoima), Bolívar (Anacoco, Río Caura, Río Parguaza, Río Suapure, Santa Elena de Uairén, Uei-tepui, Urimán), Amazonas (Puerto Ayacucho, upper Río Ventuari). Anzoátegui, Aragua, Barinas, Carabobo, Falcón, Miranda, Nueva Esparta, Táchira, Vargas, Yaracuy; Martinique, Guyana, Brazil. Passiflora longiracemosa Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 221, pl. 23. 1922. —Oroshomi thotho (Yanomami). Liana, glabrous; stipules deciduous, subulate, 0.5–1 mm long; petiole 1.5–2(–9) cm long, with 1 pair of subsessile, apical wartlike glands, oblong, ca. 2 × 1 mm, closely appressed on top surface of petiole, occasionally united into a single gland, but also extending onto base of midrib and raising union of blade base with petiole; blade 14–17 × 3.4– 15 cm, base shallowly rounded or cordate, slightly subpeltate, apex obtuse or bluntly acute, to minutely emarginate, with the midvein ending on the underside in a small dark gland, margin entire, finely revolute, above lustrous, beneath densely glaucous, frequently a few minute ocellae scattered on or near secondary or tertiary veins well within margin; inflorescence with narrow bracts to 2 mm long, shorter on old wood; peduncles 1–4 mm long; bracts deltoid to subulate, near base of peduncle, 1–2 mm long; pedicels 6–12 mm long; flowers scarlet red to coral pink, declined; floral cup 4–5 mm high, urceolate, 5-lobed without, joined to androgynoecium by 5 narrow septa 2–3 mm high; floral tube cylindric, 16–45 × 2–3 mm diameter at base, 4–5 mm at throat; sepals linear-oblong, 8–17 × 3–5 mm, obtuse; petals slightly smaller; radii in one series, ligulate, 2–5 mm long, erect, occasionally swollen, slightly dolabriform, or bifid at apex; pali in one series of minute tubercles 0.1–0.3 mm long, or a mere ridge, atropurpureous; operculum tubular, erect, ca. 3–9 mm high, upper 1/3–1/2 irregularly fimbrillate-filamentous and appressed to androgynophore; ovary

Passiflora 649

narrowly ovoid, glabrous, 3–4 mm long; fruit obovoid to elliptic, 5 × 3 cm, pericarp stiff and brittle, green with 6 longitudinal rose bands. Evergreen lowland to montane forests and forest borders, 300–1500 m; Bolívar (base of Cerro Roraima, El Paují, Kavanayén, Río Chiguao, Salto Pará, Santa Elena de Uairén, base of Soropán-tepui), Amazonas (Ocamo, Río Matacuni, upper Río Orinoco, San Carlos del Río Negro, San Juan de Manapiare, Sierra Parima, base of Sierra de la Neblina). Guyana, Brazil. ◆Fig. 557. The specimens examined may represent more than one taxon, but they are not of sufficient quality to make a decision. Passiflora maguirei Killip, Bull. Torrey Bot. Club 75: 415. 1948. Woody liana, glabrous; stipules deciduous; petiole 1.5–5 cm long, stout, glands patelliform, with raised border, 2 mm diameter, at very apex, blade 12–35 × 5–13 cm, coriaceous, lustrous, coarsely reticulate, secondary veins prominent; peduncles in the axil of a 6 mm long bract, with 2 glands above the middle, glands 1–1.5 mm diameter; floral bracts subulate-triangular, coriaceous, 1–2.5 mm long, at base of peduncle; pedicels 5 mm long; floral cup 4–10 mm long, floral tube 14– 16 mm long, both narrowly funnelform, ca. 8 mm diameter at throat; sepals linear-oblong, 2.2–4.5 cm × 5–8 mm, acute; petals oblong to elliptic-oblong, 2.2–3.5 cm × 6–15 mm, obtuse; radii in 2 or 3 series, bright yellow, the first 18–23 × 1–2 mm, linear-dolabriform, apex subulate or truncate, the second 5–10 mm long, narrowly linear or cultrate, the third 3–4 mm long, filiform; pali in 1 or more series, pink, 2–4 mm long, filiform, reflexed into tube; operculum 12–16 mm long, basally fused to floral tube in 5 ovate bands 8 mm long, above this divided into truncate or rounded segments 7–13 × 1.2–1.5 mm, just reaching throat of tube; ovary ovoid, finely and densely short-velutinous, longitudinally grooved, apex truncate; fruit ovoid to obovoid, ca. 4 × 2.5 cm, puberulous, pericarp thin, brown. Evergreen lowland forests and gallery forests, 50–200 m; Amazonas (Carmelitas, Canaripó, Coromoto, Galipero, Gavilán, Río Atabapo, Río Autana, Río Cuao, Río Sipapo, San Juan de Manapiare). Guyana, Brazil.

Passiflora maliformis L., Sp. Pl. 956. 1753. —Parcha de huesito. Stipules linear or narrowly lance-linear, to 15 mm long; petioles to 5 cm long, glands wart-like, near middle ca. 1.5 mm diameter, subsessile; blades ovate-lanceolate to orbicular-ovate, 9–15 × 5–9 cm, membranaceous to thin-membranaceous, crenate or finely serrulate, base rounded to slightly cordate, apex acute or acuminate, rarely rounded, juvenile (and occasionally mature) leaves 3-lobed 1/3–1/2 the length, lobes almost equal; peduncles solitary, to 5 cm long, declined; bracts thin-membranous, yellowish green, 4–7 × 3–4.5 cm long, united for ca. 1 cm at base, the remainder adpressed by a marginal tomentulose band and completely enveloping bud before opening, yellowish green; pedicel to 10 mm long; flowers pendent, to 10 cm diameter, floral cup 4–6 mm high, urceolate, umbilicate; floral tube broadly campanulate, 4–6 mm high, 1–1.4 cm wide at throat; sepals oblong, 4 cm long, apex cucullate, keeled and with a 5 mm corniculus without, fleshy, green; petals linear-lanceolate, to 3 cm long, light greenish white mottled with reddish purple; radii in 2 series, the outer 1.5 cm long, reflexed, the inner 3 cm long, pendent, the upper part banded with violet, base reddish purple; pali up to 10 irregular series of minute green purple-tipped tubercles; ovary glabrous; fruit globose, to 4 cm diameter, pericarp green or orange-green. Cultivated and possibly escaped, ca. 50 m; Bolívar (Ciudad Bolívar, where most likely cultivated). Aragua, Carabobo, Portuguesa, Táchira, Zulia (cultivated in Aragua and possibly the other states); West Indies, Colombia, Ecuador. ◆Fig. 564. Passiflora maliformis is widely cultivated in the Caribbean. The aril is sweet and aromatic. Passiflora misera H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 136. 1817. —Murciélago, Parcha. Plant herbaceous; stipules setaceous to linear, straight to falcate, to 3.5 mm long; petiole 0.5–4 cm long; blade membranous, 0.5–3 × 5–18 cm, midvein 1/10–1/2 the length of the laterals, base cordate to truncate or amply obtuse, apex, when blade 2-lobed, reduced to the mucro at tip of midvein, or fre-

650

P ASSIFLORACEAE

quently retuse, a shallow, broadly truncate central lobe may be present, ocellae small, dark, light-rimmed, and scattered; peduncles 1.5–11 cm long, bracts setaceous, rarely bifid or trifid, 3–6 mm long, scattered at and below apex of peduncle; pedicel 1–4 mm long; flowers 2.5–4.5 cm wide, white or tinged yellowish, greenish or violet; sepals lance- to linear-oblong, 10–20 × 5–7 mm, obtuse; petals shorter and narrower; radii in one series, filiform, attenuate, (5–)10–15 mm long; pali in one series 2–4 mm long, capitate; operculum plicate, incurved, the margin minutely fimbrillate; ovary ovoid, glabrous; fruit globose to broadly ellipsoid, 5–15 mm diameter. Riparian forests, forest edges, disturbed areas, sea level to 300 m; Delta Amacuro (Caño Mariusa), Bolívar (Caicara, Ciudad Bolívar, Río Caura, Río Paragua). Apure, Barinas, Cojedes, Distrito Federal, Falcón, Lara, Monagas, Portuguesa, Táchira, Trujillo, Zulia; Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay. Passiflora multiformis Jacq., Fragm. Bot. n. 169, t. 67, fig. 1. 1809. Plant sparsely to densely pilose-tomentose, petiole 2–5 cm long, glands minute, wart-like, sessile, near middle; blade 6–14 × 4–14 cm, 3-lobed, base occasionally with 2 or more large marginal glands, middle lobe ovate to ovate-lanceolate, acuminate, lateral lobes about 1/2 as long as middle lobe (some blades may be simple or mitten-shaped), margin finely serrulate, glandular in the acute sinuses, membranous, pilosulous on veins; peduncles solitary, 4–8 cm long, bracts 5.5–8 × 2.5–6.5 cm, light greenish white or yellowish white, tometellous externally, margins adpressed by tomentulose marginal band before opening, apex acuminate-apiculate; pedicel to 10 mm long; flowers pendulous, 5–6 cm diameter, perianth very light yellowish or greenish white spotted with red or purple, floral cup 6–10 mm high, deeply umbilicate, floral tube short-funnelform, 4–8 mm high; sepals lanceolate, 30–45 × 8–16 mm, apex rounded, keeled and with a 5–10 mm corniculus; petals linear-oblong to lanceovate, 28–40 × 5–7 mm, apex rounded; 2 series of radii, the outer thin, liguliform, ca. 1.5 cm long, reflexed, the inner thicker, flattened near base, terete above, attenuate and

crisped at tip, 2.5–3 cm long, forming a cage around androgynoecium, both series banded with red below and blue above; pali 6 or 7 indefinite series lining floral tube, the uppermost conical to short-filamentous, the inner tuberculate, the innermost 1 mm long, in a definite ring; operculum a 1–2 mm membrane horizontal to top of limen, below trochlea; ovary elliptic-ovoid; fruit 3–5 cm diameter. Savannas, secondary forests, forest borders, 100–200 m; Amazonas (Ocamo). Apure, Aragua, Barinas, Distrito Federal, Mérida, Portuguesa. Passiflora nitida H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 130. 1817. —Bejuco parcha, Parcha. Plant glabrous; stipules linear-subulate, 4–6 mm long, laterally glandular above middle; petiole 0.5–3 cm long, canaliculate above, with 2 wart-like glands at apex; blade base minutely decurrent, margin serrate, serrulate or undulate-denticulate, with the teeth minutely glandular, or crenate to subentire, coriaceous or subcoriaceous; peduncles stout, 3–7 cm long; bracts rounded at base and apex, light green; pedicel 3–4 mm long; flowers 9–11 cm wide, showy, perianth white, horizontal or reflexed; floral cup urceolate, umbilicate, 4–5 mm high; floral tube a spreading rim, 2.5–8 mm wide; sepals oblong-lanceolate, 3–5 × 1–2.3 cm, fleshy, obtuse, subulate-corniculate apically 1–4 mm, 5–13 mm below apex; petals slightly narrower and thinner; radii in 2 series 2–4 cm long, 1+ mm thick at base, subulate, long-attenuate and crisped in upper 1/2 or more, banded or mottled pink to bluish purple basally, bluish to white above and forming a spherical cage around the androgynoecium; pali in 4–6 filiform series, the outer one or two 2–4 mm long, plus 2 or 3 more ca. l mm long lining floral tube, the innermost to 1 cm long; operculum ascending, with erect 1 mm fimbrillate margin; ovary obovoid, glabrous; fruit, mesocarp to 1 cm thick. Mainly riparian habitats, 100–200 m; Amazonas (widely distributed south of Río Parhueña). Apure, Aragua, Yaracuy; Colombia, Guyana, Peru, Brazil. ◆Fig. 569. The populations in northern Venezuela are montane (1200–1300 m) and apparently Amazonian relicts. Passiflora nitida is occasionally semicultivated for the edible fruit.

Passiflora 651

Passiflora nuriensis Steyerm., Acta Bot. Venez. 3: 188, fig. 13. 1968. Woody vine, glabrous; stipules deciduous; petiole 10–20 mm long, glands apical, 2–3 mm long, patelliform; blade 8–14 × 3–5 cm, coriaceous, base minutely decurrent to pair of glands, veins and reticulations prominent beneath; peduncle with 3 deltoid bracts ca. 0.5 mm long at the base; pedicel ca. 1 mm long; floral cup urceolate, 4–5 mm high, umbilicate; floral tube 18–35 mm long, 6–10 mm diameter; sepals ca. 22 × 4–5 mm, lanceolate-oblong, fleshy, salmon red, obtuse, apex cupped; petals 16–18 × 5 mm, thinmembranous, oblanceolate-oblong, apex rounded; radii in one series, 3–5 mm long, dolabriform, ca. 1 mm wide, yellowish white, some bifid; pali in 3 series, clavate, the outer 0.5–1 mm long, the 2 inner < 0.5 mm long; operculum attached to base of androgynophore at 5 points 4 mm long, above divided into 5 erect laminae 7 mm long, bifid 2 mm at the apices; ovary angled, finely and densely white-villosulous; fruit unknown. Lower montane forests, 400–700 m; Bolívar (Cerro el Picacho, Hato de Nuria on Altiplanicie de Nuria). Endemic. Passiflora ovata Martin ex DC., Prodr. 3: 322. 1828. Shrub with few tendrils, glabrous except ovary; stipules linear, ca. 2 mm long; petiole 1.5–3 cm long, with 2 low wart-like glands at end of blade margins, purplish, 2 mm long; blade oblong to oblong-ovate, 6–11 × 3.5–6.5 cm, base rounded or subacute, apex rounded to truncate, subcoriaceous; peduncles 8–27 mm long, solitary or paired; bracts linear, 2 mm long; hypanthium 2–7 mm high, widely campanulate; sepals lanceolate, 20 × 10 mm, obtuse; petals slightly shorter; radii in one series, 10 mm long, subdolabriform, enlarged at middle, yellow below, dark brown above; pali in one series, 2 mm long; operculum tubular, clasping androgynophore, reaching or surpassing throat, purple; ovary white-tomentose. Montane shrubland, 900– 1000 m; Bolívar (southern base of Auyantepui). Guyana, French Guiana. Passiflora pedata L., Sp. Pl. 960. 1753. Plant herbaceous, stipules ovate to linearsubulate or deeply laciniate-fimbriate, 5–15 × 2–7 mm; petioles to 5 cm long; blades to 10

× 17 cm, the leaflets shallowly and distantly serrate-crenate, the teeth minutely glandtipped, the central leaflet petiolulate, base mainly attenuate, apex attenuate; peduncle solitary, to 5 cm long, stout; bracts green, 3–5 × 2–3 cm, oblong or ovate, laciniate-fimbriate, base cordate, apex rounded; pedicel 5–7 mm long; flowers bluish, 7–8.5 cm diameter; floral cup 3–4 mm high, urceolate, deeply umbilicate; floral tube funnelform, 5 mm long; sepals lance-ovate, 3–3.5 × 1–1.5 cm, pale green or violet internally, obtuse, cucullate with a dorsal corniculus 4 mm long; petals as long, 1 cm wide, oblanceolate to lanceolate, apex rounded or obtuse, pale blueviolet internally; radii in 2 series, the outer ligulate, the inner longer, broad at base and crisped at apex, 2–2.5 cm long; pali in several irregular series lining the floral tube, capillary, 0.2–0.5 mm long; operculum 2 mm high; ovary velutinous; fruit yellow, globose to obovoid, to 4.5 cm diameter, pericarp thick, seeds small and very numerous, black, aril scant, white. Cuba, Haiti, Colombia, Venezuela, Trinidad, Guyana, Suriname, Brazil; 2 varieties, 1 in Venezuela. Passiflora pedata var. pedata differs in its linear-subulate stipules, and is reported from Cuba, Haiti, Colombia, Trinidad, Guyana, Suriname, and Brazil. P. pedata var. stipularis (Killip) Killip, Publ. Field Mus. Nat. Hist., Bot. Ser. 19: 384. 1938. —Passiflora pedata subsp. stipularis Killip, J. Wash. Acad. Sci. 14: 114. 1924. —Mapuu thotho (Yanomami), Parcha. Stipules ovate, deeply lanciniate-fimbriate, 7–15 × 5–7 mm. Disturbed areas; 100– 800 m; Bolívar (Altiplanicie de Nuria, El Manteco, Guri, La Paragua, San Félix), Amazonas (Gavilán, San Juan de Manapiare, Río Ocamo). Aragua, Cojedes, Distrito Federal, Miranda, Zulia; Cuba, Guyana. Passiflora phaeocaula Killip, J. Wash. Acad. Sci. 17: 430. 1927. Plant scandent, minutely puberulous; stipules setaceous, early deciduous; petiole 10–15 mm long, minutely puberulent, obscurely flat scar-like-glandular under slightly decurrent blade margin, extending onto midrib and petiole, 2.5–4 mm long; blade base rounded to obtuse, lanceolate

652

P ASSIFLORACEAE

with apex rounded-obtuse, 6–11 × 3–6 cm, to elliptic-ovate with apex rounded to acute, 11–15 × 4–7 cm, margin finely revolute; peduncles 2 mm long, paired in axils of acicular bracts, 2–3 mm long, on a condensed rachis 1–1.5 cm long; bracts 1–1.5 mm long, subulate-triangular, below articulation; pedicel 4 mm long; floral cup 4 mm high, urceolate, 3–4 mm diameter; floral tube 12–14 mm long, obconic, 6–7 mm diameter at throat; sepals narrowly oblong, 17–20 × 6–7 mm, obtuse; petals similar, thinner and slightly smaller, cream to salmon, with violet spots; radii yellow with violet or red spots, linear below, flattened tangentially, dolabriform above, 3–4 × 1–1.5 mm, tip short, rounded obtuse; pali 0.7–2 mm long, linear-subulate, slightly flattened tangentially; operculum inclined, apex erect to clasp androgynophore, 4 mm long, upper 1.5–2 mm slightly plicate and filamentous, these flattened tangentially; ovary ovoid, minutely puberulous; fruit not seen. White-sand scrub (bana), 100– 200 m; Amazonas (between San Carlos de Río Negro and Solano). Adjacent Colombia, Brazil (Amazonas: upper Río Negro). Passiflora picturata Ker Gawl., Bot. Reg. 8: pl. 673. 1822. Stipules semiovate, 1.5–2.5 × 1–1.4 cm, apically finely aristate 2 mm; petiole thin, 2– 6 cm long, glands scattered; blade 3.5–6 × 4– 7 cm, thin-membranous, subpeltate 1 mm and truncate to subcordate or rounded at base, normally 3-lobed to middle or more, (occasionally unlobed, ovate, to 3 cm wide), lobes ovate to obovate, rounded or subacute, frequently short filamentous-glandular in the sinuses, purplish beneath, mucronulate, entire; peduncle solitary, 4–12 cm long, stout, thicker than petiole; bracts verticillate at articulation, basally (0.5–)2–4 mm petiolate, elliptic-spatulate, 15–35 × 8–15 mm, to broadly ovate (12–32 × 8–20 mm), apex obtuse, finely aristulate 3–5 mm; pedicel 1–6 mm long; flowers to 10 cm diameter, white to bluish or violet-pink; floral cup 3 mm high, 8–12 mm diameter, umbilicate, strongly urceolate, with a broadly-funnelform rim 2 mm wide; floral tube a 1 mm extension of rim; sepals oblong-elliptic, 25 × 10 mm, fleshy, with a keel terminating in a foliaceous apiculate corniculus 3–4 mm high, extended to 8 mm beyond apex; petals oblong-elliptic, slightly

smaller; radii 1 or 2 series, 5–20 mm long, banded violet and white; pali uniseriate, 2–5 mm long, banded violet on green; operculum 8–12 mm long, basal 1–2 mm membranous, strongly plicate, the 10 mm long filaments arising from apices of plications; ovary ovoid, glaucous; fruit globose, 3–3.5 cm diameter. Savannas, shrublands, gallery forests, 50– 1300 m; Bolívar (Gran Sabana, Puerto Ordaz). Anzoátegui, Carabobo, Yaracuy; Colombia, Trinidad, Suriname, Brazil. Passiflora pulchella H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 134. 1817. —Cinco llagas, Parchita, Parchita de monte, Pasionaria. Stipules 4–7 mm long; petiole 1–2 cm long; blade 2–7 × 4–10 cm, coriaceous, bilobed normally < 1/2 the length, the lobes spreading or suberect, frequently apiculate, base truncate, rounded or cuneate, apices truncate or rounded, rarely obtuse, occasionally with a reduced, broad central lobe, strongly reticulate-veined, ocellae few to numerous; peduncles to 9 cm long; bracts coriaceous to membranous, involucrate at the articulation and enclosing the young flower bud, base tapering abruptly, apex acute to obtuse; pedicel 1–2 mm long; flowers 4.5–5.5 cm diameter; floral cup saucered, umbilicate, 10–14 mm diameter; floral tube a ± flat rim 2–3 mm wide; sepals oblong, obtuse, 18–25 × 7–10 mm, greenish to yellowish; petals ovate- to oblong-lanceolate, ca. 1/2 as long, white, light to dark blue, to violet; radii in one series, filiform, as long as the petals, crisped, the tips widened and divided to lacerate, white or yellowish; pali in 2–4 series, 3–5 mm long, capitellate; operculum membranous, plicate; ovary subglobose, glabrous; fruit globose 10–20 mm diameter. Open areas, forest borders, savannas and shrublands, 50–200 m; Bolívar (Caicara, Ciudad Bolívar, Maniapure, Río Botanamo, Upata), Amazonas (San Juan de Manapiare). Apure, Aragua, Barinas, Carabobo, Cojedes, Falcón, Guárico, Lara, Portuguesa, Yaracuy, Zulia; southern Mexico, Central America, Colombia, cultivated as an ornamental in other countries. ◆Fig. 565. Passiflora pyrrhantha Harms, Notizbl. Bot. Gart. Berlin-Dahlem 9: 977. 1926. —Parcha de culebra.

Passiflora 653

Liana or subscandent shrub, puberulous on younger branches, tendrils, inflorescence, and flowers externally; tendrils often reduced to spines and thickened in basal 1 cm; stipules early deciduous; petiole 1–3 cm long, glands obscure, scar-like; blade ovate to ovate-oblong, 8–18 × 3.5–10 cm, base rounded, apex acuminate, margin finely revolute, subcoriaceous, glabrous; inflorescence a short-velutinous raceme or pseudoraceme 7–26 cm long, many-bracted at base, frequently on old wood; peduncles in pairs, stout, 0.5–1 mm long, axillary to bract-like reduced leaves or petioles with a pair of glands, rarely paired in axils of normal leaves; bracts at base of peduncle, deltoid, 1 mm long; pedicels 4–5 mm long; flowers deep orange to bright orange-red or red, 6 cm diameter, appressed-puberulous externally; floral cup 7–10 mm high, urceolate, umbilicate, 4–6 mm diameter, fused within to 5angled androgynophore by 5 septa 5–7 mm high; floral tube 23–31 mm long, ± cylindric, 4–5 mm diameter at throat; sepals ellipticoblong, 20–30 × 6–10 mm, obtuse or subacute; petals subequal to sepals, thinner; radii in one series, 0.5–6 mm long, lanceolate to lanceolate-spatulate, yellow; pali in one series, subulate 1–2 mm long; operculum erect, 7–10 mm long, filamentous 5–9 mm of this; ovary narrowly ellipsoid, velutinous, apex truncate; fruit ellipsoid, 7–11 × 3.5–4.5 cm, hard, dark and lighter green. Riparian habitats, lower montane forests, 50–600 m; Bolívar (Río Suapure), Amazonas (slopes of Cerro Huachamacari, Gavilán, Río Ocamo). Apure, Mérida, Táchira; Colombia, Ecuador, Peru. Passiflora quadrangularis L., Syst. Nat. 2: 1248. 1759. —Badea, Parcha grenadina. Cultivated vine with stout stems, glabrous; stipules ovate to ovate-lanceolate, thin-membranaceous, 2–4 × 1–2.3 cm, base narrowed, apex acute, entire or serrulate; petiole 2–5 cm long; blade broadly ovate to ovate-oblong, to suborbicular, 9–20 × 8–15 cm, thick-membranceous, base subcordate or truncate, apex rounded and abruptly acuminate, margin entire, midvein and 10–12 laterals prominent and elevated beneath; peduncles 1.5–3 cm long, 3-angled; bracts cordate-ovate, 3–5.5 × 1.5–4 cm, thin-mem-

branaceous, acute, entire or serrulate toward base; pedicel 10 mm long; flowers white, or pinkish to violet, to 12 cm wide, pendulous; floral cup urceolate, umbilicate, 4–6 mm high × 2.5–3 cm diameter, floral tube broadly campanulate, 3–4 mm high; sepals ovate to ovate-oblong, 3–4 × 1.3–2.5 cm, concave and cupped at apex, corniculus 5 mm long; petals lanceolate to ovate-oblong, 2.5–4.5 × 1–2 cm, obtuse, flat; radii in 2 series, filamentous, to 6 cm long, radiate, white banded from base with reddish purple to blue, becoming pinkish blue in upper 1/2; pali in 3 series, the outermost clavate, purplish, ca. 2 mm long, the next filamentous 1–1.5 mm long, banded reddish purple, the inner a thick membrane, 3–7 mm long, 5 or more cleft, inclined against upper surface of the trochlea; operculum 4–6 mm long, inclined against middle of trochlea, denticulate or short-filamentous at the reddish purple margin; ovary ovoid, stigmas reniform, 5–8 mm wide; fruit oblongovoid, 20–30 × 12–15 cm, frequently 3grooved longitudinally, pericarp very thick and firm-fleshy, arils translucent white, sweet. Cultivated and locally escaped, 50– 200 m; Bolívar (Ciudad Bolívar), Amazonas (Maroa, Puerto Ayacucho, Río Guainía, San Juan de Manapiare, San Simón de Cocuy). Widely cultivated at elevations below 500 m in northern Venezuela and many other tropical countries; infrequently escaped, origin uncertain, possibly Brazil. Passiflora quadrangularis is much used to make ‘batidos’ and ‘merengadas’ (whipped fruit juice and milk shakes), using both arils and the mesocarp. Passiflora quadriglandulosa Rodschied, Med. Bem. Kol. Rio Essequibo 77. 1796 [1794]. —Cámero (Baniba), Cu-da-shidé (Piaroa), Merecoya (Warao), Oroshomi thotho (Yanomami), Parcha, Parcha de culebra, Shakrakarimi (Yanomami). Stipules setaceous, 3–5 mm long; petiole 1–2.5 cm long, with 2–4 low, wart-like glands at and near base; blade deeply 3-lobed, (also entire and oblong to oblong-lanceolate or mitten-shaped, 3-veined from base), 6–18 × 3–14 cm, base rounded to truncate or subcordate, lobes acuminate, uniformly or irregularly and finely to coarsely repand-serrate, the teeth minutely gland-tipped, whitish- to

654

P ASSIFLORACEAE

ferruginous-puberulent, -tomentose or tomentulose beneath and on veins above, middle lobe ovate to ovate-lanceolate, narrowed at base, longer and wider than laterals, frequently with small glands in the sinuses of the dentations; peduncle solitary, 2.5–5 cm long, infrequently in a raceme or pseudoraceme 8–12 cm long; bracts involucrate at articulation, linear to narrowly oblong-lanceolate or elliptic, 8–15 mm long, 2 or more large glands at and near base, finely glandular-serrate or nearly pectinate; pedicel 2–5 mm long; flowers pink to scarlet red; floral cup urceolate, umbilicate, (5–)6–10 mm high; floral tube, (2–)7–15 × 6–9 mm; sepals oblong-lanceolate, 4–8 × 1–1.5 cm, often with dark marginal glands externally, largest at base, acute, carinate, corniculus acicular, 2– 10 mm long; petals slightly shorter, obtuse; 2 series of radii linear at base, subulate above, the outer 10–20 mm long, the inner equal or shorter, mainly red, infrequently one or the other purplish or white, or red basally or apically; one series of pali 8–15 mm long, basally a red membranous tube, white- or redfilamentous in upper 1/3; ovary ovoid, yellowish- to ferruginous-villosulous; fruit ovoid, 3–8 × 2.5–5 cm, speckled, arils sweet and edible. Secondary vegetation, shrublands, lowland to lower montane and riparian forests, near sea level to 700 m; Delta Amacuro (Río Amacuro, Río Cuyubini, Río Grande, Sacupana), Bolívar (scattered over most of the north of the state), Amazonas (widely distributed). Sucre; Lesser Antilles (introduced?), Trinidad, Guyana, Amazonian Peru, Brazil, and Bolivia, cultivated elsewhere. ◆Fig. 559. I have followed the bract morphology of the Passiflora vitifolia complex treatment by Gentry (1981) to determine flora area material as P. quadriglandulosa (although it is not restricted to seasonally flooded riparian forests (várzea), as Gentry suggests). Otherwise it is impossible to distinguish this from P. vitifolia in the flora area. Passiflora retipetala Mast., Bull. Misc. Inform. Kew 1893: 12. 1893. Passiflora lonchophora Harms, Notizbl. Bot. Gart. Berlin-Dahlem 10: 813. 1929. Stipules semi-ovate, 1–4 × 0.8–2 cm; petiole 2–5 cm long, glands 4–8, scattered near middle and above, sessile to short-stipitate,

< 1 mm diameter, yellowish; blade ovate, 6– 18 × 4–14 cm, membranous to subcoriaceous or chartaceous, glossy above, glaucous beneath, base obtuse or rounded to subcordate, subpeltate, entire or 3-lobed to below middle, 3- or 5-veined from base, central lobe lanceolate-ovate, narrowed at base, with reduced marginal glands in the sinus, apices acuminate or acute to obtuse; peduncle solitary, 2–4 cm long, bracts verticillate at articulation, sessile, lanceolate to cordateovate, 15–25 × 5–10 mm; pedicel 5–8 mm long; flowers 5–7 cm diameter, white; floral cup 5–7 mm high, urceolate, umbilicate; floral tube 2–3 mm long, funnelform; sepals light green, lanceolate to oblong-lanceolate, with an apical foliaceous corniculus 3–4 mm long; petals white, lanceolate, slightly longer than sepals; radii uniseriate, 1.5–2.5 cm long, white banded with purple; pali in 4 or more series 2–4 mm long, base lavender, apex yellow; operculum with erect filaments to 5 mm long; ovary ovoid, glaucous; fruit globose to ovoid, 4–6 × 4 cm. Montane forests and forest borders, 1000–1700 m; Bolívar (Gran Sabana, Ikabarú, La Escalera, Macizo del Chimantá [Chimantá-tepui, Toronótepui]). Sucre; Trinidad, Guyana, Brazil. ◆Fig. 558. Passiflora riparia Mart. ex Mast. in Mart., Fl. Bras. 13(1): 599, pl. 116. 1872. Liana; stipules linear, ca. 4 mm long, deciduous; petioles to 4.5 cm long, 2 low wartlike glands at middle; blade to 19 × 9 cm, oblong to oblanceolate, base retuse or rounded to narrowed, apex abruptly acuminate, penniveined; peduncles stout, 2–5.5 cm long, in pendulous axillary branches with or without reduced leaves, occasionally axillary to normal leaves; bracts 3–5 × 2–3 cm, reddish; pedicels 5–10 mm long; floral cup urceolate, umbilicate, ca. 5 mm high, 16 mm diameter; floral tube broadly funnelform, 9 mm high, 20 mm diameter at throat; sepals oblong, quite fleshy, 4–5 × 2 cm, obtuse, cupped, opening only partially, corniculus to 5 mm long, thick and conical; petals shorter, oblong-linear, apex obtuse; 2 (rarely 1) series of radii, 4–5 cm long, the inner to 2 mm thick, filamentous, fleshy, banded red to purplish, curved inwards forming a spherical cage around the androgynoecium, outer series more slender; pali tubercular to filamentous

Passiflora 655

and capitellate, to 2 mm long in 2 or more series within the floral tube; operculum horizontal, the recurved margin with short, capitellate filaments; ovary ovoid, normally densely velutinous; fruit ovoid or globose, 3– 10 × 2.5–5 cm, glabrous or tomentose, pericarp 1 cm thick. Evergreen lowland forests, 100–200 m; Amazonas (Gavilán, Tamatama). Colombia, Ecuador, Peru, Brazil. Passiflora rubra L., Sp. Pl. 956. 1753. Plant herbaceous; stem, and generally leaves, moderately to densely gray- or yellow-brown-pubescent, rarely glabrate; stipules setaceous to linear-subulate, 5–8 mm long; petiole to 5 cm long, frequently reddish; blade to 10 cm long, broadly oblong, the 2 lobes broadly lanceolate to ovate, acute or obtuse, apiculate or mucronulate, usually widely divergent, infrequently a middle lobe present, reduced or equal to the laterals, membranous; peduncle solitary, or rarely in pairs, 1.5–6.5 cm long; pedicel 2–5 mm long; flowers to 5 cm diameter, white, often tinged red; sepals linear-lanceolate, 10–30 × 3–6 mm, petals linear, 10–20 × 2–4 mm; radii in one series, narrowly ligulate, or filiform above, 5–18 mm long, geniculate, purplish red or lavender basally, white or greenish above; pali frequently present as filaments < 2 mm long, 1–2 mm below radii; operculum membranous, slightly plicate, limen a shallow, fleshy cup, frequently lobed; ovary subglobose, 6-ribbed, densely white- or brown-hirsute; fruit obovoid or pyriform, sparsely hirsute to glabrous, acutely 6angled, red, or only the angles and base red, 2–6 × 1.5–3 cm, base tapering, apex rounded or obtuse. Semideciduous to evergreen lower montane forests, 200–900 m; Bolívar (El Manteco, Guri, Santa Elena de Uairén), probably adventive in flora area. Aragua, Barinas, Carabobo, Distrito Federal, Falcón, Lara, Miranda, Sucre, Táchira, Vargas, Yaracuy, and probably other Venezuelan states; West Indies, Trinidad, Colombia, Ecuador, Peru, Brazil, Bolivia. Passiflora sclerophylla Harms, Notizbl. Bot. Gart. Berlin-Dahlem 6: 347. 1917. Scandent shrub, finely puberulous; stipules subulate, thick, 1–2 mm long, early deciduous; petiole 5–12 mm long, appressedpuberulous, with 2 scar-like glands at apex

as continuation of slightly decurrent margin of the blade, and onto base of midrib; blade thick, stiff-coriaceous to -chartaceous, apex rounded, emarginate, or subacute and finely apiculate, drying brown on upper surface, olive on lower surface; peduncles 1–3, 2–5 mm long, appressed-pubescent; bracts deltoid, ca. 1 mm long, at or near apex of peduncle; pedicels 8–15 mm long; floral cup and tube funnelform, densely tomentulose, the cup 3.5–5.5 mm high, neither urceolate nor umbilicate, the tube 4–12 mm long, 6–10 mm wide at throat; sepals narrowly oblong, 20– 25 × ca. 7 mm, obtuse, densely olivevillosulous externally; petals narrowly oblong to lanceolate, 18–25 × ca. 6 mm; radii dolabriform to subdolabriform, laterally compressed, slightly geniculate, tip shortly acuminate, or upper 1/2 narrow linear-attenuate, verrucose, dull yellow spotted brownish purple; pali filiform to linear, 1.3–4 mm long, in same flower subdolabriform or branched above in a flat, unequal Y shape; operculum variable, an erect ridge 1–2 mm high cleft nearly to base in filaments, or linear to linear-clavate filaments 2.5–3 mm long, or these nearly 4 mm long and widened (1.5 mm) geniculate above the middle; ovary narrow-oblong, densely olivaceous-villosulous to glabrescent; fruit ellipsoid, rounded 6-angled, 4–7.5 × 2–4 cm, apex, with bases of styles extended in a narrow nipple 5–10 mm long, the pericarp thin-coriaceous, sparsely puberulent. Montane savannas and shrublands, riparian forests, 600–2200 m; Bolívar (Gran Sabana, likely on most tepuis), Amazonas (Cerro Aracamuni, Cerro Duida, Serranía Uasadi). Guyana (Mount Roraima). ◆Fig. 555. On different tepuis, plants of Passiflora sclerophylla are quite variable in leaf form, pubescence, and corona characters, but there are insufficient collections to erect subspecific taxa at this point. Passiflora securiclata Mast., Bull. Misc. Inform. Kew 1893: 12. 1893. —Ahorca vena’o, Bejuco parcha, Parcha diente de culebra, Parcha montañera. Scandent vine to subscandent shrub, glabrous, the tendrils often reduced to spines; stipules minute, setaceous, deciduous; petiole 1–1.5 cm, glands apical, scar-like or with a slightly raised border, extended slightly

656

P ASSIFLORACEAE

onto midrib and base of blade, reddish; blade 5–18 × 2–8 cm, rarely mucronulate, thinly coriaceous or subcoriaceous; inflorescence of slender pseudoracemes, very finely puberulous; peduncle 2 mm long; bracts triangulardeltoid, 0.5 mm long, at base of peduncle; pedicels 3–12 mm long; flowers reddish orange to deep red; floral cup 7–10 mm high, enlarged urceolate, umbilicate, 2.5 mm diameter, 5-lobed at base and joined within 4–5 mm to androgynophore by 5 septa, tubular above; floral tube 15–35 mm long, cylindric, 3–4 mm diameter at throat; sepals narrowly oblong, 15–20 × 3–6 mm, obtuse; petals similar, slightly smaller; radii in one series, cultrate to dolabriform, 3–5 mm long, finely and irregularly lobed along inner margin; pali in one series, filiform, 0.5–2 mm long, filamentous, swollen at base; operculum erect, tubular, 5–10 mm high, upper half fimbrillate; ovary narrowly ellipsoid, 4 mm long, densely and minutely velutinous, truncate, with the style bases well separated; fruit ovoid 3.5–5 × ca. 3 cm, pericarp glabrous, thin, stiff and brittle, green or yellowish with longitudinal green stripes. Gallery forests, seasonally flooded forests, near sea level to 300 m; Delta Amacuro (Piacoa), Bolívar (Caicara, Ciudad Bolívar, Hato La Vergareña, Moitaco, Río Caura, Río Chiguao, Río Cuchivero, Río Oris, Río Paragua, San Pedro de las Dos Bocas), Amazonas (Caño Iguapo, Isla Ratón, La Esmeralda, Puerto Ayacucho, Río Casiquiare, Río Manapiare, Río Ocamo, San Antonio, Tamatama, Venado, Yutajé). Anzoátegui, Apure; Colombia, Guyana, northern Brazil. ◆Fig. 567. Passiflora seemannii Griseb., Bonplandia (Hanover) 6: 7. 1858. —Chãmore (Piaroa), Parcha, Parcha de ratón. Stipules narrow, glandular-serrulate medially, apically attenuate, 9–14 × ca. 1 mm; petiole 3–9 cm long, glands ca. 2 mm long, apical on petiole, hidden just beneath lobes of blade, infrequently with a second pair of glands at the middle; blade broadly ovate, deeply cordate, 7–15 × 5–11 cm, the basal lobes usually overlapping, margin denticulate (occasional leaves with 1 or 2 lateral lobes); peduncles solitary, 5–10 cm long; bracts broadly lanceolate, 2.5–4 × 1.5–2 cm, at articulation, acuminate, united margin-

ally 1/3 or more, whitish, purple-tinged, enclosing 1/3–1/2 of perianth; pedicel 4–10 mm, 1/2 within umbilicate base of flower; flowers to 10 cm diameter, pendulous; floral cup 6–8 mm high, urceolate-campanulate, deeply umbilicate; floral tube funnelform, 1.3–2 cm long; sepals ovate-lanceolate, 2.5–4 × 1.5–2 cm greenish white tinged with violet, shortcorniculate near apex; petals oblong, 2–3.5 × 1–1.4 cm, pale purplish or lilac; radii biseriate, outer 14–20 mm long, the inner 20–30 mm long, filamentous-ligulate, attenuate, banded lavender on white, tips crisped; pali innumerable short, slightly-filamentous tubercles < 0.3 mm long, irregularly lining entire floral tube, the innermost a definite ring of filaments 0.7–1 mm long; operculum a horizontal membrane 1–2 mm wide, border curved up and ciliate 0.5 mm long; ovary ovoid or ellipsoid, glabrous; fruit ovoid, to 5 × 3.5 cm. Forest edges, near cultivated areas, 50–500 m; Amazonas (Caño Marieta, La Esmeralda, Puerto Ayacucho, Río Sipapo, San Fernando de Atabapo, San Juan de Manapiare, Yavita). Táchira; Mexico, Central America, Colombia. ◆Fig. 563. Passiflora seemannii is edible and is frequently cultivated in the flora area, in the Caribbean, and elsewhere. Passiflora serrulata Jacq., Observ. Bot. 2: 26, t. 46, fig. 2. 1767. Older stem with abundant cork; petiole to 3 cm long, two wart-like glands near middle or slightly below, slightly cupped, rarely a second pair above; blade 5–16 × 3.5–11 cm, lateral lobes 1/3–1/2 as long as middle lobe, middle lobe ovate to ovate-lanceolate, much narrowed at base, obtuse or acute (some blades may be simple and broadly ovate, or mitten-shaped), base truncate to shallowly cordate, margin finely glandular-serrate, 2– 4-glandular near rounded sinus, membranous or coriaceous, glabrous to densely pilosulous or white-tomentose on petioles, underside of leaf blade, and veins on the upper leaf surface; peduncles solitary, 2–6 cm long; bracts 3–6.5 × 1.6–3.7 cm, initially completely united by tomentulose marginal band, margin serrulate, the teeth minutely glandular; pedicel 5–12 mm long; flowers creamy white, 3.5–7 cm diameter, pendulous, fragrant, bracts and perianth reflexed in an-

Passiflora 657

thesis; floral cup urceolate, umbilicate, 3–4 mm high; floral tube conical, 1–1.5 mm long; sepals oblong-lanceolate, 15–25 × 9–12 mm, obtuse; petals lanceolate, ca. 13 × 4 mm; 2 series of radii linear, outer 20–25 mm long, crisped at tips, inner 12–15 mm, purple banded above white base; 2–4 series of minute pali in floral tube, tuberculate to filamentous, ultimate series 1–1.5 mm long; operculum a horizontal membrane 1 mm wide, row of filaments 0.3 mm long from below outer margin; ovary ovoid, glabrous; fruit edible, spherical, 2–3 cm diameter, yellow, pericarp coriaceous. Savannas, shrublands, deciduous forests. 50–800 m; Bolívar (Ciudad Bolívar, El Manteco, Guasipati, Upata, probably much Llanos vegetation in northern part of state). Anzoátegui, Aragua, Carabobo, Cojedes, Falcón, Guárico, Lara, Monagas, Portuguesa, Sucre, Vargas, Zulia; coastal Colombia Trinidad, Tobago. ◆Fig. 562. Passiflora spinosa (Poepp. & Endl.) Mast., Trans. Linn. Soc. London 27: 630. 1871. —Tacsonia spinosa Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 59, t. 181. 1838. Liana; stems, petiole, tendril, inflorescence, and flower externally minutely puberulous, tendrils often reduced to stout spines with the basal 5 mm thickened; stipules deciduous; petiole stout, 10–30 mm long; blade 10–21 × 3–9 cm, base rounded, with 2 scar-like glands on lower surface and extended from the minutely decurrent margin, lustrous, apex acuminate; peduncles 1–5 mm long, subtended with leaves reduced to short petioles with 2 large glands; bracts at articulation, deltoid, 1 mm long; pedicels 3–6 mm long; flowers bright red to orange, floral cup 10–14 mm high, to 6 mm diameter at base, urceolate, umbilicate, joined within to androgynophore by 5 septa; floral tube, 20– 26 mm long, 2.5–4 mm wide at base, 6–8 mm wide at throat; sepals narrowly oblong, obtuse; petals similar; radii yellow, in one series, cultrate-dolabriform, 2–5 mm long, ca. 1 mm wide radially, slightly thickened-verrucose along upper inner margin, apex acute or filiform; pali yellow, in 1 or 2 series, filiform, 1–2 mm long, the base bulbous; operculum erect, tubular, 2.6–12 mm long, with subulate filaments in apical 1/3–2/3; ovary narrowly oblong, densely yellow-tan short-

villosulous; fruit ovoid or ellipsoid, 5–6 × 2.5 cm, rounded hexagonal in section, ochre. Evergreen lowland to montane forests, 50–1300 m; Bolívar (Altiplanicie de Nuria, Enaña, Los Pijiguaos, Río Suapure), Amazonas (Cerro Huachamacari, Gavilán, Río Casiquiare, San Carlos del Rio Negro, base of Sierra de la Neblina). Colombia, French Guiana, Peru, Brazil. Passiflora suberosa L., Sp. Pl. 958. 1753. Passiflora pallida L., Sp. Pl. 955. 1753. Stipules linear-subulate, to 8 mm long; petiole 0.5–4 cm long; blade extremely variable in form and size, even on the same branch, 3–11 × 0.3–13 cm, usually membranous, infrequently subcoriaceous, normally 5-veined from base and shortly (or deeply) 3lobed (or simple or mitten-shaped), base obtuse to cordate, the lobes broad to linear, the laterals quite reduced to as long as the central, ocellae obscure if present; peduncles normally in pairs, to 15 mm long, pedicel 2–7 mm long; bracts, if present, setaceous to linear, above articulation, minute, early deciduous; flowers yellowish green, 8–30 mm diameter; sepals lanceolate to ovate-lanceolate, 5–8 mm long, obtuse to acute, the one innermost often petaloid; radii in one series, filiform, recurved, to 4 mm long, yellowish, spotted purplish at base; pali shorter, capitellate, purplish; ovary subglobose or ovoid; fruit soft and juicy. Savannas, shrublands, disturbed areas, 100–600 m; Bolívar (Altiplanicie de Nuria, El Manteco), Amazonas (Capibara). Aragua, Carabobo, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Sucre, Táchira, Vargas, Zulia; U.S.A. (Florida, Texas), Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, introduced in the Paleotropics. Passiflora tuberosa Jacq., Pl. Hort. Schoenbr. 4: 49, t. 496. 1804. —Kaso’ kana (Panare). Plant herbaceous, stipules narrowly linear-subulate, falcate, 2–5 mm long, coriaceous; petiole 1–3.5 cm long; blade oblong, 4– 16 × 4–12 cm, base rounded or truncate bilobed ± below middle, often deeply, lobes often nearly parallel, lanceolate to oblong-

658

P ASSIFLORACEAE

lanceolate, obtuse, mucronulate, frequently with lighter or white splotching along the principal veins, sinus rounded, mucronulate or apiculate; peduncles paired, to 4 cm long; bracts setaceous, scattered on peduncle, 2–3 mm long; pedicel 5–7 mm long; flowers 4.5–6 cm diameter, white; floral cup patelliform, slightly umbilicate, floral tube a mere rim; sepals lanceolate-deltoid, 15–25 mm long; petals ovate-lanceolate to narrowly oblong, 6–15 mm long, somewhat reflexed between sepals; radii in 1 or 2 (or more?) series, purple-banded, ligulate or cultrate, 3–7 mm long, the apex occasionally clavellate; pali in 1 or 2 (+?) series, filiform or slightly clavellate, 0.5–2 mm long; operculum plicate; ovary (sub)globose, 2.5 mm diameter; fruit black, ovoid, 10–15 mm diameter. Riparian forests, near sea level to 400 m; Delta Amacuro (expected), Bolívar (Caicara, Isla Anacoco, Río Cuyuní, Río Maniapure, San Antonio, Sierra Imataca). Aragua, Distrito Federal, Falcón, Lara, Miranda, Portuguesa, Sucre, Yaracuy; West Indies (St. Thomas, Trinidad), Guyana, Suriname, French Guiana, Brazil. Specimens from northern Venezuela, other than Sucre, may represent a separate taxon. Passiflora variolata Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 58, t. 179. 1838. —Hoja de culebra, Motodi (Baniba), Motodipicada (Curripaco), Parcha, Parcha de culebra de agua. Passiflora bomareifolia Steyerm., Mem. New York Bot. Gard. 17: 455. 1967. Woody vine, glabrous; stipules linear-falcate, 2–3 mm long, deciduous; petiole 6–15 mm long, biglandular near base, the glands almost scar-like to very slightly raised warts; blade elliptic, lance-elliptic to ovate to oblong, 5–14.5 × 1.5–6 cm, penniveined, base rounded to subacute, apex acute to acuminate, aristulate, minute, dark submarginal ocellae normally present, especially visible on young leaves; peduncles solitary, occasionally in pseudoracemes with reduced leaves; bracts verticillate at articulation, narrowly lance-linear, 3–8 × 1 mm, each side with 1 or 2 glands near base; pedicel 2–11 mm long; flowers pink to deep red, or pale violet, 9 cm wide, floral cup urceolate, 2–3 ×

8–10 mm, floral tube cylindric-campanulate, 3–5 × 10 mm; sepals linear-oblong, 35–45 × 7–10 mm, obtuse, carinate, corniculus green, 2–5 × 1–1.5 mm; petals subequal, narrow; radii in one series, normally white or cream, membranous-tubular, divided above in filaments 2–7 mm long; pali in one series, 2–3 mm high, membranous-tubular in lower half, linear-subulate filamentous above; operculum dependent then erect, membranous, 3–5 mm high, denticulate-laciniate; ovary ovoid-ellipsoid, glabrous to densely and minutely tomentulose; fruit 4.5–7 × 2–4 cm, ovoid with acute apex, exocarp ca. 2 mm thick, hard and brittle, spotted with green and with 6 green longitudinal lines, mesocarp pithy, ca. 3 mm thick. Forest borders, shrublands, secondary vegetation, 50–1200 m; widespread in Bolívar and Amazonas. Amazonian Colombia, Peru, and Brazil. ◆Fig. 568. Passiflora vespertilio L., Sp. Pl. 597. 1753. —Parcha montañera. Stipules 2–5 mm long; petiole 0.5–3 cm long; blade 2(3)-lobed, lunate to semi-circular or nearly square to transversely rectangular, 1–10 × 3–15 cm, 3-veined with 2 strong secondary veins from middle of midvein, base rounded or subtruncate to shallowly cordate, lobes divaricate, rounded or acute, upper margin of blade normally truncate or shallowly lunate to broadly emarginate between acuminate to broadly rounded lobes, blade normally coriaceous, the veins strong and prominent, basal pair of ocellae usually larger than the others and occupying the angle between the primary lateral veins and midrib; peduncles 3–10 mm long; bracts scattered just below the articulation; pedicel equal or slightly shorter; flowers to 5 cm diameter, white to yellowish green; sepals broadly lance-oblong, 12–20 × 6–9 mm, obtuse, subcoriaceous; petals narrowly oblong 8–12 × 2–5 mm; radii in one series of narrow ligulate filaments, 10–15 mm long, yellowish green, tips purplish; pali in one series, capillary, 3–4 mm long, greenish white; operculum closely plicate, white; ovary ovoid, glabrous; fruit subglobose, 1–3.5 cm diameter. Mainly riparian habitats, gallery forests, scrub, secondary vegetation, near sea level to 1000 m; Delta Amacuro (wide-

Passiflora 659

spread), Bolívar (widespread in the north to Altiplanicie de Nuria, La Escalera, and Salto Pará), Amazonas (widely distributed at low elevations along Río Orinoco to Río Negro, some tributaries and foothills). Aragua, Monagas, Portuguesa, Sucre, Táchira; Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 566. Passiflora sp. A Plant herbaceous, thinly tomentosepuberulous; stipules narrowly linear-subulate, subfalcate, 3–4 mm long; petiole 5–10 mm long; blade broadly elliptic, 5–7.5 × 4–5 cm, base rounded, shallowly bilobed at apex, young blades with a very short middle lobe, apex of lobes, and sinus, rounded, minutely mucronulate, firmly membranous, ocellae various; peduncles in pairs or solitary, ca. 1.5 cm long; bracts narrow-subulate, ca. 3 mm long, reddish brown, nearly verticillate 2–3 mm above base of peduncle; pedicels ca. 3 mm long; flowers 2.5 cm diameter; sepals broadly oblong-lanceolate, 15 × 8 mm, obtuse, puberulous and pale green without; petals oblong, 8–9 × 4 mm long, obtuse, pale green bordered with lavender; radii in one series, liguliform to filiform, 7–9 mm long, broadened at base to ca. 1 mm wide, green at base, maroon or lavender at center, tips yellow; pali in one series, 3–5 mm long, capillary, or some with base broadened, fused to outer base of operculum; operculum plicate, ca. 2–3 mm long, margin fimbriate; ovary globose, densely white-velutinous; fruit unknown. Montane forests, 1300–2000 m; Bolívar (La Escalera at km 134 south of El Dorado), Amazonas (Pico Zuloaga on Sierra de la Neblina). Endemic. Passiflora sp. A is closely related to the Andean P. andreana Mast. Passiflora sp. B. —Mapuu ke thotho (Yanomami), Mapuu thotho (Yanomami). Plant somewhat delicate, glabrous throughout; stipules 10–19 × 10–15 mm, apex obtuse, mucronulate; petiole thin, 2.5– 4.2 cm; blade thin membranous, ca. 4 × 6 cm, lower surface prominently 5-veined, base truncate, subpeltate 1 mm, 3-lobed to middle, the central slightly the larger, lobes

oblong-ovate, broadly rounded, mucronulate, sinus rounded, with 2–4 minute marginal glands, margin finely revolute; peduncles 25–35 mm long, in flower thicker than stem, petioles, and tendrils; bracts verticillate at articulation, broadly and deeply cordate, to 16 × 12 mm, shortly (< 1 mm) apiculate; pedicels 6–7 mm long; perianth reflexed in anthesis; floral cup 5 mm high, ca. 7 mm diameter, urceolate, umbilicate ca. 3 mm; floral tube a broadly funnelform rim, ca. 3 mm long; sepals ovate-lanceolate, 20–26 × 10 mm, cupped at apex, broadly carinate keeled with a foliaceous corniculus 6–7 mm high, extending to 7 mm beyond apex; petals ca. 16 × 5 mm, ovate-lanceolate, thinner; radii in 2 linear, filamentous series, outer 8–10 mm long, inner ca. 6 mm long; pali in 2 series, outer 0.5 mm tubercles, inner filaments < 1 mm long; operculum a 2.5 mm membrane, the apex triangular-plicate inward 0.5–1 mm long, with erect, flattened filaments to 5 mm long from the sinuses; limen surrounding enlarged base of androgynophore, the free margin expanded, ca. 1 mm wide, undulatelobed, at level of base of operculum; ovary ± globose, on 3 mm long gynophore, shortly beaked; fruit broadly ovoid, ca. 5 × 3 cm, the mesocarp of thin-filamentous aerenchyma. Gallery forests, 100–300 m; Amazonas (Río Ocamo). Endemic. Passiflora sp. B is known from only two collections to date. It resembles P. subpeltata Ortega in the foliaceous-corniculate sepals and aerenchyma of the fruit. Passiflora sp. C Woody climber, minutely tomentulose on all parts of developing apex, barely discernible on mature parts except flowers; stipules 0.1 mm tomentulose bulges, disappearing almost completely with growth; petiole 2–3.5 cm long, apically 2 scar-like glands 3 mm long; blade broadly ovate, 13–17 × 8–13 cm, base rounded to truncate or subcordate, apex sharply attenuate 1–1.5 cm long, margin finely revolute; peduncles paired in the leaf axils, 4–5 mm long, on a 1 mm stipe, light brown; bracts deltate, 1 mm long, at base of peduncle; pedicel 6–8 mm long, dark brown, densely minutely sericeous, as also exterior of hypanthium and calyx; floral cup 8–9 mm high, cylindric; floral tube 8 mm high, cylin-

660

P ASSIFLORACEAE

dric-funnelform; sepals green, thick, ca. 15 × 3.5 mm in late bud; petals cream with purple spots, smaller and thinner; radii in one series 12(–15?) mm long, purplish at base, yellowish green above, curved outward, tangentially flattened basally to 1+ mm, outer border verruculose, as also both borders of long-attenuate and crisped apical portion; pali in various series, the outer 3–4(+?) mm long, linear, the apex T-shaped, 0.5+ mm wide or with various short tubercular protuberances, these also along inner border of some filaments, or even short-lobed from the base, this series followed by < 10 irregular series down the upper 1/2 of the floral tube, diminishing in size from 2 mm to ca. 0.2 mm, the initial series tuberculate at the apex;

Fig. 551. Passiflora ambigua

operculum erect, 3–3.5 mm long, the upper 2 mm radially linear filaments; limen an undulate ring at midpoint of androgynophore, basal 1/2 of the androgynophore above this densely white-sericeous; ovary obovoid, 9lobed, densely white-sericeous, constricted at basally united styles, these white sericeous in lower 1/2; fruit ellipsoid, 10–13 × 4–6 cm, including the ± abruptly narrowed base 1– 1.5 cm long, in addition the apex surmounted by an umbo 15 × 10–12 mm, pericarp thin, stiff-coriaceous. Evergreen lowland forests, 100–200 m; Amazonas (San Simón de Cocuy). Endemic? The floral data are taken from a nearly mature bud and the remains of the flower at base of fruit.

Fig. 552. Passiflora capparidifolia

Passiflora 661

Fig. 553. Passiflora auriculata

Fig. 555. Passiflora sclerophylla Fig. 554. Passiflora glandulosa

662

P ASSIFLORACEAE

Fig. 556. Passiflora edulis

Fig. 558. Passiflora retipetala

Fig. 557. Passiflora longiracemosa

Passiflora 663

Fig. 559. Passiflora quadriglandulosa

Fig. 560. Passiflora garckei

Fig. 561. Passiflora foetida var. foetida

664

P ASSIFLORACEAE

Fig. 562. Passiflora serrulata Fig. 563. Passiflora seemannii

Fig. 564. Passiflora maliformis

Passiflora 665

Fig. 565. Passiflora pulchella

Fig. 566. Passiflora vespertilio

666

P ASSIFLORACEAE

Fig. 567. Passiflora securiclata

Fig. 568. Passiflora variolata

P E D A L I A C E A E 667

Fig. 569. Passiflora nitida

PEDALIACEAE by James S. Miller Herbs, annual or perennial, terrestrial or aquatic (Trapella), rarely shrubs, pubescence of short-stalked hairs with a multicellular head of mucilage-secreting cells, the plants often viscid. Leaves opposite or the upper leaves sometimes alternate, simple or lobed, entire or serrate, exstipulate. Inflorescence a terminal raceme, simple axillary dichasia, or flowers solitary and axillary, often subtended by 1 or 2 bracts each with an aborted flower functioning as an extrafloral nectary. Flowers bisexual, zygomorphic. Sepals (4)5, nearly free or basally connate; corolla sympetalous, irregular, sometimes with a basal spur, 5-lobed, the lobes imbricate, the limb often oblique or bilabiate. Stamens 4, epipetalous, in 2 pairs and alternate with the corolla lobes, the fifth reduced to a staminode, or the stamens 2 (Trapella), sometimes with 2 staminodes; anthers bithecal, dehiscing longitudinally. Nectary disk usually annular, surrounding the base of the ovary. Ovary superior, or rarely inferior (Trapella), 2-carpellate, usually 2-locular, sometimes 4-locular or 1-locular, the placentation axile or parietal; style terminal, the stigma 2-lobed; ovules 1–many per locule, anatropous. Fruit a loculicidal capsule or less commonly drupaceous or a nut, often with horns or prickles; seeds with a straight embryo and little or no endosperm. Pantropical, with a few temperate zone species; ca. 17 genera and ca. 80 species, 2 genera and 2 species in the flora area.

668

P EDALIACEAE

Craniolaria is sometimes placed in the segregate family Martyniaceae. Key to the Genera of Pedaliaceae 1. 1.

Flowers in a terminal inflorescence; floral tube > 10 cm long; fruit ± drupaceous, with a curved beak ........................................... 1. Craniolaria Flowers axillary; floral tube or corolla < 5 cm long; fruit a ± cylindrical capsule, not or only slightly beaked ........................................ 2. Sesamum

1. CRANIOLARIA L., Sp. Pl. 618. 1753. Herbs with viscid-villous pubescence. Leaves opposite, simple, 5-lobed, exstipulate. Inflorescences terminal, racemose. Flowers bisexual, zygomorphic, hypogynous. Calyx unequally 5-lobed, submembranous, campanulate-spathaceous. Corolla zygomorphic, sympetalous, 5-lobed, the tube long, cylindrical, narrow, campanulate at the mouth. Stamens 5, 4 fertile and didynamous, the fifth reduced to a staminode; filaments terminating in a glandular connective; anthers bithecal, dehiscing longitudinally. Ovary superior, 2-carpellate, 1-locular but appearing nearly 4-locular by intrusion of the parietal placenta; style elongate. Fruit drupaceous, ovate, apically rostrate. Seeds few or solitary in each locule. West Indies, Colombia, Venezuela, Bolivia, Paraguay, Argentina; 3 species, 1 in Venezuela. Craniolaria annua L., Sp. Pl. 618. 1753. Viscous herb; flowers white with purple splotches in the throat. Weedy areas, disturbed sites, sandy edges of rivers, near sea level to 200 m; Bolívar (La Fortuna near

Represa Guri, San Félix), Amazonas (Puerto Ayacucho). Anzoátegui, Apure, Aragua, Cojedes, Dependencias Federales, Guárico, Monagas, Nueva Esparta, Sucre, Zulia; Hispaniola, Puerto Rico, Colombia. ◆Fig. 570.

2. SESAMUM L., Sp. Pl. 634. 1753. Herbs, erect or prostrate, often viscid pubescent, rarely glabrous. Leaves opposite, subopposite to alternate on upper parts of the plant, simple or lobed, entire or serrate, usually pubescent with glandular hairs, petiolate. Flowers solitary in the leaf axils, rarely 2 or 3 per axil, subtended by small, aborted flowers functioning as extrafloral nectaries. Calyx 5-lobed, the lobes free to nearly the base. Corolla often basally saccate, 5-lobed and bilabiate, often showy, white, yellow, red, purple, or pink. Stamens 4 in 2 pairs, included in the corolla tube; anthers oblong, the connective dorsally thickened and with a terminal gland. Ovary superior, oblong, 2-carpellate, 2-locular but usually appearing 4-locular due to invagination of the ovary wall. Fruit capsular, oblong or ovoid, acuminate at the apex, 4-angular, opening apically. Seeds numerous. Tropical and southern Africa, India, Sri Lanka, the east Indies, and Australia; ca. 20 species, 1 introduced and cultivated in Venezuela. One species of this somewhat confused genus is widely cultivated in tropical and subtropical regions, and several others are less commonly cultivated in limited areas. Sesamum orientale L., Sp. Pl. 634. 1753. —Ajonjolí. Sesamum indicum L., Sp. Pl. 634. 1753. Erect annual herb to 2 m tall; flowers

white, pink, or lavender. Cultivated fields, disturbed areas, 50–200 m; Bolívar (near Caicara del Orinoco, El Dorado), Amazonas (Puerto Ayacucho). Cojedes, Distrito Federal,

Craniolaria 669

Fig. 570. Craniolaria annua

670

P EDALIACEAE

Falcón, Guárico, Monagas, Portuguesa, Yaracuy. ◆Fig. 571. Sesamum orientale is a cultivated plant of obscure origin that probably originated in east Africa or peninsular India (Nayar & Mehra, Econ. Bot. 24: 20–31. 1970.). It is widely cultivated in tropical and subtropical regions and occasionally escapes, although naturalized populations are probably shortlived. Seeds are used as a garnish in breads and cakes. They also have about 50% content of a high quality, semi-drying oil used in much the same manner as olive oil.

Fig. 571. Sesamum orientale

PERIDISCACEAE by Bruce K. Holst Trees. Leaves alternate, entire, lustrous above, strongly 3-veined from the base, lower surface with a small pit in the axil of each basal nerve; petioles pulvinate; stipules intrapetiolar, early deciduous and leaving a narrow oblique scar. Inflorescence an axillary cluster of very short racemes covered with short branched hairs, or flowers fasciculate; bracts fairly large, persistent; pedicels short. Flowers densely pubescent. Sepals 5–7, imbricate; petals lacking. Stamens numerous and inserted on a large cupular several-lobed nectary disk or outside an annular fleshy nectary

Peridiscus 671

disk; filaments partially and rather irregularly connate in the lower half; anthers very small, monothecal, 2-locular, didymous, opening from the middle by lateral valves, introrse. Gynoecium of 3 or 4 united carpels; ovary superior, sessile and half immersed in or subtended by the disk, depressed, 1-locular, with 6–8 ovules pendulous from the top of the ovary; styles 3 or 4, short, free, spreading. Fruits drupaceous, contracted at the base and so slightly stipitate, ovoid, 1-locular and 1seeded. Seeds with endosperm, the embryo bent. Southern Venezuela, Guyana, northern Brazil; 2 genera and 2 species, 1 species in the flora area. The second genus in the family, Whittonia Sandwith, is endemic to the Potaro River basin in Guyana and may eventually be found in Venezuela. It differs from Peridiscus by having the flowers in fascicles (versus racemes), and an annular disk that subtends the ovary (versus half-immersing the ovary). The family Peridiscaceae is nearly endemic to the Guayana Shield, but extends slightly south of the Shield into Amazonian Brazil. 1. PERIDISCUS Benth., Gen. Pl. 1: 127. 1862. Trees. Leaf blades glabrous, coriaceous, ovate to elliptic or oblong, acute to acuminate, strongly 3-veined from the base, the tertiary venation finely reticulate. Inflorescences short, axillary racemes, ferruginous-villous. Sepals usually 5, ferruginous-villous. Stamens inserted outside the large several-lobed disk; filaments partly united at the base. Ovary half immersed in the nectary disk. Fruit a globose drupe. Seed one; embryo small; endosperm copious, horny. Southern Venezuela, northern Brazil; 1 species. Peridiscus lucidus Benth., Gen. Pl. 1: 127. 1862. —Pariente cuajo. Tree to 26 m tall; flowers whitish, fruits ca. 2 cm diameter, yellow-green. Nonflooded

evergreen lowland forests, 50–500 m; Bolívar (Río Icabarú), Amazonas (Río Baría, Río Negro near Piedra de Cocuy). Brazil (Amazonas). ◆Fig. 572.

Fig. 572. Peridiscus lucidus

672

P HYTOLACCACEAE

PHYTOLACCACEAE by Gerardo A. Aymard C. and Nidia L. Cuello A. Herbs or succulent subshrubs, erect or clambering, rarely larger woody shrubs or trees, glabrous or sparsely and minutely puberulent. Leaves alternate, simple and entire, pinnately veined, often with short, linear crystals within; stipules absent or small and inconspicuous. Inflorescences terminal, axillary, or leaf-opposed, usually solitary, racemose, spicate, or compound-paniculate with spicate or racemose branches; bracts and minute bracteoles present or absent. Flowers small, actinomorphic or rarely somewhat bilaterally symmetrical, bisexual or unisexual by abortion. Perianth usually of 1 whorl of 4 or 5 parts (sepals or tepals), free or united near the base, imbricate in bud, usually persistent in fruit, greenish white to pink or purple. Stamens often as many as the perianth parts and alternate with them, or 3–many and variously arranged; filaments free or united only at the base, often borne on a hypogynous disk, slender; anthers dorsifixed or basifixed, dehiscing laterally. Pistil simple or compound, 1-carpellate or of 2–many carpels, superior, free or variously united, each locule with a single basal ovule; stigmas linear to capitate, sessile or borne on a short style. Fruit of 1 or more free or united carpels, fleshy or dry (rarely winged), subtended by the persistent perianth. Seed subglobose to lenticular or reniform, erect, embryo annular or curved; endosperm present. U.S.A., Mexico, Pantropics (mostly South America), also eastern Mediterranean area; ca. 25 genera and ca. 125 species, 6 genera and 10 species in the flora area. Key to the Genera of Phytolaccaceae 1. 1. 2(1). 2. 3(1). 3. 4(3). 4. 5(4).

5.

Scandent or climbing shrubs or woody lianas ........................................... 2 Herbs or decumbent or erect shrubs ......................................................... 3 Thorns present; tepals 5; stigma pappilose; stamens 15–65; fruit winged (samara) .................................................................................... 5. Seguieria Thorns absent; tepals 4; stigma penicillate; stamens 8–25; fruit a globose fleshy drupe ........................................................................ 6. Trichostigma Decumbent or sometimes erect herbs to 40 cm tall; fruits tuberculate or spiny ........................................................................................... 1. Microtea Erect herbs or small subshrubs; sometimes woody at the base; fruits smooth .................................................................................................... 4 Tepals 5, glabrous externally; ovary 5–16-carpellate ............... 3. Phytolacca Tepals 4, pilose externally; ovary 1-carpellate .......................................... 5 Plants with a strong odor of garlic; stipules present; tepals linear, pubescent basally without; stamens 8; pedicels ca. 1 mm long; fruit an achene .................................................................................................... 2. Petiveria Plants without odor of garlic; stipules absent; tepals obovate-oblong, pilose without; stamens 4; pedicels 4–8 mm long; fruit a globose red berry ....................................................................................................... 4. Rivina

1. MICROTEA Sw., Prodr. 4: 53 1788. Small annual herbs, erect, prostrate, or decumbent. Stems usually somewhat succulent and glabrous. Leaves alternate and usually small, petiolate or subsessile;

Microtea 673

stipules absent or represented by minute tubercles; blade simple and entire, glabrous. Inflorescences terminal and axillary, 1–several at a node, racemose, spicate, or paniculate, with floral bracts and with or without bracteoles. Flowers very small, bisexual. Tepals 5(4), free above a thickened receptacle, persistent in fruit. Stamens (3–)5(–9) usually alternating with the tepals, free or united near the base; anthers dorsifixed; thecae globose and dehiscing laterally. Ovary with 1 locule and 1 ovule; style short; stigmas 2 or 3. Fruit fleshy or hard, smooth to tuberculate or spiny, the pericarp adhering to the seed. Seed erect; embryo curved; endosperm fleshy. Neotropics (one species in temperate North America); ca. 9 species, 2 in Venezuela, both in the flora area. Key to the Species of Microtea 1.

1.

Decumbent herbs; leaves elliptic to ovate; inflorescence 3–5 cm long; pedicels 0.8–1 mm long or absent; bracteoles absent; stamens 5 ...................................................................................................... M. debilis Erect herbs; leaves linear-lanceolate; inflorescence 8–12 cm long; pedicels 2–5 mm long; bracteoles 2; stamens 8 ............................... M. maypurensis

Fig. 573. Microtea debilis

674

P HYTOLACCACEAE

Microtea debilis Sw., Prodr. 53. 1788. —Albajaca, Escobilla. Decumbent herb to 45 cm tall; flowers white. Wet areas, secondary vegetation, edges of towns, near sea level to 300 m; Delta Amacuro (Boca de Macareo, Sacupana), Amazonas (Maroa, upper Río Orinoco, near San Carlos de Río Negro, San Fernando de Atabapo). Aragua, Barinas, Carabobo, Distrito Federal, Falcón, Lara, Miranda, Portuguesa, Sucre, Táchira, Yaracuy, Zulia; Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 573.

Microtea maypurensis (H.B.K.) G. Don in Loudon, Hort. Brit. 98. 1830. —Ancistrocarpus maypurensis H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 186, pl. 122. 1817. —Ceratoccoca maypurensis (H.B.K.) Willd. ex Schult., Syst. Veg. 6: 800. 1820. Microtea maypurensis var. angustifolia Moq. in A. DC. Prod. 13(2): 18. 1849. Erect herb to 40 cm tall, with many stems. Rocky riversides, 50–200 m; Amazonas (Raudal de Maipures). Apure; Colombia, Guyana, Suriname, Peru, Brazil, Bolivia, Paraguay.

2. PETIVERIA L., Sp. Pl. 342. 1753. Erect herbs or subshrubs, minutely puberulent or glabrescent, tissue often with the odor of garlic. Stems with longitudinal ridges. Leaves spirally alternate or distichous; stipules paired and lateral; petiole grooved adaxially; blade acute to acuminate. Inflorescences slender, elongate, racemes lax, subtended by short bracts, terminal or axillary and solitary or several from a node. Flowers short-pedicellate or subsesssile, actinomorphic, small, bisexual. Perianth of 4 parts, free, equal and acute, narrow, spreading at anthesis, persisting and erect in fruit, stiff and dry. Stamens 4–9, alternate with the perianth parts or irregularly placed, shorter than perianth; anthers dorsifixed and linear, 2-cleft at apex and base. Ovary 1-locular, oblong, tomentose, with 4 or 6 reflexed spines near the apex; stigma solitary, sessile and penicillate, ovule erect. Fruit a long, narrow, cuneate achene, laterally compressed with 2 slightly flattened carinate sides, apically 2-lobed with 4 or 6 recurved spines, pericarp adhering to the seed. Seeds erect and linear. Neotropics and subtropics; 2 species, 1 in Venezuela. Petiveria alliacea L., Sp. Pl. 342. 1753. —Anaucé, Mapurite. Petiveria foetida Salisb., Prodr. Stirp. Chap. Allerton 214. 1796. Herb or small shrub, with a strong odor of garlic. Dry or wet forest understory, near sea level to 300 m; Delta Amacuro (Pedernales), Bolívar (La Paragua, Tumeremo). Aragua, Cojedes, Carabobo, Distrito Federal, Falcón, Lara, Mérida, Miranda, Portuguesa, Sucre,

Yaracuy, Zulia; U.S.A (Florida), Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, also cultivated and naturalized in the Old World. ◆Fig. 574. Petiveria alliacea is well known as “mapurite,” and all parts of the plant have a reputation to treat cancer.

3. PHYTOLACCA L., Sp. Pl. 441. 1753. Herbs or subshrubs, rarely trees. Stems erect or clambering, often somewhat succulent, glabrous or sparsely puberulent. Leaves spirally alternate, simple, sessile or petiolate; stipules absent; pinnately veined. Inflorescences at first terminal but soon becoming leaf-opposed or extra-axillary, spikes, racemes, or raceme-like panicles, pedicels subtended by narrow bracts and often with 1 or 2 small bracteoles along the length of the pedicel. Flowers small and actinomorphic, bisexual or unisexual. Perianth of 1 whorl of 5 usually free tepals, the tepals greenish or white to pink and purple. Androecium of 5–30 stamens in 1 or 2 whorls; filaments free or

Petiveria

Fig. 574. Petiveria alliacea

675

676

P HYTOLACCACEAE

united at the base and borne on a disk; anthers dorsifixed. Ovary usually broader than long, composed of a whorl of 5–16 partly or completely united carpels, 5–16 locules with 1 erect ovule each. Fruit usually a fleshy, juicy berry, broader than long with a centrally depressed apex, 5–16-ribbed or -parted. Seed compressed; embryo curved. Tropics and subtropics (mostly in the Americas), warm-temperate North America; ca. 30 species, 9 in Venezuela, 3 of these in the flora area. Key to the Species of Phytolacca 1. 1. 2(1). 2.

Inflorescence 30–55(–80) cm long; ovary globose, 10–16-carpellate .................................................................................................. P. rivinoides Inflorescence 5–25 cm long; ovary subglobose, 6–9-carpellate ................. 2 Pedicels 3.5–4 mm long; tepals oblong-elliptic; carpels free at the apex ....................................................................................................... P. rugosa Pedicels 5–7 mm long; tepals elliptic to oval-ovate; carpels completely united ...................................................................................... P. thyrsiflora

Fig. 575. Phytolacca rivinoides

Rivina 677

Phytolacca rivinoides Kunth & Bouché, Index Seminum (Berlin). 1848: 15. 1849. —Aurosá (Pemón), Carurú (Baré), Chipachi, Nirguo rebalsero, Pebajú (Guahibo), Pescuezo de pava. Succulent herb to subshrub 1–3 m tall; flowers white to rose; leaves edible. Disturbed forests, open areas in secondary forests, edges of towns, old plantations, 100–1500 m; Delta Amacuro (Serranía de Imataca), Bolívar (widespread), Amazonas (Puerto Ayacucho, San Carlos de Río Negro, San Fernando de Atabapo, Río Cataniapo, Río Matacuni, Río Yatúa). Apure, Aragua, Carabobo, Distrito Federal, Falcón, Lara, Mérida, Miranda, Portuguesa, Sucre, Trujillo, Yaracuy, Zulia; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 575.

Phytolacca rugosa A. Braun & Bouché, Index Seminum (Berlin). 1851: 13. 1852. Herb or subshrub 1–4 m tall; flowers white to pink. Edges of wet forests, ca. 1200 m; Bolívar (slopes of Auyán-tepui). Distrito Federal, Lara, Mérida, Portuguesa, Trujillo, Táchira; Mexico, Central America, Colombia, Ecuador, Peru. Phytolacca thyrsiflora Fenzl ex J.A. Schmidt in Mart., Fl. Bras. 14(2): 343. 1872. Herb or subshrub 1–3 m tall; flowers white to pink. Lower montane wet forests, open areas, 700–1000 m; Bolívar (Cerro Bolívar, El Paují). Aragua; Hispaniola, Guyana, French Guiana, Peru, Brazil, Bolivia, Paraguay.

4. RIVINA L., Sp. Pl. 121. 1753. Perennial herbs or subshrubs to 1 m tall, sometimes woody near the base, glabrous or minutely puberulent. Leaves spirally alternate, on slender petioles, simple and entire, pinnately veined; stipules absent. Inflorescences erect, solitary, open racemes, terminal or becoming axillary, usually minutely puberulent; pedicels subtended by small, narrow, deciduous bracts, 1 or 2 minute bracteoles often present at the apex of the pedicel beneath the perianth. Flowers small, actinomorphic, bisexual. Perianth of 4 subequal tepals in 2 decussate pairs, slightly imbricate in bud, oblong-spatulate and usually rounded at the apex, corolla-like, becoming white, persisting and enlarging only slightly in fruit. Stamens 4, free, alternating with the perianth parts and inserted on the slightly elevated receptacle; disk hypogynous, slightly shorter than the perianth; anthers elongate and dorsifixed, extrorse-lateral. Pistil simple, ovary with 1 locule and 1 basal ovule; style terminal or subterminal, often curved. Fruit a globose, red drupe with pericarp adherent to the seed, subtended by the persistent perianth. Seed lenticular, often with a covering of short hairs; embryo annular. Southern U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay, also cultived and naturalized in India, Java, and New Caledonia; 1 species. Rivina humilis L., Sp. Pl. 122. 1753. —Cuentica roja. Herb or subshrub 0.5–2 m tall, usually woody at the base; fruit red or orange. Distribution as in genus; 3 varieties, 1 in the flora area.

R. humilis var. humilis Open areas, secondary vegetation, 100– 300 m; Delta Amacuro (Sacupana), Bolívar (near El Manganeso, near El Manteco). Aragua, Carabobo, Cojedes, Distrito Federal. Falcón, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Yaracuy, Zulia. ◆Fig. 576.

678

P HYTOLACCACEAE

Fig. 576. Rivina humilis var. humilis

5. SEGUIERIA Loefl., Iter Hispan. 191. 1758. Trees, scandent shrubs, or lianas. Branches terete or subterete, glabrous, striate by small furrows, younger branches round or angled, rarely sparsely pubescent, usually with paired thorns above the bases of the petioles, sometimes minute or absent. Leaves alternate, petiolate, entire, glabrous or sparsely pubescent in young leaves. Inflorescences axillary or terminal, racemes or panicles; bracts present in the axis of the inflorescence, sometimes leaf-like, membranous, lanceolate to triangular. Flowers pedicellate. Tepals 5, imbricate, reflexed in fruit. Stamens 15–70; anthers linear, dorsifixed, extrorse, opening by longitudinal slits. Ovary superior, 1-carpellate, 1-locular, subglobose or laterally compressed basally; stigma papillose; ovule 1, basal, campylotropous. Fruit winged, globular to pear-shaped. Seed 1, erect, glabrous, embryo curved. Neotropics; ca. 6 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Seguieria 1. 1.

Leaves chartaceous to subcoriaceous; filaments 5–6 mm long; samara green or yellow when dried ....................................................... S. aculeata Leaves coriaceous; filaments < 4.5 mm long; samara black when dried ............................................................................................. S. macrophylla

Seguieria aculeata Jacq., Select. Stirp. Amer. Hist. 170. 1763. Scandent shrub, with scandent or decumbent branches; thorns recurved. Semidecid-

uous to evergreen lowland forests, 200–300 m; Bolívar (El Temblador on lower Río Caura, lower Río Botanamo, lower Río Caroní). Aragua, Carabobo, Distrito Federal, Falcón,

Seguieria 679

Fig. 577. Seguieria aculeata

Fig. 578. Seguieria macrophylla

680

P HYTOLACCACEAE

Mérida, Miranda, Táchira, Zulia; Panama, Colombia, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 577. Seguieria macrophylla Benth., Trans. Linn. Soc. London 18: 235. 1841. Seguieria cordata Britton, Bull. Torrey Bot. Club 48: 331. 1921.

Tall woody liana or climbing shrub; thorns recurved. Semideciduous to lowland evergreen forests, near sea level to 500 m; Delta Amacuro (east of Caño Corial, Piacoa, Sacupana, San Antonio), Bolívar (north of Cerro Ichún). Apure, Barinas, Miranda, Zulia; Panama, Colombia, Guyana, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 578.

6. TRICHOSTIGMA A. Rich. in Sagra, Hist. Phys. Cuba, Bot. Pl. Vasc. 627. 1845. Shrubs or woody climbers, usually glabrous. Leaves spirally alternate, petiolate, entire; stipules minute or caducous. Inflorescences terminal or axilary, solitary, elongate, lax racemes. Flowers pedicellate (pedicels often with small bracts and 2 minute bracteoles, deciduous), bisexual. Perianth of 4 subequal parts in 2 whorls, broadly imbricate in bud, reflexed in fruit. Stamens 8–28, free, borne on the elevated receptacle, disk present; filaments free; anthers dorsifixed, longer than wide. Pistil simple with sessile or subsessile penicillate stigma, ovary subglobose, 1-carpellate, 1-locular with 1 ovule. Fruit a globose drupe, reddish or purple. Seed with annular embryo; testa crustaceous. Neotropics; ca. 3 species, 2 in Venezuela, 1 of these in the flora area.

Trichostigma octandrum (L.) H. Walter in Engl., Pflanzenr. IV. 83(Heft 39): 109. 1909. —Rivina octandra L., Cent. Pl. II, 9. 1756. Woody climbing vine; flowers white to whitish green; fruit purple or red. Moist forests, 50–200 m; Delta Amacuro (Punta Prada, Río Grande). Apure, Aragua, Barinas, Carabobo, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Portuguesa, Táchira, Yaracuy, Zulia; U.S.A. (Florida), Mexico, Central America, West Indies, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 579.

Fig. 579. Trichostigma octandrum

P I P E R A C E A E 681

PIPERACEAE by Julian A. Steyermark and Ricardo Callejas-Posada Erect or rarely climbing shrubs, small trees, or succulent, herbaceous, terrestrial or epiphytic plants. Prophyll (in Piper) resembling a solitary bract. Leaves alternate, opposite, verticillate, or basal, often aromatic when crushed, petiolate or sessile; petioles often with sheathing foliar bases and often with stipule-like margins; blades simple, entire, without lobes or lobed only at base, glabrous or pubescent, palmately or pinnately veined, or pliveined (subpinnate but with strong lateral veins arising from the basal 1/2 or 2/3 of midrib and strongly curving toward apex; acrodromous), sometimes peltate. Inflorescence axillary, terminal, opposite the leaf with a solitary spike, or racemose or compound, umbellate or paniculate when compound, white, cream, green, lavender, or sometimes roseate or reddish, the rachis usually thick and fleshy, glabrous, papillate, or pubescent. Flowers numerous, usually close together in spiral or ringlike arrangement, without a perianth, bisexual (all American species), in the axils of solitary, small, peltate or subpeltate, glabrous, pubescent, or fimbrillate bracts, the bracts enlarged apically with a triangular, rounded, or lunulate apex. Stamens usually 1(2–4)–6, free and/or arising near base of ovary; filaments basally articulate, usually very short; anthers with 1, 2, or 4 thecae, laterally dehiscent, often in a horizontal plane, the connective sometimes prominent and forming a disk similar to a gland at apex. Ovary superior, sessile or rarely stipitate, unilocular with 1 basal ovule; stigmas usually 1–4, sessile or on a short to elongate style. Fruit a drupe, dry or fleshy. Seed small, solitary; endosperm scanty, perisperm abundant; embryo very small. Tropics and subtropics worldwide; 4 genera and ca. 2500 species, 2 genera and 124 species in the flora area. Key to the Genera of Piperaceae 1.

1.

Plants herbaceous, succulent, terrestrial or more often epiphytic; nodes usually not swollen or thickened; leaves alternate, opposite, verticillate, or basal; floral bracts usually round and flat at apex, orbicular as seen from above, rarely puberulent; stigma 1, often minutely papillate or fimbriate ............................................................................. 1. Peperomia Plants usually woody, sometimes climbing, rarely epiphytic; nodes usually swollen or thickened; leaves alternate; floral bracts (when viewed from above) usually triangular, concave, inflexed, or narrowed below into a pedicel, triangular or deltoid as seen from above, usually fimbriate marginally at apex or less frequently glabrous throughout; stigmas (2)3(4), fimbriate .............................................................................. 2. Piper

1. PEPEROMIA Ruiz & Pav., Fl. Peruv. Prodr. 8, pl. 2. 1794. Herbaceous annuals or perennials, terrestrial, saxicolous, or more frequently epiphytic, usually succulent. Stems erect, spreading, creeping, pendent, or climbing. Leaves usually petiolate; blades entire, sometimes peltate, glabrous or pubescent, often with yellowish or black gland-like dots, frequently with pellucid dots when viewed by transmitted light. Inflorescence a simple solitary spike or compound with numerous paniculate spikes, white, cream, or green, erect, pendent, or recurved, the

682

P IPERACEAE

rachis glabrous, papillate, or puberulous. Flowers crowded or loosely arranged, sessile, superficial, or arising from slight depressions, subtended by an orbicular or peltate, glabrous or fimbriate, often gland-dotted bract. Stamens 2, early deciduous; anthers 2-locular, subglobose or ellipsoid; filament generally shorter than anther. Stigma simple or sometimes cleft but usually fimbrillate, often borne in the center of translucent tissue which develops into a beak, sessile or stylose, the apex rounded and mammiform, more often oblique and subapical on abaxial side of the usually oblique beak, or at base of a rostrate beak. Fruit 1-seeded, drupe-like with a thin pericarp and a slightly hard endocarp, basally to subbasally attached to rachis, sessile or pedicellate, or frequently articulated to a persistent pseudopedicel developing at late maturity, globose, ovoid, ellipsoid, subcylindric, or obovoid, the surface smooth or verrucose, usually adhesive-sticky. Pantropics; ca. 1100 species, 125 in Venezuela, 51 of these in the flora area. Key to the Species of Peperomia 1.

Leaves ± peltate (leaf margin continuous above point of union with petiole) .......................................................................................................... 2 1. Leaves not peltate (leaf margin ending at its junction with petiole) ....... 5 2(1). Inflorescence with usually 2 or 3 spikes ...................................... P. distachya 2. Inflorescence simple, with 1 spike ............................................................. 3 3(2). Leaves pinnately veined, the lateral veins extending upward along greater part of midvein ............................................................. P. maculosa 3. Leaves palmately veined or slightly pliveined, with all or most of the veins originating in or near the peltate point of union with petiole .... 4 4(3). Petiole and lower surface of leaf blade pubescent, at least at junction with petiole, or along length of the principal veins ................. P. hernandiifolia 4. Petiole and lower surface of leaf blade glabrous ............................. P. venusta 5(1). Leaves mainly opposite or verticillate ....................................................... 6 5. Leaves mainly alternate (although sometimes opposite at some nodes) .............................................................................................................. 24 6(5). Leaves mainly opposite (some nodes with 1, 3, or 4 leaves) ..................... 7 6. Leaves partly to totally verticillate (some species with opposite leaves on fertile stems and verticillate leaves on sterile stems) ........................ 16 7(6). All or most leaves of sterile and fertile stems opposite ............................ 8 7. Some leaves of sterile and fertile stems alternate, or in groups of 3, 4, or more ...................................................................................................... 12 8(7). Some inflorescences compound, with 2 or 3 spikes arising from a peduncle; foliar leaves oblanceolate ...................................P. quadrangularis 8. Inflorescence simple, with 1 spike on a peduncle; leaves suborbicularovate, elliptic, rhombic, or sometimes obovate ..................................... 9 9(8). Peduncle usually with short spreading pubescence; stems and petioles generally with spreading pubescence; leaves soft, membranous ....................................................................................................... P. blanda 9. Peduncles either glabrous or with short appressed pubescence; stems and petioles glabrous or with appressed pubescence; leaves coriaceous or thick-fleshy ........................................................................................... 10 10(9). Leaves mainly rounded or obtuse at apex, nearly as wide as long or only

Peperomia 683

10. 11(10). 11. 12(7). 12. 13(12). 13. 14(13). 14. 15(12). 15. 16(6). 16. 17(16). 17. 18(17).

18. 19(16). 19. 20(19). 20.

21(19). 21. 22(21). 22. 23(22). 23. 24(5). 24.

slightly longer than wide, 1–3(–3.5) cm long; petioles mostly 1–5 mm long .................................................................................. P. quadrangularis Leaves mainly narrowed to an acute apex, slightly to much longer than wide, (2–)3–6.5 cm long; petioles generally (2–)4–20 mm long .......... 11 Leaves rather abruptly narrowed at apex and cuneately or acutely narrowed at base ........................................................................... P. angustata Leaves gradually narrowed to apex and mainly rounded, obtuse, or subacute at base ............................................................................ P. rotundata Stem mainly glabrous .............................................................................. 13 Stem ± pubescent ..................................................................................... 15 Leaves pliveined .................................................................. P. guaiquinimana Leaves either palmately 3–5-veined or with only 1 midvein faintly visible .............................................................................................................. 14 Leaves rounded at apex ................................................................... P. cladara Leaves mainly acute or subacute at apex ............................... P. dendrophila Leaves emarginate at apex .............................................................. P. cladara Leaves not emarginate at apex .................................................... P. rotundata Stems glabrous ......................................................................................... 17 Stems with various degrees of pubescence, sometimes only papillatepuberulent ............................................................................................ 19 Leaves 1.5–4 cm wide and 2.5–9 cm long .................................... P. angustata Leaves 0.2–1.2 cm wide and 0.4–1.5 cm long .......................................... 18 Leaves widest at middle, narrowing toward apex, almost as wide as long or to 1.5 times longer than wide; rachis of inflorescence minutely pilosulous on ribs ................................................................... P. tetraphylla Leaves widest above middle, 2–5 times longer than wide; rachis of inflorescence glabrous ....................................................................... P. galioides Leaves 15–40 mm wide ............................................................................ 20 Leaves 1.5–13(–15) mm wide ................................................................... 21 Leaves dimorphic, the lower obovate and rounded at apex; pliveined; stems moderately crisp-villosulous ......................................... P. yutajensis Leaves ± uniform, obtusely acute to obtusely subacuminate at apex; inconspicuously palmately veined; stems at most sparsely papillatepuberulent ................................................................................ P. angustata Stems and leaves exfoliating with a gray-white scaliness; leaves strongly revolute and marginally folded lengthwise ............................... P. loxensis Stems and leaves not exfoliating as described above; leaves neither revolute nor folded lengthwise ................................................................... 22 Leaves widest above middle, not narrowing at apex .................... P. galioides Leaves widest at middle, narrowing toward apex .................................. 23 Peduncle 0.3–0.7 cm long; spike 1 mm or less wide, commonly with pseudopedicels; petiole almost absent, 0.5–1 mm long ............. P. quaesita Peduncle 1–3 cm long; spike 1.5–2.5 mm wide, without pseudopedicels; petiole 1–5 mm long ............................................................... P. tetraphylla Leaves either pliveined or pinnately veined, some or all of the lateral veins arising above very base of leaf blade ......................................... 25 Leaves palmately veined, the lowest principal veins arising together at very base of leaf blade ......................................................................... 54

684

P IPERACEAE

25(24). Inflorescence compound, with 2 or more spikes arising from principal axis ....................................................................................................... 26 25. Inflorescence a solitary spike or with 2 or more spikes arising from the same base ............................................................................................. 39 26(25). Base of leaf obtuse, rounded, or subcordate ............................................ 27 26. Base of leaf acute ..................................................................................... 31 27(26). Leaves pinnately veined with 5–7 veins on each side; stems mainly 10– 15 mm thick .................................................................................. P. striata 27. Leaves pliveined, or if pinnately veined, then the stems 2–8 mm thick, or very elongate, pendent, or creeping .................................................... 28 28(27). Plants terrestrial, the stems rooting at nodes, creeping; uppermost pair of veins arising 3–11 mm above base of leaf blade ................................. 29 28. Plants epiphytic, the stems pendent, not creeping; uppermost pair of veins arising 10–50 mm above base of leaf blade ............................... 30 29(28). Primary and secondary peduncles glabrous; uppermost pair of veins arising 3–5 mm above the base of leaf blade; beak of fruit much shorter than body ................................................................................... P. distachya 29. Primary and secondary peduncles at least partly puberulous; uppermost pair of veins arising 5–11 mm above base of leaf blade; beak of fruit conspicuous, nearly as long as body .................................................. P. gentryi 30(28). Petioles eciliate ...................................................................... P. macrostachya 30. Petioles ciliate ................................................................................ P. elongata 31(26). Inflorescence paniculate, the spikes subverticillate on an elongate axis; spikes 1–1.5(–2) cm long ................................................ P. pernambucensis 31. Inflorescence not as above; spikes 2–25 cm long .................................... 32 32(31). Leaves 5–7 times longer than wide, narrowly lanceolate or oblanceolate ................................................................................................... P. lancifolia 32. Leaves 1.5–3.5 times longer than wide, elliptic-ovate, oblong-, or ovateelliptic, obovate, or subelliptic ............................................................. 33 33(32). Stems elongate, pendent, creeping, or rooting, 1 m or more long; mature fruits cylindric or ovoid-cylindric, 1–3 mm long, not rostrate, the apex obliquely shield-shaped; stigma central .............................................. 34 33. Stems erect or ascending from a decumbent or rooting base, if creeping or rooting then < 1 m long; mature fruits subglobose, ovoid, or cylindric, 0.7–1.8 mm long, short- to long-rostrate; stigma near base of beak .............................................................................................................. 35 34(33). Petioles ciliate; floral bracts mainly not fimbrillate, flat, uniformly opaque; mature fruits 1–1.8 mm long; spikes generally solitary, rarely 2 ....................................................................................... P. elongata 34. Petioles eciliate; floral bracts ± fimbrillate, densely papillose, umbonate, scarious throughout; mature fruits 2–3 mm long; spikes usually 2– 4, sometimes solitary ......................................................... P. macrostachya 35(33). Apex of leaf acute to acuminate; leaves membranous upon drying ....... 36 35. Apex of leaf rounded or obtuse to subacute; leaves thick-coriaceous upon drying ................................................................................................... 37 36(35). Stems usually completely glabrous; inflorescence bearing 1–3 spikes; leaves 4–9 cm long, 1.5–2.6 times longer than broad; stems 0.15–0.35 m tall; body of fruit shorter than or equaling beak, the beak 1–1.5 mm long ................................................................................................ P. alpina

Peperomia 685

36.

37(35).

37.

38(37).

38.

39(25). 39. 40(39). 40. 41(40). 41. 42(41).

42.

43(40). 43. 44(43). 44. 45(44). 45. 46(44).

Stems, at least on upper internodes, appressed-pubescent; inflorescence bearing 4–11 spikes; leaves 15–23 cm long, mainly (2–)2.5–3 times longer than broad; stems 0.5–0.9 m tall; body of fruit much longer than beak, the beak 0.3 mm long .......................................................... P. striata Leaf apex usually usually rounded or obtuse; apex of beak of mature fruit strongly bent or recurved, basal part of beak slender; floral bracts 0.2– 0.4(–0.5) mm diameter ............................................................ P. obtusifolia Leaf apex usually tapering above the middle to a subacute or broadly acute apex; apex of beak of mature fruit slightly curved, basal part of beak manifestly swollen or thickened; floral bracts 0.5–0.7 mm diameter ....................................................................................................... 38 Secondary upper peduncle immediately below inflorescence usually glabrous or glabrescent; spikes often in groups of 2 or 3, or solitary; leaves greenish gray when dry; spikes white or green when dry ............................................................................................ P. magnoliifolia Secondary upper peduncle immediately below inflorescence usually puberulent or glabrous; spikes in groups of 6–20, never solitary; leaves green to orange when dry; spikes deep orange when dry .................. P. uaupesensis Leaf blades, petioles, and stems densely black-dotted when dried ..................................................................................................... P. glabella Leaf blades not as above .......................................................................... 40 Leaves rounded or obtuse at apex ........................................................... 41 Leaves mainly subacute to acute or acuminate at apex ......................... 43 Peduncle longer than mature spike; mature spikes generally 2–4 cm long ............................................................................................. P. haematolepis Peduncle shorter than mature spike; mature spikes generally (3.5–)5– 26 cm long ............................................................................................ 42 Apex of beak of mature fruit slightly curved; basal part of beak manifestly swollen or thickened; floral bracts 0.5–0.7 mm diameter; secondary upper peduncle immediately below inflorescence usually glabrous or glabrescent; spikes often in groups of 2 or 3, or solitary ....... P. magnoliifolia Apex of beak of mature fruit strongly bent or recurved; basal part of beak slender, less conspicuously swollen; floral bracts 0.2–0.4(–0.5) mm diameter; secondary upper peduncle sparsely to moderately minutely puberulent, sometimes glabrescent or glabrous; spikes often solitary, also in groups of 2 or 3 ............................................................ P. obtusifolia Leaf blade rounded, cordate, or broadly obtuse at base ......................... 44 Leaf blade mainly acute, subacute, or cuneate at base .......................... 47 Leaves pubescent on upper surface along midvein and/or on principal veins; plants creeping, rooting at nodes; fruit shortly rostrate ......... 45 Leaves completely glabrous on upper surface; plants epiphytic, mainly pendent; fruit not rostrate ................................................................... 46 Leaves 5-veined, only the midvein pubescent on upper surface; fruit with a short beak < 0.5 mm long ...................................................... P. yatuensis Leaves 7–11-veined, all the veins of the upper surface pubescent; fruit with a beak 0–5 mm long ................................................. P. fundacionensis Petioles eciliate; floral bracts ± fimbrillate, densely papillose, umbonate, scarious throughout; mature fruits 2–3 mm long; spikes usually 2– 4, sometimes solitary ........................................................ P. macrostachya

686

P IPERACEAE

46.

47(43). 47. 48(47). 48. 49(48). 49. 50(49).

50.

51(48).

51.

52(47). 52. 53(52). 53. 54(24). 54. 55(54). 55. 56(54). 56. 57(56). 57. 58(57). 58. 59(58).

Petioles ciliate; floral bracts mainly not fimbrillate, flat, uniformly thickened; mature fruits 1–1.8 mm long; spikes generally solitary, rarely 2 .................................................................................................. P. elongata Leaves pinnately veined the greater part of their length from base nearly to apex .................................................................................................. 48 Leaves pliveined, the upper veins arising below middle half ................ 52 Stigma apical or subapical; ovary and fruit not beaked ......................... 49 Stigma situated at base of beak; fruit terminating in a slender, elongate beak ...................................................................................................... 51 Plant terrestrial, with erect stem; stigma apical; fruit terminating in an abrupt, flattened, or truncate apex ........................................ P. acuminata Plant epiphytic, usually associated with ant nests; stigma subapical; fruit terminating in an obliquely shield-shaped beak ................................ 50 Petioles eciliate; floral bracts ± fimbrillate, densely papillose, umbonate, scarious throughout; mature fruits 2–3 mm long; spikes usually 2– 4, sometimes solitary ......................................................... P. macrostachya Petioles ciliate; floral bracts mainly not fimbrillate, flat, uniformly thickened; mature fruits 1–1.8 mm long; spikes generally solitary, rarely 2 ....................................................................................... P. elongata Apex of beak of mature fruit slightly curved; basal part of beak manifestly swollen or thickened; floral bracts 0.5–0.7 mm diameter; secondary upper peduncle immediately below inflorescence usually glabrous or glabrescent; spikes often in groups of 2 or 3, or solitary ...... P. magnoliifolia Apex of beak of mature fruit strongly bent or recurved; basal part of beak slender, less conspicuously swollen; floral bracts 0.2–0.4(–0.5) mm diameter; secondary upper peduncle sparsely to moderately minutely puberulent, sometimes glabrescent or glabrous; spikes often solitary, also in groups of 2 or 3 ............................................................ P. obtusifolia Internodes of stem with a gray, scaly epidermal covering ..................... ......................................................................................... P. guaiquinimana Internodes of stem glabrous .................................................................... 53 All principal veins of leaves arising in lowest 10 mm of leaf base ..... P. alata Upper principal veins of leaves arising (5–)10–60 mm above leaf base .................................................................................................... P. elongata Both surfaces of leaf blades, petioles, and stem black-dotted when dried .............................................................................................................. 55 Not as above ............................................................................................. 56 Stems glabrous, except for ciliate decurrent angles below the petioles; leaves glabrous except below the ciliate petioles ....................... P. glabella Stems ± pubescent between the ciliate decurrent angles below petioles; leaves often pubescent ........................................................ P. venezueliana Leaves 2–10 mm long; stem nodes glabrous ........................................... 57 At least some leaves > 10 mm long; stem nodes often pubescent .......... 62 Leaves ± emarginulate at apex ............................................................... 58 Leaves not emarginulate at apex ............................................................ 60 Leaves longer than wide ................................................................... P. tenella Leaves wider than long or as wide as long .............................................. 59 Petioles and stems minutely hirtellous with spreading trichomes; leaf

Peperomia

59. 60(57). 60. 61(60).

61.

62(56). 62. 63(62).

63.

64(63). 64. 65(64). 65. 66(65). 66. 67(65). 67. 68(62). 68. 69(68). 69. 70(69). 70. 71(70).

71.

687

blades not ciliate; uppermost leaves opposite or alternate; stems ascending to suberect ................................................................ P. delascioi Petioles and stems glabrous; leaf blades ciliate; all leaves alternate; stems creeping and rooting ............................................................. P. emarginella Most leaves orbicular or suborbicular; petioles (2–)4–13 mm long ............................................................................................... P. rotundifolia Most leaves longer than wide, or if suborbicular then narrowed at apex; petioles 1–4(–5) mm long ..................................................................... 61 Uppermost leaves always alternate, widest near or below the middle; fruits pedicellate, the body > 1 mm long, > 2 times longer than wide; internodes glabrous to pilose ........................................................ P. tenella Uppermost leaves often opposite or whorled, widest near or above the middle; fruits not pedicellate, shorter than 0.9 mm, < 2 times longer than wide; internodes pilose ............................................... P. jamesoniana Leaves emarginate or emarginulate at apex .......................................... 63 Leaves not emarginate or emarginulate at apex .................................... 68 Leaves broadly rounded and not narrowed above middle; stems strongly longitudinally grooved, constricted at the nodes, the epidermis scalyexfoliating; upper surface of leaf drying wrinkled, the lateral veins obsolete or nearly so .................................................................... P. cladara Leaves narrowed above middle to the apex; stems neither longitudinally grooved nor constricted, without a scaly exfoliating epidermis; upper surface of leaf not drying wrinkled, the lateral veins evident ........... 64 Upper leaves often opposite; spikes 2.9–3.5 mm diameter; leaves mainly (2.5–)4–7 cm long; pseudopedicels frequent ....................... P. dendrophila All leaves alternate; spikes 0.5–3 mm diameter; leaves 0.6–3.8 cm long; pseudopedicels absent ......................................................................... 65 Stem 1.5–2.5(–4) mm thick; leaves 12–38 × 6–15 mm; spikes 1.5–3 mm diameter ............................................................................................... 66 Stem 0.5–1(–1.5) mm thick; leaves 6–18(–21) × 2–6.5(–12) mm; spikes 0.5–1 mm diameter .............................................................................. 67 Fruits sessile; leaves reddish on lower surface ..................... P. purpurinervis Fruits pedicellate; leaves greenish on lower surface ....................... P. tenella Fruit pedicellate; stigma apical; leaves distichous; petioles 1–2 mm long ....................................................................................................... P. tenella Fruit sessile; stigma subapical; leaves not distichous; petioles 3–7 mm long ...................................................................................... P. jamesoniana Leaves slightly to deeply cordate at base ................................................ 69 Leaves neither cordate nor cordulate at base ......................................... 76 Leaves acute, acuminate, or subacute at apex ........................................ 70 Leaves rounded, obtuse, or subobtuse at apex ........................................ 74 Stem erect; fruit not rostrate; stigma apical, mammiform .................... 71 Stem prostrate or creeping; fruit rostrate; stigma subapical, at the base of a slender beak ...................................................................................... 72 Annual plants of disturbed or cultivated ground; leaf blades 1–3.7 cm long, thin-membranous, transparent; peduncle 0.2–0.7 mm thick; fruit longitudinally ribbed ................................................................ P. pellucida Perennial plants of rocky habitats; leaf blades 4–5.2 cm long, mem-

688

P IPERACEAE

72(70). 72. 73(72). 73. 74(69). 74. 75(74). 75. 76(68). 76. 77(76). 77. 78(77). 78. 79(78). 79. 80(78). 80.

81(80).

81.

82(77). 82. 83(82).

83.

branous, opaque; peduncle 1–1.5 mm thick; fruit verruculose ......... ..................................................................................... P. lanceolato-peltata Stems and petioles glabrous; leaf blades conspicuously rugose ................. .................................................................................................. P. spruceana Stems and petioles pubescent; leaf blades scarcely or not rugose ......... 73 Stems and petioles densely retrorsely pubescent; leaves conspicuously cordate ........................................................................................ P. duidana Stems and petioles antrorsely pubescent or crisp-pubescent; leaves scarcely cordulate ........................................................................ P. serpens Stem erect; spikes loosely flowered, 0.5 mm or less wide; annual plants of disturbed or cultivated ground ................................................. P. pellucida Stem prostrate or creeping; spikes densely flowered, 1–2 mm wide; perennial plants of undisturbed or natural habitats ................................... 75 Stems and petioles densely retrorsely pubescent; leaves conspicuously cordate ........................................................................................ P. duidana Stems and petioles antrorsely pubescent or crisp-pubescent; leaves scarcely cordulate ........................................................................ P. serpens Leaves mainly acute or subacute at base ................................................ 77 Leaves mainly rounded, truncate, obtuse, or subobtuse at base ........... 87 Leaves rounded, obtuse, or subobtuse at apex ........................................ 78 Leaves acute, acuminate, or broadly acute at apex ................................ 82 Stem 2–5 mm thick, erect or ascending; peduncles 15–22 mm long; leaves reddish on lower surface ...................................................................... 79 Stem 0.5–2 mm thick, creeping, prostrate, erect, or ascending; leaf blades to 2.1 cm long and to 1.8 cm wide; peduncles to 15 mm long ............. 80 Leaves narrowly ovate to elliptic, rounded or acute at the base but never cordulate; stems greenish; leaf blades to 3.5 cm long ...... P. purpurinervis Leaves broadly ovate to rounded, often cordulate; stems often reddish; leaf blades to 5 cm long ............................................................ P. rotundata Leaves 0.2–0.5(–0.6) cm wide; mature spikes 1.2–3.5 cm long; stigma apical; leaves at least 2 times longer than wide ................ P. jamesoniana Leaves (0.5–)0.6–1.8 cm wide; mature spikes 0.8–6 cm long; stigma subapical or apical; leaves ca. 1.5–3.2 times as long as wide, but if stigma apical then leaves < 2 times longer than wide .................................... 81 Pseudopedicels frequent; stigma subapical; stems 0.5–1 mm thick; mature spikes 1.5–6 cm long; plants of eastern Bolívar and Delta Amacuro states ....................................................................................... P. ouabianae Pseudopedicels not present; stigma apical, mammiform; stems 1–2 mm thick; mature spikes 0.8–1.5(–2) cm long; plants of Sierra de la Neblina .................................................................................................. P. neblinana Stems and leaves (except the ciliolate apex) glabrous ............................ 83 Stems and/or leaves pubescent ................................................................ 85 Leaves thin and delicate, as wide as or wider than long, or only slightly longer than wide; spikes delicate, loosely flowered, 0.5 mm or less wide; stigma apical; plants of disturbed or cultivated ground .......... P. pellucida Leaves fleshy-membranous to coriaceous, obviously longer than wide; spikes densely flowered, 1.5–3 mm wide; stigma subapical; plants of undisturbed or natural habitats .......................................................... 84

Peperomia 689

84(83). Stems usually ± zig-zag in distal portion, usually conspicuously winged; leaves drying ± concolorous; leaves usually distichous, long acuminate at apex, usually markedly acute at base; pseudopedicels usually absent; spikes 1–2 mm thick ........................................................... P. alata 84. Stems neither zig-zag nor winged; leaves drying bicolorous; leaves not distichous, mostly shortly acute with the very apex obtuse or blunt, usually broadly acute or subacute at base; pseudopedicels on rachis present or absent; spikes 2–3.5 mm thick........................... P. dendrophila 85(82). Leaves 3–9(–15) mm wide and stems filiform or very slender, 1 mm or less thick; pseudopedicels frequent ............................................... P. ouabianae 85. Leaves wider or stems 2–5 mm thick; pseudopedicels frequent or absent .............................................................................................................. 86 86(85). Spikes secund, solitary, completely opposite the leaves; spikes 3–5 mm thick, with numerous pseudopedicels; stigma apical ..... P. enantiostachya 86. Spikes not secund, sometimes 2–4 together, not all opposite the leaves; spikes 1–2 mm thick, lacking pseudopedicels; stigma subapical .................................................................................................. P. rotundata 87(76). Stems and leaves (except for apex) glabrous .......................................... 88 87. Stems and/or leaves partly or wholly pubescent .................................... 90 88(87). Leaves 5–17 cm long, thick-fleshy .......................................... P. maypurensis 88. Leaves 0.5–3.5 cm long, membranous or thin-fleshy .............................. 89 89(88). Plants annual; stems erect; spike loosely or sparsely flowered, 0.5–0.6 mm thick; plants of disturbed or cultivated ground ....................... P. pellucida 89. Plants perennial; stems creeping; spike densely flowered, 1–3 mm thick; plants of undisturbed or natural habitats ........................... P. rotundifolia 90(87). Stems and petioles retrorsely pubescent ...................................... P. duidana 90. Stems and petioles not retrorsely pubescent .......................................... 91 91(90). Stems, petioles, and leaves inconspicuously to conspicuously pubescent .............................................................................................................. 92 91. Stems glabrous, glabrescent, or puberulent; leaves minutely puberulent, sometimes abundantly ciliolate along margins, or glabrous .............. 97 92(91). Petioles shorter than leaf blades; stems erect to suberect ..................... 93 92. Petioles frequently longer than leaf blades; stems creeping or prostrate .............................................................................................................. 94 93(92). Stem 3–5 mm thick; leaf blades to 5 cm long and to 3.2 cm wide; peduncles to 22 mm long; stigma subapical ............................................. P. rotundata 93. Stem 0.5–2 mm thick; leaf blades to 1.8 cm long and to 1.8 cm wide; peduncles to 9 mm long; stigma apical, mammiform ............. P. neblinana 94(92). Peduncles shorter than spikes; peduncles without bracts; leaves (5–)6– 16 × (4–)6–15 mm ................................................................. P. rotundifolia 94. Peduncles equaling or longer than spikes; peduncles with 1 or 2 bracts; leaves 7–40 × 6–40 mm ........................................................................ 95 95(94). Petioles pilosulous; upper surface of leaf with only the veins pilosulous, the trichomes lax, not appressed; plants of ca. 2500 m elevation ......................................................................................... P. marahuacensis 95. Petioles and upper surface of leaf both appressed-puberulent; plants of near sea level to 1000 m elevation ...................................................... 96 96(95). Leaves mainly obtuse to almost rounded at apex, 0.7–2.3(–2.7) × 0.8–

690

P IPERACEAE

2.5(–2.7) cm; spikes 0.7–1 cm long; upper surface of leaf scarcely or not sulcate along length of the 5 veins (best seen in living plant) ...................................................................................................... P. serpens 96. Leaves shortly narrowed to a subacute or broadly acute apex, mainly 2– 4 × 2–4 cm; spikes 3–5.5 cm long; upper surface of leaf conspicuously sulcate along length of the 7 veins ............................................ P. urocarpa 97(91). Leaves fleshy-coriaceous, ± thick when dried, 1.5–4 × 1.5–3 cm; leaf margins densely appressed-ciliate, the ciliation extending downward as 2 lines on petiole; spikes 5–12 cm long, 2 mm thick ...................... P. celiae 97. Leaves thinner, submembranous, 1–1.8 × 0.7–1.8 cm; ciliation of leaf not extending downward on petiole; spikes 2 cm long, 1 mm thick .................................................................................................. P. neblinana Peperomia acuminata Ruiz & Pav., Fl. Peruv. 1: 32, pl. 51, fig. a. 1798, non (L.) Dahlst. 1900. Peperomia pyrifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 61. 1815 [1816]. Terrestrial herb. Steep forested talus slopes, cool forested streamsides, shaded sandstone ledges in dwarf forest of upper slopes and summits of tepuis, 1900–2800 m; Bolívar (Auyán-tepui, Ilú-tepui, Macizo del Chimantá, Roraima-tepui), Amazonas (Cerro Marahuaka, Sierra de la Neblina). Venezuelan Coastal Cordillera and Andes, Sierra de Perijá; Central America, West Indies, northern South America. Peperomia alata Ruiz & Pav., Fl. Peruv. 1: 31, pl. 48. 1798. Peperomia alata var. angustifolia C. DC. in A. DC., Prodr. 10(1): 418. 1869. Peperomia nilssonii Yunck., Bol. Soc. Venez. Ci. Nat. 23(101): 64, fig. 2. 1962. Peperomia microreticulata Steyerm., Bol. Soc. Venez. Ci. Nat. 26(110): 412, fig. 1. 1966. Succulent epiphytic, terrestrial, or saxicolous herb, 7–40 cm tall. Humid forests, shaded sandstone ledges and tree trunks, generally at 200–1400 m; Bolívar (Altiplanicie de Nuria, Cerro Jaua, Cerro Sarisariñama, Cerro Venamo, El Dorado, Isla Anacoco, northeast of Luepa, Ptari-tepui, near Río Aponguao, near Río Venamo, upper Río Cuyuní Serranía de Imataca), Amazonas (Cerro Duida, Sierra de la Neblina, Sierra Parima). Venezuelan Coastal Cordillera and Andes, Sierra de Perijá; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil.

Peperomia alpina (Sw.) A. Dietr., Sp. Pl. 185. 1831. —Piper alpinum Sw., Prodr. 15. 1788. Peperomia longemucronata C. DC., Notizbl. Bot. Gart. Berlin-Dahlem 6: 497. 1917. Peperomia longirostrata Yunck., Piperac. N. South Amer. 2: 676. 1950. —Peperomia longemucronata f. longirostrata (Yunck.) Steyerm., Fl. Venez. 2(2): 133. 1984. Herb, principally terrestrial, sometimes epiphytic, with erect stems 15–35 cm tall. Forested tepui slopes and escarpments, 1900–2200 m; Bolívar (Auyán-tepui, Ilútepui, Macizo del Chimantá, Roraima-tepui). Venezuelan Andes, Coastal Cordillera; Mexico, Central America, Antilles, Colombia. ◆Fig. 603. Peperomia angustata H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 68. 1815 [1816]. —Piper angustatum (H.B.K.) Poir. in Lam., Encycl., Suppl. 4: 469. 1816. Peperomia victoriana C. DC. in A. DC., Prodr. 16(1): 449. 1869. Peperomia victoriana var. marginata C. DC., Proc. Amer. Acad. Arts 40: 685. 1905. Peperomia viridispica Trel., Contr. U.S. Natl. Herb. 26: 44. 1927. Cespitose succulent, epiphytic or saxicolous, perennial herb, with 4-angled stems 20–30 cm or more long. Crevices of rocks, along ledges, sometimes epiphytic on tree trunks and branches, deciduous and gallery forests, 50–600 m; Delta Amacuro (Los Castillos de Guayana), Bolívar (40 km west of El Manteco, near Puerto Ordaz, between Río Caura and Cuchivero, middle Río Orinoco, east slopes of Serranía de los Pijiguaos,

Peperomia 691

Platanal, Represa Guri, slopes of Cerro Bolívar), Amazonas (ca. 5 km north-northwest Samariapo, 10 km. south Puerto Ayacucho, drainage of Río Arari). Northern and central Venezuela west to Sierra de Perijá and Apure, absent from the Andes and most of Coastal Cordillera; Colombia, Guyana. ◆Fig. 604. Peperomia blanda (Jacq.) H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 67. 1815 [1816]. —Piper blandum Jacq., Collectanea 3: 211. 1789 [1791]. —Salvaje. Peperomia ciliata H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 68. 1815 [1816]. Peperomia blanda var. langsdorfii (Miq.) Henschen, Nova Acta Regiae Soc. Sci. Upsal. ser. 3, 8: 39. 1873. Peperomia decipiens C. DC., Notizbl. Bot. Gart. Berlin-Dahlem 6: 493. 1917. Peperomia salvaje C. DC., Annuaire Conserv. Jard. Bot. Genève 21: 244. 1920. Peperomia macaroana Trel. ex V. Badillo in Pittier et al., Cat. Fl. Venez. 1: 245. 1945, nom. nud. Pubescent terrestrial, epiphytic, or saxicolous herb with stems 10–60 cm tall. Semideciduous forests, 200–800 m; Bolívar (Altiplanicie de Nuria, Cerro Bolívar, near Santa Elena de Uairén). Widespread elsewhere in Venezuela; widely distributed from Mexico, West Indies, parts of Central America, and South America to Brazil, Bolivia, and Argentina. Peperomia celiae Yunck., Mem. New York Bot. Gard. 9(3): 324. 1957. Succulent herb with spreading stems 6– 15 cm long. Forested tepui slopes, ca. 1500 m; Amazonas (Cerro Yutajé). Endemic. Peperomia cladara Yunck., Mem. New York Bot. Gard. 9(3): 323. 1957. Peperomia cladara f. ciliata Yunck., Mem. New York Bot. Gard. 9(3): 324. 1957. Peperomia cladara f. perglabra Yunck., Mem. New York Bot. Gard. 9(3): 323. 1957. Stoloniferous, succulent herb with purplish or reddish brown, strongly longitudinally grooved stems 10–20 cm long, with scaly exfoliating epidermis. Rock crevices and ledges of sandstone, and along slopes of

canyons and forested streams on tepui summits, 1500–2000 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Macizo del Chimantá), Amazonas (Cerro Coro-Coro, Cerro Guaiquinima, Cerro Huachamacari, Cerro Sipapo, Cerro Yutajé). Endemic. ◆Fig. 592. Peperomia delascioi Steyerm., Brittonia 40: 294, fig. 1. 1988. Procumbent fleshy herb 5–20 cm tall; leaves with the uppermost opposite, others alternate, very small, 5–8 mm long, suborbicular; spikes slender, sparsely flowered. Sandstone exposures and shaded crevices, tepui summits, 2400–2500 m; Bolívar (Camarcaibarai-tepui, Ilú-tepui), Amazonas (Cerro Marahuaka). Guyana (Mount Roraima). Peperomia dendrophila Schlecht. & Cham., Linnaea 5: 74. 1830. Peperomia pennellii Trel. & Yunck., Piperac. N. South Amer. 2: 608, fig. 533. 1950. Terrestrial or epiphytic, coriaceous-leaved herbaceous plant with erect to ascending stems 3–25 cm tall; pseudopedicels on the rachis of fruiting specimens generally conspicuous. Wet montane forests, 1200–2000 m; Bolívar (Macizo del Chimantá), Amazonas (Sierra de la Neblina). Venezuelan Coastal Cordillera and Andes; southern Mexico, Central America, Colombia, Ecuador, possibly Peru. Peperomia distachya (L.) A. Dietr., Sp. Pl. 1: 156. 1831. —Piper distachyon L., Sp. Pl. 30. 1753. Peperomia geminispica Trel. & Yunck., Piperac. N. South Amer. 2: 700. 1950. —Peperomia distachya f. geminispica (Trel. & Yunck.) Steyerm., Fl. Venez. 2(2): 86. 1984. Peperomia foveolata Steyerm., Fl. Venez. 2(2): 99, fig. 12. 1984. Terrestrial herb, rooting at nodes; leaves alternate, 7–11-veined, narrowed to a subacute base; spikes paired. Humid forests, ca. 900 m; Amazonas (Sierra de la Neblina). Venezuelan Coastal Cordillera, locally on lower slopes of the Andes of Barinas and Táchira, Sierra de Perijá, Zulia; widely distributed from Mexico, Central America, Antilles, South America to Peru and Brazil.

692

P IPERACEAE

Peperomia duidana Trel., Bull. Torrey Bot. Club 58: 354. 1931. Creeping terrestrial, epiphytic, or saxicolous herb with elongate stems. Lower montane forests, common on sandstone mountains but also in granitic areas, 600–1200 m; Bolívar (Auyán-tepui Serranía de Maigualida), Amazonas (Cerro Autana, Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú, Cerro Sipapo, Cerro Yutajé, Sierra de la Neblina). Guyana, Ecuador. ◆Fig. 582. Peperomia elongata H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 62. 1815 [1816]. Succulent terrestrial, epiphytic, or saxicolous herb. Evergreen lowland to lower montane forests, commonly along streams, 100–1000 m; Delta Amacuro, Bolívar, Amazonas. Venezuelan Coastal Cordillera; Colombia, Guyana, Suriname, French Guiana, Brazil; 3 varieties, all in Venezuela, 2 of these in the flora area. The typical variety has not been recorded from the flora area. It has ovate or ovate-elliptic shorter leaves rounded or obtuse at base, and occurs in the Coastal Cordillera of Venezuela. Key to the Varieties of P. elongata 1. Stems and leaf surfaces glabrous; petioles glabrous except for ciliate margins .................................... var. guianensis 1. Stems, leaf surfaces, and petioles pubescent .............................. var. piliramea P. elongata var. guianensis Yunck. in Maguire, Bull. Torrey Bot. Club 75: 290. 1948. —Ashudu. Herbaceous, pendent epiphyte, also saxicolous or spreading over ground, with elongate stems to 1 m long. 50–1000 m; Delta Amacuro (Caño Araguao), Bolívar (middle Río Caura, Río Paragua), Amazonas (lower Río Ventuari, La Esmeralda, Maroa, near Santa Bárbara del Orinoco, Río Arari, Río Cunucunuma, Río Temi, upper Río Padamo, upper Río Yatúa). Sierra de Perijá, Barinas, Portuguesa; Guyana. P. elongata var. piliramea Trel. & Yunck., Piperac. N. South Amer. 2: 659. 1950. Epiphyte. Ca. 100 m; Amazonas (near San Carlos de Río Negro). Falcón; Guyana.

Peperomia emarginella (Sw. ex Wikstrom) C. DC. in A. DC., Prodr. 16(1): 437. 1869. —Piper emarginellum Sw. ex Wikstrom, Kongl. Vetensk. Acad. Handl. 56. 1828. Peperomia emarginella var. glabrior C. DC., Anales Inst. Fís.-Geogr. Nac. Costa Rica 9: 177. 1897. Delicate, minute creeping herb with filiform stems, epiphytic or saxicolous on moist, mossy rocks. Humid evergreen lowland to lower montane forests, 50–700 m; Delta Amacuro (Serranía de Imataca), Bolívar (Altiplanicie de Nuria, El Dorado to Santa Elena, La Escalera, Río Paragua), Amazonas (Cerro Huachamacari). Venezuelan Coastal Cordillera and lower slopes of the Andes of Táchira and Sierra de Perijá; Central America, West Indies, northern South America. ◆Fig. 594. Peperomia enantiostachya C. DC., Bull. Herb. Boissier 6: 514. 1898. Succulent epiphyte with pendent to ascending stems. Wet, lower montane forests along rocky streams, 1800–1900 m; Amazonas (Sierra de la Neblina). Andes of Colombia and Ecuador. Peperomia fundacionensis Steyerm., Fl. Venez. 2(2): 101, fig. 12. 1984. Terrestrial or subepiphyte on tree trunk. Wet bases of tepui bluffs, 1700–1800 m; Amazonas (Serranía Yutajé). Táchira (sandstone areas). Peperomia galioides H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 71, pl. 17. 1815 [1816]. Branching, cespitose succulent herb, terrestrial, saxicolous, or epiphytic with stems 10–80 cm tall. Shady sandstone ledges, montane and dwarf forests on upper tepui slopes and summits, 2100–2800 m; Bolívar (Ilútepui, Roraima-tepui, Sororopán-tepui), Amazonas (Cerro Marahuaka). Venezuelan Coastal Cordillera and Andes, Sierra de Perijá; Mexico, Central America, West Indies, South America. Peperomia gentryi Steyerm., Ann. Missouri Bot. Gard. 74: 87, fig. 2A and 2B. 1987. Herbaceous plant with creeping and widely spreading stems. Wet montane forests, 1200–1300 m; Bolívar (Sierra Maigualida), Amazonas (Sierra de la Neblina). Ecuador.

Peperomia 693

Peperomia glabella (Sw.) A. Dietr., Sp. Pl. 1: 156. 1831. —Piper glabellum Sw., Prodr. 16. 1788. Herbaceous plant, usually epiphytic, rarely saxicolous, with stoloniferous, erect, pendent, or prostrate, usually branched stems. Venezuela; Central America, West Indies, South America; 2 varieties, both in the flora area. When dried, Piper glabella is easily recognized by black dots which appear on all vegetative parts as well as on peduncles and rachis of inflorescence. Key to the Varieties of P. glabella 1. Leaves ovate or ovate-elliptic to ellipticlanceolate, 1.5–3.5 × 0.5–1.3 cm, acute to acuminate at both extremities ............... ........................................ var. glabella 1. Leaves lanceolate, 3–8(–9) × 1–3 cm, acute to long attenuate at both extremities ............................... var. nervulosa P. glabella var. glabella Humid evergreen forests, 50–500 m; Delta Amacuro (between Margarita and Puerto Miranda, between Tucupita and La Horqueta, Caño Capure, Caño Pablo, Río Araguao, Río Cuyubiní), Bolívar (90 km from El Dorado to Santa Elena, Altiplanicie de Nuria, middle Río Caura). Venezuelan Coastal Cordillera and Andes, Sierra de Perijá; Central America, West Indies, South America. P. glabella var. nervulosa (C. DC.) Yunck., Ann. Missouri Bot. Gard. 37: 98. 1950. —Peperomia melanostigma var. nervulosa C. DC. in A. DC., Prodr. 16(1): 409. 1869. Leaves lanceolate, acute to long-acuminate. Montane forests, near sea level to 1100 m; Delta Amacuro (west Isla Cocuina), Bolívar (Altiplanicie de Nuria, km 33-35 highway El Dorado-Santa Elena de Uarién, Río Botanamo, Río Caura, Salto Pará, Santa Elena de Uarién), Amazonas (Río Casiquiare). Venezuelan Coastal Cordillera, Sierra de Perijá, absent from the Andes; Central America, West Indies, South America. Peperomia guaiquinimana Trel. & Yunck., Piperac. N. South Amer. 2: 596, fig. 518. 1950. Peperomia guaiqui-

nimana Trel. ex V. Badillo in Pittier et al., Cat. Fl. Venez. 1: 245. 1945, nom. nud. Epiphytic glabrous herb with ascending pale brown stems. Riparian forests, ca. 300 m; Bolívar (base of Cerro Guaiquinima, Río Paragua). Endemic. ◆Fig. 583. Peperomia haematolepis Trel., Field Mus. Nat. Hist., Bot. Ser. 13(2): 52. 1936. Saxicolous plant; leaves succulent; spikes erect; floral bracts pale yellow or ochraceousbuff. Wet sandstone bluffs in montane forests, 1500–1700 m; Amazonas (Cerro Yutajé). Venezuelan Andes, Lara, Portuguesa, Yaracuy; Colombia, Ecuador, Peru, Brazil. In the rest of its range outside the flora area, this species has the floral bracts rufous with prominent reddish maroon gland dots. Peperomia hernandiifolia (Vahl) A. Dietr., Sp. Pl. 1: 157. 1831. —Piper hernandiifolium Vahl, Enum. Pl. 1: 344. 1804. Peperomia choroniana var. puberulenta Yunck. in Trel. & Yunck., Piperac. N. South Amer. 2: 730. 1950. Terrestrial or saxicolous herb with elongate, prostrate, wine-red stems. Wet forests on tepui slopes and summits, 300–1900 m; Bolívar (Guyana border west to Cerro Jaua), Amazonas (Sierra Parima and Cerro Duida west to Sierra de la Neblina). Sucre; Mexico, Central America, West Indies, South America south to Brazil. ◆Fig. 589. Peperomia jamesoniana C. DC.,J. Bot. 4: 137. 1866, non (Miq.) DC. 1869. Peperomia chimantana Yunck., Mem. New York Bot. Gard. 9(3): 423. 1957. Peperomia pseudojamesoniana Steyerm., Fl. Venez. 2(2): 199, fig. 27. 1984. Small epiphytic or terrestrial herb with erect-ascending stems 4–10 cm long. Wet mossy forests over sandstone, (100–)1600– 2000 m; Bolívar (Macizo del Chimantá, Ptari-tepui), Amazonas (Sierra de la Neblina). Venezuelan Andes. ◆Fig. 606. Peperomia lanceolato-peltata C. DC., J. Bot. 4: 136. 1866. Peperomia boomii Steyerm., Brittonia 38: 220, fig. 1. 1986. Herb 6–10 cm tall. Along dry stream beds on igneous rocks, ca. 200 m; Bolívar (Ser-

694

P IPERACEAE

ranía Cerbatana). Northern Venezuela; Central America, Colombia. Peperomia lancifolia Hook., Icon Pl. 4, pl. 332. 1841. Peperomia erasmiaeformis Trel., Publ. Field Mus. Nat. Hist., Bot. Ser. 22(1): 9. 1940. —Peperomia lancifolia subsp. erasmiaeformis (Trel.) Steyerm., Fl. Venez. 2(2): 131. 1984. Terrestrial, epiphytic, or saxicolous herb with green or purplish, erect or ascending stems 10–30 cm tall; spikes 1 or 2 per peduncle. Wet montane forested slopes overlying sandstone, ca. 500–1800 m; Bolívar (Auyán-tepui, 118 km south of El Dorado, Río Cuyuní drainage), Amazonas (Sierra de la Neblina). Venezuelan Coastal Cordillera; Mexico, Central America, Colombia, Guyana, northern Ecuador, Peru, Brazil. ◆Fig. 580. Peperomia loxensis H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 70. 1815 [1816]. —Piper loxense (H.B.K.) Poir. in Lam., Encycl., Suppl. 4: 470. 1816. Peperomia diffusa C. DC., J. Bot. 4: 133. 1866. Epiphytic or saxicolous herb with erect stems 8–15 cm tall, the epidermis flaky and loosening. Montane and riparian forests at base of tepuis, ca. 1100 m; Amazonas (Cerro Huachamacari). Venezuelan Coastal Cordillera and Andes; the Andes of Colombia and Ecuador, Brazil. Peperomia macrostachya (Vahl) A. Dietr., Sp. Pl. 1: 149. 1831. —Piper macrostachyon Vahl, Enum. Pl. 1: 341. 1804. Piper myosuroides Rudge, Pl. Guian. 1: 9, pl. 5. 1805. —Peperomia myosuroides (Rudge) A. Dietr., Sp. Pl. 1: 157. 1831, “myusuroides.” Epiphytic, usually pendulous, succulent herb, with elongate stems to 1 or more m tall. Rainforests and riparian forests, 100–1200 m. Venezuelan Andes, Miranda; Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Brazil; 2 varieties, both in the flora area. Peperomia macrostacyha is commonly found in stinging ant nests, associated with species of Codonanthe and Aechmaea.

Key to the Varieties of P. macrostachya 1. Stems glabrous; leaves glabrous on both sides ...................... var. macrostachya 1. Stems, at least when young, puberulent; lower surface of leaves, at least along midrib, puberulent .................................. ............................. var. nematostachya P. macrostachya var. macrostachya 100–500 m; Delta Amacuro (Caño Capure, Jotajana, Serranía de Imataca), Bolívar (south of Cerro Venado, northwest of El Manteco, Kilómetro 88, Macizo del Chimantá, middle Río Caura, Río Paramichi, 40 km west of Santa Elena de Uairén, Serranía de Imataca), Amazonas (44–45 kilometers southeast of Puerto Ayacucho, between Paso del Diablo and Caño de Culebra, between Yavita and Pimichín, Cerro Duida, Cerro Sipapo, near San Carlos de Río Negro, Río Cuao). Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 602. P. macrostachya var. nematostachya (Link) Trel. & Yunck., Piperac. N. South Amer. 2: 661. 1950. —Peperomia nematostachya Link in Spreng. et al., Jahrb. Gewächsk. 1(3): 63. 1820. 400–1200 m; Bolívar (Pijiguaos, base of Ptari-tepui and Sororán-tepui, Río Suapure). Miranda; Colombia, Guyana, Ecuador. Peperomia maculosa (L.) Hook., Exot. Fl. pl. 92. 1824. —Piper maculosum L., Sp. Pl. 30. 1753. —Yerba salvaje. Terrestrial or saxicolous, succulent herb with stems 3–40 cm tall. Wet montane forests, 1000–1200 m; Bolívar (Cerro Wa-kaosen, southwest of Santa Elena de Uairén), Amazonas (Sierra de la Neblina). Venezuelan Coastal Cordillera and Andes, Sierra de Perijá; Central America, West Indies, northern South America. ◆Fig. 588. A striped-leaf variant sometimes occurs. Peperomia magnoliifolia (Jacq.) A. Dietr., Sp. Pl. 1: 153. 1831. —Piper magnoliifolium Jacq., Collectanea 3: 210. 1789 [1791], “magnoliaefolium.” Thick-leaved herb, saxicolous, epiphytic,

Peperomia 695

or rarely terrestrial, with dull lavender or magenta, reddish-marked stems 10–30 cm long. Evergreen lowland to lower montane forests, 50–500 m; Delta Amacuro (near El Palmar, near Jotajana, Río Acure, west of Isla Cocuina), Bolívar (Cerro Pijiguaos), Amazonas (Cerro Yapacana, near Siquita, San Simón de Cocuy). Venezuelan Coastal Cordillera, Sierra de Perijá. Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil. Peperomia marahuacensis Steyerm., Ann. Missouri Bot. Gard. 74: 87, fig. 2C. 1987. Terrestrial or saxicolous herb with stems elongated to 80 cm; leaves palmately 5veined. Cave-like depressions on tepui summit, ca. 2600 m; Amazonas (Cerro Marahuaka). Endemic. Peperomia maypurensis H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 66. 1815 [1816]. —Piper maypurense (H.B.K.) Poir. in Lam., Encycl., Suppl. 4: 468. 1816. Peperomia ornata Yunck., Brittonia 8: 61, fig. 1. 1954. Succulent, cespitose, saxicolous herbaceous plant with magenta or reddish purple stems 2–17 cm tall; leaves striped. On sandstone or granitic boulders or ledges in lowland to lower montane forests, 100–300 (–1000) m; Bolívar (Río Parguaza), Amazonas (Cerro Marahuaka, Cerro Sipapo, Puerto Ayacucho to Gavilán, San Pedro de Cataniapo). Endemic. ◆Fig. 591.

var. emarginata (Ruiz & Pav.) Dahlst., Kongl. Svenska Vetenskapsakad. Handl. 33(2): 66. 1900. Peperomia cuneata Miq. in Hook., London J. Bot. 4: 429. 1845. —Peperomia obtusifolia var. cuneata (Miq.) Griseb., Fl. Brit. W. I. 166. 1864. Peperomia magnoliaefolia var. emarginulata C. DC. in A. DC., Prodr. 16(1): 427. 1869. —Peperomia obtusifolia var. emarginulata (C. DC.) Trel. & Yunck., Piperac. N. South Amer. 2: 680, fig. 596. 1950. Epiphytic or terrestrial herb with thick, fleshy, erect or ascending green or reddish purple-marked stems 3–15 cm or more tall. Montane and humid forests, 50–1100 m; Delta Amacuro (near Caño Araguao), Bolívar (Altiplanicie de Nuria, alto Río Cuyuní, Perai-tepui, Río Chicanán basin, Serranía de Imataca), Amazonas (23 km northeast from Puerto Ayacucho). Venezuelan Coastal Cordillera and Andes, Sierra de Perijá; Mexico, Central America, West Indies, South America. ◆Fig. 595.

Peperomia neblinana Yunck., Mem. New York Bot. Gard. 10(5): 43. 1964. Much-branched, epiphytic herbaceous plant with spreading, elongate stems 7–12 cm long. Wet montane slopes, 1200–1700 m; Bolívar (Sierra Maigualida), Amazonas (Sierra de la Neblina). Endemic.

Peperomia ouabianae C. DC., Candollea 1: 400. 1923. Peperomia silvestris C. DC., Notizbl. Bot. Gart. Berlin-Dahlem 6: 494. 1917, non C. DC., 1866. Peperomia tafelbergensis Yunck. in Maguire, Bull. Torrey Bot. Club 75: 292. 1948. Peperomia wurdackii Yunck., Mem. New York Bot. Gard. 9(3): 434. 1957. Creeping, diminutive epiphytic or saxicolous herbaceous plant, decumbent, ascending, to suberect, with elongate stems 3–8 cm long. Humid lower montane to montane forests, 300–1200 m; Bolívar (El Paují, Urimán), Amazonas (Cerro Neblina, west of Serranía de Yutajé). Venezuelan Coastal Cordillera; Guyana, Suriname, French Guiana. ◆Fig. 585.

Peperomia obtusifolia (L.) A. Dietr., Sp. Pl. 1: 154. 1831. —Piper obtusifolium L., Sp. Pl. 30. 1753. Peperomia emarginata Ruiz & Pav., Fl. Peruv. 1: 33, pl. 49, fig. a. 1798. —Piper emarginatum (Ruiz & Pav.) Vahl, Enum. Pl. 1: 339. 1804. —Peperomia obtusifolia

Peperomia pellucida (L.) H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 64. 1815 [1816]. —Piper pellucidum L., Sp. Pl. 30. 1753. Delicate terrestrial herbaceous annual, stems 10–40 cm tall. Disturbed or cultivated areas of gardens and plant nurseries, also on shaded alluvial ground, mostly near sea

696

P IPERACEAE

level to 300 m; Delta Amacuro (Capure, Sacupana, Tucupita), Bolívar (north of Isla Anacoco), and Amazonas (20 km south of confluence of Río Negro and Brazo Casiquiare, middle and upper Orinoco, Puerto Ayacucho). Common throughout Venezuela; southern U.S.A., Central America, West Indies, South America, tropical portions of the Old World. ◆Fig. 593. Peperomia pernambucensis Miq. in Hook., London J. Bot. 4: 420. 1845. Epiphytic succulent herbaceous plant with erect to decumbent stem 3–5 cm tall. Wet lower montane forests, 600–800 m; Bolívar (near headwaters of Río Paramichi, near Serranía Pia-zoi). Central America, Trinidad, Guyana, Suriname, French Guiana, Colombia, northeastern Brazil. ◆Fig. 599. Peperomia purpurinervis C. DC., Notizbl. Bot. Gart. Berlin-Dahlem 6: 496. 1917. Terrestrial saxicolous or epiphytic herb with dull magenta or lavender, erect to spreading stems 10–25 cm tall. Wet mossy slopes, moist sandstone ledges, and rocks along streams, near waterfalls, and montane forests, 1200–2000 m; Bolívar (Auyán-tepui, Cerro Venamo, headwaters of Río Cuyuní, Ilú-tepui, Macizo del Chimantá, Roraimatepui west to Auyán-tepui). French Guiana. ◆Fig. 605. Peperomia quadrangularis (J.V. Thomps.) A. Dietr., Sp. Pl. 1: 169. 1831. —Piper quadrangulare J.V. Thomps., Trans. Linn. Soc. London 9: 202, pl. 21, fig. 1. 1808. Peperomia saginans Trel. ex V. Badillo in Pittier et al., Cat. Fl. Venez. 1: 246. 1945, nom. nud. Peperomia atabapoensis Steyerm., Fl. Venez. 2(2): 52, figs. 1, A3, B3. 1984. Epiphytic or saxicolous herbaceous plant with creeping, pendent, elongate stems. Wet forests, frequently along streams, 100–600 m; Bolívar (Altiplanicie de Nuria, near Río Parguaza, Río Ventuari, Salto Pará, middle Río Caura, Sierra de Guayapo, south of Cerro Venado, south to El Dorado), and Amazonas (25-30 km southwest Puerto Ayacucho, confluence between Río Atabapo and Guaviare, San Fernando de Atabapo, near Río Manapiare, near Río Ventuari,

Síquita). Generally common in Venezuela; Panama, West Indies, northern South America. ◆Fig. 587. Peperomia quaesita Trel. in J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(2): 83. 1936. Epiphytic or rarely terrestrial herbaceous plant with erect or spreading stems 8–15 cm tall and usually 4-verticillate (rarely 3- or 5-) leaves. Rainforests, 100–1000 m; Bolívar (near Santa Teresita de Kavanayén) and Amazonas (middle Río Matacuni). Western Venezuela (Apure, Táchira); Colombia, Ecuador, Peru, Bolivia. ◆Fig. 584. Peperomia rotundata H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 67, pl. 12. 1815 [1816]. Terrestrial, epiphytic, or saxicolous herbaceous plant with wine-red, brick-red, or reddish brown stems; upper surface of leaves dark green and shining, the lower surface pale green, maroon-purple, or reddish purple; peduncles and inflorescences often reddish purple. 2000–2700 m. Colombia, Venezuela, Ecuador; 3 varieties, all in Venezuela, 2 of these in the flora area. Key to the Varieties of P. rotundata 1. Most of the leaves of the fertile branches opposite ....................... var. rotundata 1. Most of the leaves of the fertile branches alternate ..................... var. trinervula P. rotundata var. rotundata Peperomia epilobioides Trel. & Yunck., Piperac. N. South Amer. 2: 551, fig. 483. 1950. Peperomia trinervula var. suboppositifolia Trel. & Yunck., Piperac. N. South Amer. 2: 648. 1950. Terrestrial plant with reddish brown stem, ascending to 40 cm. Open rocky plateau, 2500–2600 m; Amazonas (summit of Cerro Marahuaka). Venezuelan Andes; the Andes of Colombia, Ecuador. P. rotundata var. trinervula (C. DC.) Steyerm., Fl. Venez. 2(2): 217. 1984. —Peperomia trinervula C. DC. in A. DC., Prodr. 16(1): 420. 1869. (?)Peperomia moritzii C. DC. in A. DC., Prodr. 16(1): 420. 1869.

Peperomia 697

Peperomia purpurascens Moritz ex C. DC. in A. DC., Prodr. 16(1): 420. 1869. Peperomia roraimana C. DC., Notizbl. Bot. Gart. Berlin-Dahlem 6: 493. 1917. Terrestrial or saxicolous herbaceous plant with wine-red or brick-red, erect or ascending stems 15–60 cm tall. Shaded sandstone ledges, wet upper montane forests, 2000– 2700 m; Bolívar (Aparamán-tepui, Roraimatepui), Amazonas (Cerro Marahuaka). Venezuelan Coastal Cordillera and Andes; Colombia. Peperomia rotundifolia (L.) H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 65. 1815 [1816]. —Piper rotundifolium L., Sp. Pl. 30. 1753. Piper nummularifolium Sw., Prodr. 16. 1788, non Lam. & Willd. ex C. DC. 1869. Peperomia nummularifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 66. 1815 [1816]. Peperomia rotundifolia f. ovata Dahlst., Kongl. Svenska Vetenskapsakad. Handl. 33(2): 101. 1900. —Peperomia rotundifolia var. ovata (Dahlst.) C. DC. in Urb., Symb. Antill. 3: 230. 1902. Peperomia bartlettii C. DC., Notizbl. Bot. Gart. Berlin-Dahlem 6: 470. 1917. Delicate, creeping or climbing, epiphytic or saxicolous herb and filiform stems; leaves pale green. Wet forests, 50–1000 m; Delta Amacuro (Altiplanicie de Nuria, near Caño Mariusa, Río Acure, Río Cuyuní, Río Orocoima, Salto Pará, southwest of Tumeremo), Bolívar (El Palmar, Gran Sabana, middle Rio Caura, upper Río Cuyuní), Amazonas (middle Caño Yagua, east of Cerro Yapacana, near Maroa, near Río Cataniapo, Río Cunucunuma, upper Río Padamo, upper Río Yatua, near Samariapo, near San Carlos de Río Negro, Sierra Parima, Solano, upper Yavita). Venezuelan Coastal Cordillera and Andes; widespread in Mexico, Central America, West Indies, South America. ◆Fig. 586.

Amacuro), Bolívar (Altiplanicie de Nuria, Río Caura, Serranía de Imataca, upper Río Cuyuní), Amazonas. Coastal Cordillera, Sierra de Perijá, Apure, Táchira, but absent from the Andes; Central America, West Indies, South America to Peru and Brazil. ◆Fig. 598. Peperomia spruceana Benth. ex. C. DC. in A. DC., Prodr. 16(1): 402. 1869. Terrestrial, succulent herbaceous plants, often occurring in colonies, with spreadingascending stems; leaves rugose. Evergreen lowland forests, 100–200 m; Amazonas (Río Mawarinuma, Sierra de la Neblina). Amazonian Brazil. Peperomia striata Ruiz & Pav., Fl. Peruv. 1: 32, t. 52. 1798. Peperomia omnicola C. DC., Bull. Herb. Boissier 6: 507. 1898. Peperomia omnicola var. oblanceolata Trel., J. Wash. Acad. Sci. 16: 206. 1926. Large erect herb, principally terrestrial, with stems 0.5–1 m tall. Wet forests of tepuis, 1200–2300 m; Bolívar (Ptari-tepui), Amazonas (Sierra de la Neblina). Venezuelan Andes; Guatemala, Costa Rica, Colombia, Ecuador, Peru, Bolivia. ◆Fig. 590. Peperomia tenella (Sw.) A. Dietr., Sp. Pl. 1: 153. 1831. —Piper tenellum Sw., Prodr. 16. 1788. —Acrocarpidium tenellum (Sw.) Miq., Syst. Piperac. 53. 1843. Small, terrestrial, epiphytic, or saxicolous herb with erect, slender stems. Shady sandstone ledges, moist mossy rocks, wet forested tepui slopes, 1400–2600 m. Central America, northern South America; 2 varieties, both in the flora area. Some specimens are intermediate between the two varieties, and between Peperomia tenella and P. tovariana C. DC. Key to the Varieties of P. tenella

Peperomia serpens (Sw.) Loudon, Hort. Brit. 13. 1830, non C. DC. —Piper serpens Sw., Prodr. 16. 1788. Peperomia ionophylla Griseb., Mem. Amer. Acad. new ser. 8: 175. 1861. Creeping or climbing epiphytic or saxicolous herb with elongate stems. Evergreen lowland to lower montane forests, 50–300 (–700) m; Delta Amacuro (Araguao, Río

1. Leaves 6–18(–21) × 2–6.5(–12) mm; petiole 1–2 mm long; stem 0.5–1(–1.5) mm thick; spikes 0.5–1 mm thick .................. .......................................... var. tenella 1. Leaves 12–25(–38) × 6–15 mm; petiole 2– 5 mm long; stem 1.5–2.5(–4) mm thick; spikes 1.5–2(–3) mm thick ...................... ............................................. var. tyleri

698

P IPERACEAE

P. tenella var. tenella 1500–2600 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Ilú-tepui, Cerro Jaua, Macizo del Chimantá, Ptari-tepui, Roraimatepui west to Cerro Jaua, Sierra de la Neblina), Amazonas (Cerro Duida, Cerro Marahuaka, Cerro Parú, Cerro Sipapo). Venezuelan Coastal Cordillera and Andes; Central America, northern South America. P. tenella var. tyleri (Trel.) Steyerm., Fl. Venez. 2(2): 246. 1984. —Peperomia tyleri Trel., Bull. Torrey Bot. Club 58: 355. 1931. 1400–2000 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Macizo del Chimantá [Chimantá-tepui], [Toronó-tepui], Serranía Marutaní, Sierra Pakaraima), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Sipapo). Colombia. ◆Fig. 596. Peperomia tetraphylla (G. Forst.) Hook. & Arn., Bot. Beechey Voy. 97. 1841 [1832]. —Piper tetraphyllum G. Forst., Fl. Ins. Austr. 5. 1786. Piper reflexum L. f., Suppl. Pl. 91. 1781. —Peperomia reflexa (L. f.) A. Dietr., Sp. Pl. 1: 180. 1831, non H.B.K. 1815 [1816]. Peperomia reflexa f. americana Miq., Syst. Piperac. 173. 1843. —Peperomia reflexa var. americana (Miq.) Dahlst., Kongl. Svenska Vetenskapsakad. Handl. 33(2): 175, pl. 8, fig. 5, 1900. —Peperomia tetraphylla var. americana (Miq.) Yunck., Bol. Inst. Bot. (São Paulo) 3: 178. 1966. Peperomia cryptotricha Trel. ex V. Badillo in Pittier et al., Cat. Fl. Venez. 1: 244. 1945, nom. nud. Dwarf herb, epiphytic, rarely saxicolous, stems 5–20(–25) cm or more tall; leaves usually 4-verticillate. Rare on wet forested tepui slopes and summits, 1800–2500 m; Bolívar (Macizo del Chimantá [Sarvén-tepui], Roraima-tepui), Amazonas (Sierra de la Neblina). Cloud forests of Venezuelan Coastal Cordillera and Andes, Sierra de Perijá; pantropics. Peperomia uaupesensis Yunck., Bol. Inst. Bot. São Paulo 3: 194, t. 171. 1966. Succulent, epiphytic herb with ascending stems; leaves coriaceous; spikes deep orange when dry. Gallery forests, often growing on rotting logs, 50–400 m; Bolívar (Serranía

Cerbatana), Amazonas (Puerto Ayacucho to Samariapo, Río Autana). Colombia (Meta), Peru (Madre de Dios), Brazil (Acre, Amazonas). Peperomia urocarpa Fisch. & C.A. Mey., Index Sem. Hort. Petrop. no. 1577. 1837. —Acrocarpidium urocarpum (Fischer & C. Mey.) Miq., Syst. Piperac. 60. 1843. Acrocarpidium majus Miq., Syst. Piperac. 60. 1843. Peperomia hederacea Miq. in Mart., Fl. Bras. 4(1): 20. 1853. Peperomia major C. DC. in A. DC., Prodr. 16(1): 432. 1869. Epiphytic, saxicolous, or rarely terrestrial subscandent succulent herb, with elongate, rooting stems. Lower montane forests, 500– 1000 m; Bolívar (Cerro Wa-kao-ien, 80 km southwest of El Dorado, Río Chicanán, upper Río Cuyuní, southwest of Santa Elena de Uairén). Sierra de Perijá; West Indies, Colombia, Ecuador, Brazil. ◆Fig. 597. Peperomia venezueliana C. DC., J. Bot. 4: 139. 1866. Peperomia ernstiana C. DC. in A. DC., Prodr. 16(1): 422. 1869. Peperomia horrescens Trel. ex V. Badillo in Pittier et al., Cat. Fl. Venez. 1: 245. 1945, nom. nud. Principally terrestrial, rarely epiphytic or saxicolous herb with black-dotted vegetative parts, as in P. glabella, the stems erect or ascending, 10–35 cm tall; upper surface of leaves often variegated, the lower surface often lavender-spotted. Wet forested tepui slopes, 1200–1300 m; Bolívar (upper Río Caura, near El Palmar), Amazonas (Sierra de la Neblina). Venezuelan Coastal Cordillera and Andes; Colombia. Peperomia venusta Yunck., Mem. New York Bot. Gard. 9: 325. 1957. Terrestrial glabrous herb with prostrate or decumbent stems 3.5–10 cm long. Wet, forested tepui slopes, ca. 1000 m; Amazonas (Cerro Marahuaka). Endemic. ◆Fig. 601. Peperomia yatuensis Steyerm., Fl. Venez. 2(2): 275, fig. 36. 1984. Creeping, epiphytic or saxicolous herbaceous plant. Wet lower montane forests, ca. 900 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 581.

Peperomia 699

Peperomia yutajensis Steyerm., Fl. Venez. 2(2): 275, fig. 37. 1984. Terrestrial, subscandent herb with elon-

gate stems 60–100 cm long. Wet, forested tepui slopes, ca. 1800 m; Amazonas (Cerro Yutajé). Endemic. ◆Fig. 600.

Fig. 580. Peperomia lancifolia

Fig. 581. Peperomia yatuensis

Fig. 582. Peperomia duidana

Fig. 583. Peperomia guaiquinimana

Fig. 584. Peperomia quaesita

700

P IPERACEAE

Fig. 585. Peperomia ouabianae

Fig. 586. Peperomia rotundifolia

Fig. 588. Peperomia maculosa

Fig. 589. Peperomia hernandiifolia

Fig. 587. Peperomia quadrangularis

Peperomia 701

Fig. 590. Peperomia striata

Fig. 592. Peperomia cladara

Fig. 591. Peperomia maypurensis

Fig. 593. Peperomia pellucida

Fig. 594. Peperomia emarginella

702

P IPERACEAE

Fig. 595. Peperomia obtusifolia

Fig. 596. Peperomia tenella var. tyleri

Fig. 598. Peperomia serpens

Fig. 597. Peperomia urocarpa

Peperomia 703

Fig. 599. Peperomia pernambucensis

Fig. 601. Peperomia venusta

Fig. 600. Peperomia yutajensis

704

P IPERACEAE

Fig. 602. Peperomia macrostachya var. macrostachya

Fig. 603. Peperomia alpina

Fig. 604. Peperomia angustata

Fig. 605. Peperomia purpurinervis

Fig. 606. Peperomia jamesoniana

Piper 705

2. PIPER L., Sp. Pl. 28. 1753, Gen. Pl. 333. 1737. Ottonia Spreng., Neue Entd. 1: 255. 1820. Peperidia Kostel., Allg. Med.-pharm. Fl. 2: 455. 1833, non Reichenb. 1828. Amalago Raf., Sylva Tellur. 84. 1838. Lepianthes Raf., Sylva Tellur. 85. 1838, non Lepanthes Sw. 1799. Enckea Kunth, Linnaea 13: 590. 1839. Heckeria Kunth, Linnaea 13: 564. 1839, non Raf. 1838. Pothomorphe Miq., Bull. Sci. Phys. Nat. Néerl. 2: 447, 450. 1839. Schilleria Kunth, Linnaea 13: 676. 1839. Steffensia Kunth, Linnaea 13: 609. 1839. Artanthe Miq., Comm. Phytogr. 40, t. 7, 8. 1840. Generally trees or shrubs, ± ligneous with soft and subherbaceous wood, sometimes climbing or subscandent with elongate stems, rarely herbs (P. peltatum, P. umbellatum) or truly ligneous epiphytes (P. subsessilifolium var. morii). Stems usually with enlarged nodes, with a lateral prophyll frequently in the form of a flap similar to a bud scale over apex of shoot. Leaves alternate, simple, petiolate; petiole with margins narrowly to conspicuously winged, ± sheathed only at base (at flowering nodes) or throughout the petiolar length to base of leaf blade (at vegetative nodes); blades entire, or lobed only at base (in Venezuelan species), sometimes peltate, membranous to thick-coriaceous, the upper surface smooth, rugose, or verrucose to alveolate, glabrous to densely pubescent, sometimes scabrous, the lower surface often with pale or dark glandular dots or diminutive translucent dots; venation palmate to pinnate, the secondary lateral veins arising, when pinnately arranged, from below the middle only, or to the upper third only, or extending from base to apex throughout the length. Inflorescence axillary or terminal, appearing opposite the leaf, pedunculate, with a single spike; spike linear-cylindric (in Venezuelan species), erect, ascending, or pendent, straight, arched, or recurved, apically rounded, obtuse, mucronate, or caudate; rachis generally ribbed between flowers, smooth, papillate, fimbrillate, or villous. Flowers numerous, usually densely massed or less frequently more distant from one another, principally sessile, each subtended by a bract of diverse form; bract usually subpeltate or peltate, broadened above, rounded or triangular at summit (when viewed from above), narrowed below to a pedicellate part, glabrous or pubescent, or concave and cucullate upward with an inflexed apex, ligulate, spathulate, or concave and shell-like, the apex itself glabrous or pubescent in the center, but usually with a fimbrillate margin with short to elongate trichomes. Stamens (1)2, 4, or 5; anthers with lateral or vertical dehiscence, the connective sometimes prominent and disk-like; filaments usually short. Ovary unilocular, ovoid, subcylindric, conic, or obovoid; stigmas (2)3(4 or 5), filiform, short or rounded, sessile or with a short to somewhat elongate style. Fruit 1-seeded, fleshy, dry, globose, obpyramidal, cylindric, or turbinate, rounded or angled, sometimes laterally compressed, trigonous, or quadrangular, sessile or long-pedicillate, smooth and glabrous or papillate-puberulent or hispidulous. Pantropics; ca. > 1000 species, 139 in Venezuela, 73 of these in the flora area. Key to the Species of Piper 1. 1.

Spikes several to many, umbellate on a common peduncle; leaves orbicular, orbicular-ovate, or broadly reniform ............................................... 2 Spikes solitary on a peduncle; leaves with other shapes .......................... 3

706

P IPERACEAE

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

7.

8(3). 8. 9(8). 9. 10(9). 10. 11(10). 11. 12(10).

12.

13(12).

13. 14(9). 14.

Leaves peltate; leaves, peduncles, and petioles glabrous ........... P. peltatum Leaves conspicuously cordate with subequal lobes; leaves, peduncles, and petioles puberulent .............................................................. P. umbellatum Leaves palmately veined, with some or all of the veins arising from the very base ................................................................................................ 4 Leaves pinnately veined, with some or all of the veins arising above the very base ................................................................................................ 8 One pair of veins arising from below the middle of leaf, the others from the very base .......................................................................... P. foveolatum All veins arising from the very base of leaf blade ..................................... 5 Leaves 9–13-veined .................................................................. P. marginatum Leaves 5–7(–9)-veined ................................................................................ 6 Leaves mainly drying dark or blackish (sometimes green); tertiary veinlets obscure, neither prominent, reticulate, nor elevated ........ P. amalago Leaves drying yellow- or bronze-green; tertiary veinlets prominent, reticulate, elevated .................................................................................... 7 Petiole glabrous; leaves mainly 15–35 cm long, 7–28 cm wide, to 2.2 times longer than wide; ovary truncate, densely papillate-puberulent, with an apical glabrous disk ......................................................... P. reticulatum Petiole ciliate-pilose with spreading trichomes; leaves mainly (8–)10– 21 cm long, (2–)3–11 cm wide, (1.5–)2–3.5 times longer than wide; ovary acutely pointed, usually glabrous or sparsely papillate above .......................................................................................................... P. tenue Longest main lateral veins all arising from midrib within the lower 3/8 of leaf blade ................................................................................................ 9 Longest main lateral veins arising throughout the length of leaf blade, or at least extending to the middle, or upper 1/3, or 1/4 of the length ...... 16 Plants with climbing stems...................................................................... 10 Plants without climbing stems ................................................................ 14 Lower surface of leaf completely glabrous .............................................. 11 Lower surface of leaf with at least some veins pubescent ...................... 12 Tertiary venation of lower surface of leaf obsolete or inconspicuous .................................................................................................... P. vitaceum Tertiary venation of lower surface of leaf conspicuously reticulate and elevated ........................................................................................... P. politii Leaves subcordate or slightly so at base, oblong-ovate, to suborbicularovate, slightly longer than wide to 2.6 times longer than wide; rachis glabrous; tertiary venation conspicuously grossly reticulate on lower surface .................................................................................................. 13 Leaves not subcordate at base, grossly elliptic-lanceolate, 2.8–3 times longer than wide; rachis hispidulous; tertiary venation finely alveolateareolate on lower surface ................................................ P. pseudocreanum Leaves with 2 pairs of basal veins arising palmately from the base, and 1 additional pair of lateral veins arising from midrib below the middle ................................................................................................ P. foveolatum Leaves pinnately veined with 3–6 lateral veins on each side ........... P. politii Ovary and fruit without any evident style; fruit exserted ............. P. nigrum Ovary and fruit with a ± developed style; fruit not exserted ................. 15

Piper 707

15(14). Upper surface of leaf densely puberulent; midvein and lateral veins densely puberulent below ......................................................... P. tepuiense 15. Upper surface of leaf glabrous; midvein and lateral veins usually glabrous below or glabrescent ......................................................... P. crassinervium 16(8). Leaves 20–60 × 10–35 cm, often with conspicuous basal lobes .............. 17 16. Leaves mainly smaller than 20–60 × 10–35 cm, basal lobes absent, inconspicuous or rarely conspicuous ............................................................ 25 17(16). Leaves pinnately veined along entire length, the principal lateral veins 12 or 13 on each side ................................................................ P. augustum 17. Leaves pinnately veined at most to the upper 1/3, the principal lateral veins 4–9 on each side ......................................................................... 18 18(17). Upper surface of leaf, as well as some veins on the upper side, pubescent ......................................................................................... P. toronotepuiense 18. Upper surface of leaf, excluding the veins, glabrous .............................. 19 19(18). Leaf base lacking lobes; spikes erect or ascending ................................. 20 19. Leaf base with lobes ± developed, if not, then spike pendent or eventually so ........................................................................................................... 21 20(19). Spikes terminating abruptly into a short conical protuberance; leaf blades broadly ovate or suborbicular-ovate, usually rounded or subcordate at base; mature spikes 8–15 cm long, 2–5 mm thick .................. P. coruscans 20. Spikes not terminating abruptly into a short conical protuberance; leaf blades broadly elliptic or ovate-elliptic, obutse or rounded at base; mature spikes 7–9.5 cm long, 2–3 mm thick ....................... P. perstipulare 21(19). Upper surface of leaf with glandular dots ....................................... P. julianii 21. Upper surface of leaf lacking glandular dots .......................................... 22 22(21). Tertiary venation of lower surface of leaf with all principal areoles 2– 7 mm long or wide ...................................................................... P. javitense 22. Tertiary venation of lower surface of leaf with all or most principal areoles exceeding 7 mm long or wide .............................................................. 23 23(22). Leaf lobes, if present, mainly equal or subequal, or if unequal, then usually shorter than petiole .......................................................... P. coruscans 23. Leaf lobes mainly unequal ....................................................................... 24 24(23). Peduncle of inflorescence glabrous or glabrescent; upper internodes glabrous or glabrescent; petioles glabrous, glabrescent, or appressedpubescent .................................................................................. P. obliquum 24. Peduncle of inflorescence usually densely pubescent; upper internodes and petioles usually densely pubescent .................................... P. cernuum 25(16). Principal lateral veins arising from midrib throughout the length of the leaf blade, or at least to the upper 1/4 .................................................. 26 25. Principal lateral veins arising from midrib to as far as the middle or to the upper 1/3 of leaf blade, but not beyond ................................................. 66 26(25). Lower surface of the leaf completely glabrous (in P. hermannii with minute, appressed marginal trichomes on lower surface) ................. 27 26. Lower surface of the leaf with at least some veins pubescent ............... 45 27(26). Leaf blades linear-lanceolate to narrowly lanceolate, 1–2.7 cm wide, 4.2– 6.8 times longer than wide .......................................... P. dunstervilleorum 27. Leaf blades without the above combination, if < 3 cm wide, then 4 times or less longer than wide ........................................................................... 28

708

P IPERACEAE

28(27). Base of leaf blade conspicuously unequal, a space of 4–30 mm separating the 2 sides at base .................................................................... P. arboreum 28. Base of leaf blade equal, or if somewhat unequal, then a space of only 1– 3 mm separating the 2 sides at base ................................................... 29 29(28). Ovary and fruit stipitate .................................................................. P. ovatum 29. Ovary and fruit not stipitate ................................................................... 30 30(29). Style ± developed or evident .................................................................... 31 30. Style absent .............................................................................................. 33 31(30). Stem with at least upper internodes pubescent; petiole 1–6 mm long ................................................................................................ P. otto-huberi 31. Stem glabrous; petiole (4–5)–32 mm long ............................................... 32 32(31). Mature peduncles 2.2–2.8 cm long, exceeding the length of mature spikes; tertiary veinlets prominent on lower surface of leaf and uniting with main lateral veins ........................................................... P. parapeltobryon 32. Mature peduncles 0.5–2.3 cm long, usually shorter than, or equaling, mature spikes; tertiary veinlets not prominent on lower surface of leaf .................................................................................................... P. jauaense 33(30). Rachis of inflorescence minutely puberulent to pilose ........................... 34 33. Rachis of inflorescence glabrous .............................................................. 37 34(33). Stem with at least upper internodes with 2 lines of recurved or retrorse trichomes extending to below base of petiole ..................... P. anonifolium 34. Stem either completely glabrous or lacking the 2 lines of retrorse trichomes described above ....................................................................... 35 35(34). Stems glabrous; petiole 10–17 mm long; fruit lobulate with lobes projecting at apex ........................................................................... P. alatabaccum 35. Stems glabrous or granular-puberulent; petiole 5–10 mm long; fruit not lobulate ................................................................................................. 36 36(35). Lower surface of leaf with elevated tertiary veins; leaves drying pale brown to brownish green; internodes granular-puberulent, scarcely or not canaliculate, the ribs or ridges obtuse or rounded when present; shrub 1–2.5 m tall ............................................................ P. bartlingianum 36. Lower surface of leaf without elevated or prominent tertiary veins; leaves drying pale green; internodes mainly glabrate, conspicuously canaliculate, the ribs or ridges acutely angled alternating with papillate depressions; subherbaceous plant 0.3–1 m tall .................... P. piscatorum 37(33). Leaf margins ± ciliate .............................................................................. 38 37. Leaf margins not ciliate ........................................................................... 39 38(37). Petioles, peduncles, and stems glabrous; petioles at sterile nodes 6– 23 mm long; spike 2.5–9 cm long ............................................. P. trigonum 38. Petioles, peduncles, and at least upper internodes of stem with short ascending to spreading trichomes; petioles 2–6 mm long; spike 1.5–2.5 cm long ......................................................................................... P. otto-huberi 39(37). Lower surface of leaf not obviously gland-dotted ................................... 40 39. Lower surface of leaf obviously gland-dotted .......................................... 41 40(39). Stem with at least upper internodes with 2 lines of recurved or retrorse trichomes extending to below base of petiole ..................... P. anonifolium 40. Stem either completely glabrous or lacking the 2 lines of retrorse trichomes described above ............................................................... P. aequale

Piper 709

41(39). Lower margin of leaf blade with minute appressed trichomes; summit of fruit conspicuously verruculose .............................................. P. hermannii 41. Lower margin of leaf blade glabrous; summit of fruit not verruculose .............................................................................................................. 42 42(41). Lower surface of leaf with tertiary veinlets prominently anastomosing, pustulose, or irregularly twisted; spikes 6–15 mm thick ..... P. tortivenulosum 42. Lower surface of leaf not as above described; spikes 2–5 mm thick ...... 43 43(42). Principal lateral veins 8–12 on each side; leaf base obtuse to rounded; petiole conspicuously vaginate-winged to or above the middle; spikes 2–2.5 cm long ....................................................................... P. cuyunianum 43. Principal lateral veins 4–8 on each side; leaf base acute to cuneiform; petiole vaginate only at base; spikes 3–4 cm long .................................... 44 44(43). Leaf blades 4–5 cm wide, 2.7–2.9 times longer than wide; petiole 7–13 mm long; peduncle 12–17 mm long; lateral veins 5–8 each side ..... P. jauaense 44. Leaf blades 2–2.7 cm wide, 3–4 times longer than wide; petiole 2– 8 mm long; peduncle 10–11 mm long; lateral veins 4–5 each side ............................................................................................... P. neblinanum 45(26). Base of leaf blade conspicuously unequal ............................................... 46 45. Base of leaf blade either subequal or slightly unequal ........................... 51 46(45). Lower lobe of leaf incurved or ± overlapping petiole .............................. 47 46. Lower lobe of leaf not as above described ............................................... 49 47(46). Ovary and fruit without a style; shrub or subherbaceous plant 0.5–2.5 m tall ..................................................................................... P. demeraranum 47. Ovary and fruit with a ± developed style; subherbaceous or suffruticose plants 0.1–0.6 m tall ............................................................................ 48 48(47). Lower surface of leaf with prominently elevated tertiary veins densely hirsute with spreading trichomes; stems retrorsely villous with trichomes to 1 mm long ................................................................... P. liesneri 48. Lower surface of leaf with faint or slightly evident tertiary veins with appressed pubescence; stems with diminutive appressed, retrorse pubescence ...................................................................... P. consanguineum 49(46). Plants subherbaceous or suffruticose, 0.1–0.5 m tall; stems with diminutive dense retrorse pubescence; spikes 1.5–2.5 cm long ......................................................................................... P. consanguineum 49. Subshrubs or small trees 0.6–8 m tall; stem not densely retrorse-pubescent; spikes 5–18 cm long .................................................................... 50 50(49). Apex of leaf blade obtuse, rounded, or shortly acute; petioles usually with minute, tuberculate enlargements .................................... P. tuberculatum 50. Apex of leaf blade slenderly acute to acuminate; petioles usually lacking tuberculate enlargements ........................................................ P. arboreum 51(45). Leaves sparsely to densely pubescent on both surfaces ......................... 52 51. Leaves with upper surface, excluding the midvein, glabrous ................ 54 52(51). Spikes 1–1.7 cm long; upper internodes of stem sparsely puberulent to glabrescent; leaves broadest below the middle or toward base .. P. guianense 52. Spikes (2–)3–11.5 cm long; upper internodes densely pubescent; leaves broadest at, near, or above the middle ................................................ 53 53(52). Principal lateral veins 6–10 on each side, fully extended to margins; petiole 1–5 mm long; spikes (2–)3–4.5 cm long ................................ P. baccans

710

P IPERACEAE

53.

54(51). 54. 55(54). 55. 56(55). 56. 57(56).

57.

58(55). 58. 59(58).

59.

60(58). 60. 61(60). 61. 62(60). 62. 63(62). 63. 64(62). 64. 65(64). 65. 66(25).

Principal lateral veins 4–6(–7) on each side, the upper ones usually ascending to leaf tip; petiole 6–15 mm long; spikes 4.5–10 cm long ................................................................................................. P. avellanum Stems with elongate spreading hispid trichomes to 3–4 mm long ....................................................................................... P. francovilleanum Stems with shorter pubescence or not as above ..................................... 55 Leaves conspicuously rugose or bullate on both sides ............................ 56 Leaves not conspicuously rugose or bullate ............................................ 58 Internodes retrorsely hirtellous .............................................................. 57 Internodes glabrous or sparsely puberulous at or just below attachment of petiole ........................................................................................ P. gentryi Leaf blades 8.5–14 × 2.5–4 cm, conspicuously mottled, the upper surface not sulcate, the veins on lower surface not elevated; spikes 17 mm long; peduncles 4.5 mm long ............................................................ P. mosaicum Leaf blades 11.5–25 × 3.5–9.5 cm, not mottled, the upper surface sulcate, the veins on the lower surface strongly elevated; spikes 20–30 mm long; peduncles 5–7 mm long .......................................................... P. holtii Ovary and fruit with a ± developed style ................................................ 59 Ovary and fruit lacking a style ................................................................ 60 Leaf blades conspicuously gland-dotted on both sides, especially on lower surface, with dark dots; petiole vaginate to base of leaf blade; stem densely short-hirtellous ......................................................... P. otto-huberi Leaf blades inconspicuously or minutely dotted, without dark dots; petioles vaginate only near base; stem minutely and densely retrorsely pubescent ........................................................................ P. consanguineum Leaf blade mainly unequal at base, with one side 3–5 mm shorter than the other ............................................................................................... 61 Leaf blade mainly equal or subequal at base, or if slightly unequal then with one side 1–2 mm shorter than the other .................................... 62 Fruit densely hirtellous or hispidulous; floral bracts densely pilose centrally at apex, densely long-fimbriate elsewhere ................ P. kegelianum Fruit glabrous or papillate; floral bracts glabrous centrally at apex, fimbriate on margins .......................................................... P. hostmannianum Leaves not ciliate on margins .................................................................. 63 Leaves ciliate on margins ........................................................................ 64 Spikes (4–)7–10 cm long; fruit densely hirtellous or hispidulous; floral bracts densely pilose centrally at apex ................................ P. kegelianum Spikes 2–6 cm long; fruit glabrous; floral bracts glabrous centrally at apex ......................................................................................... P. pubivaginatum Ciliation of leaves conspicuous, abundant, with trichomes 0.1–1 mm long .................................................................................................. P. guianense Ciliation of leaves inconspicuous, sparse, the shorter trichomes usually appressed and ascending ..................................................................... 65 Stems subretrorsely hirtellous or crisp-puberulous with trichomes to 0.2– 0.3 mm long ......................................................................... P. adenandrum Stems minutely puberulous or papillate-puberulent with non-retrorse trichomes to 0.2 mm long ........................................................... P. morilloi Stems climbing or subclimbing ................................................................ 67

Piper 711

66. Stems not climbing or subclimbing ......................................................... 69 67(66). Leaf base unequal, the upper margin 23–25 mm shorter than the lower margin ........................................................................... P. hostmannianum 67. Leaf base equal ......................................................................................... 68 68(67). Leaf blades 7–15 cm long, acute or acuminate at base ........... P. steyermarkii 68. Leaf blades 3–7 cm long, obtuse, rounded, or cordiform at base ....... P. politii 69(66). Spikes arched, recurved in the distal half ................................... P. aduncum 69. Spikes straight, erect-ascending, or nodding .......................................... 70 70(69). Base of leaf blade conspicuously unequal or at least obliquely cordiform or with the lower side 0.4–2.5 cm longer on the petiole than the upper side .............................................................................................................. 71 70. Base of leaf blade equilateral or ± inequilateral with one side 0.1–0.3 cm longer than the other side, but not cordiform ..................................... 76 71(70). Upper surface of leaf not scabrous or scabridulous ................................ 72 71. Upper surface of leaf ± scabrous or scabridulous ................................... 74 72(71). Spikes nodding, 1.5–5.7 cm long; margins of leaf blades ciliate ................................................................................................ P. cililimbum 72. Spikes erect to ascending, (5–)7–11 cm long; margins of leaf blades not ciliate .................................................................................................... 73 73(72). Midvein of upper side of leaf usually puberulous or minutely puberulent; petiole mainly (3–)5–18(–22) mm long .................................... P. dilatatum 73. Midvein of upper side of leaf glabrous; petiole 2–4 mm long (plus an additional 2 or 3 on the longer side of leaf blade) ........................ P. cumanense 74(71). Leaves mainly widest toward base; peduncle 15–25 mm long, much exceeding petiole; petiole 2–5 mm long; spikes 3.5–7 cm long; floral bracts densely pubescent over the greater part of the apex ........... P. sabanaense 74. Leaves widest at or near middle; peduncle 4–16 mm long, shorter than, equaling, or longer than petiole; petiole 4–22 mm long; spikes 5–13 cm long; floral bracts densely fimbriate on the apical margins, but the central portion of apex glabrous ............................................................... 75 75(74). Upper surface of leaf usually conspicuously scabrous; anther thecae opening from above, spreading wide horizontally; fruit laterally compressed, usually sparsely to densely puberulent ..................... P. hispidum 75. Upper surface of leaf scarcely or slightly scabrous; anther thecae opening from the sides, the thaece only slightly divergent and not spreading horizontally; fruit trigonal-obpyramidal (as seen from above), mainly glabrous or sparsely papillate ................................................. P. dilatatum 76(70). Lower surface of leaf completely glabrous .............................................. 77 76. Lower surface of leaf with at least some of veins pubescent, or elsewhere pubescent ............................................................................................. 88 77(76). Ovary and fruit with a short style 0.2–2 mm long, more conspicuous in fruit ....................................................................................................... 78 77. Ovary and fruit without a style ............................................................... 79 78(77). Spikes (5.5–)6–16.5 cm long; leaves elliptic-ovate to ovate, 5.5–12.5 cm wide ................................................................................... P. crassinervium 78. Spikes 1–4.5 cm long; leaves broadly lanceolate or oblong-lanceolate, 4– 5 cm wide .................................................................................... P. jauaense 79(77). Ultimate tertiary veinlets of the lower surface of leaf forming an elevated

712

P IPERACEAE

79. 80(79). 80. 81(80). 81. 82(81). 82. 83(82). 83. 84(82). 84. 85(84). 85.

86(81). 86. 87(86). 87. 88(76). 88.

89(88). 89. 90(89). 90. 91(90). 91. 92(91). 92. 93(92). 93. 94(93).

network irregularly branched within the larger areoles ........... P. aequale Ultimate tertiary veinlets not as described above .................................. 80 Floral bracts seen from above horseshoe-shaped ............. P. hippocrepiforme Floral bracts seen from above triangular, oval, or round ....................... 81 Lower surface of leaf obviously gland-dotted .......................................... 82 Lower surface of leaf inconspicuously or not gland-dotted .................... 86 Upper surface of leaf gland-dotted .......................................................... 83 Upper surface of leaf not gland-dotted .................................................... 84 Subherbaceous plant, 0.6 m tall; leaves firmly membranous; leaf margins ciliate; spike 2 mm thick at anthesis ................................... P. venamoense Shrub 1.5–3 m tall; leaves subcoriaceous or chartaceous; leaf margins not ciliate; spike 3–4 m thick at anthesis .................................. P. divaricatum Peduncle 12–22 mm long; fruiting spikes 8–12 cm long .............. P. lemaense Peduncle 6–12 mm long; fruiting spikes 2–7.5 cm long .......................... 85 Floral bracts densely fimbriate on margins; leaves with lateral veins usually 4 on each side ............................................................... P. tamayoanum Floral bracts mainly glabrous or sparsely papillate on margins, or shortly puberulent at the extremities; leaves with 5–7 lateral veins on each side ............................................................................. P. pseudoglabrescens Peduncle 15–25 mm long; leaves 1–4.5 cm wide, (2.7–)3.4–5.7(–8) times longer than wide ................................................................. P. bolivaranum Peduncle 6–13 mm long; leaves 4.2–10 cm wide, 1.9–2.7 times longer than wide ...................................................................................................... 87 Spike 2–7.5 cm long, apiculate; petiole 10–18 mm long ............................. .................................................................................... P. pseudoglabrescens Spike 7–8 cm long, not apiculate; petiole 2–4 mm long (plus an additional 2–3 mm on the longer side of leaf blade) .............................. P. cumanense Upper surface of leaf obviously scabrous; anther thecae opening from above, spreading wide horizontally .......................................... P. hispidum Upper surface of leaf smooth or scarcely scabridulous; anther thecae opening from the sides, the thecae either slightly divergent, or if divergent, not spreading horizontally ......................................................... 89 Peduncle 18–37 mm long .................................................... P. rupununianum Peduncle 17 mm or less long ................................................................... 90 Stems glabrous throughout their length ................................................. 91 Stems with at least upper internodes sparsely to densely pubescent or puberulent ............................................................................................ 97 Spikes when mature 1–1.7 cm long ............................................. P. guianense Spikes when mature (3–)4–16.5 cm long ................................................ 92 Leaf margins ciliate ................................................................................. 93 Leaf margins not ciliate ........................................................................... 95 Lower surface of leaf, as well as the midvein and lateral veins, pubescent ............................................................................................ P. subduidaense Lower surface of leaf with only the midvein and lateral veins pubescent .............................................................................................................. 94 Spikes 3.5 cm long; lower midvein and some of lateral veins of lower surface of leaf scarcely or sparsely and minutely appressed-pubescent ............................................................................................... P. venamoense

Piper 713

94.

Spikes 6.5–11 cm long; lower midvein and all the lateral veins of the lower surface of leaf moderately to densely appressed–pubescent ......................................................................................... P. rupununianum 95(92). Petiole 17–40 mm long; stipular development 5–7 cm long .... P. perstipulare 95. Petiole 4–15(–25) mm long; stipular development 0.6–2 cm long .......... 96 96(95). Leaf with principal lateral veins branched to the upper 5/8 or 2/3 of the length of the leaf blade, (3)4–7(–10) on each side; petiole vaginate between 1/3–1/2 its length ............................................... P. hostmannianum 96. Leaf with principal lateral veins branched from slightly below or slightly above the middle, 3–5 on each side; petiole vaginate at or near base .................................................................................................. P. dilatatum 97(90). Leaf margins ciliate ................................................................................. 98 97. Leaf margins not ciliate ......................................................................... 104 98(97). Stems with at least upper internodes retrorsely pubescent ................... 99 98. Stems with internodes not retrorsely pubescent .................................. 101 99(98). Ciliate trichomes of leaf margins conspicuously spreading, 0.3–0.5 mm long; mature fruiting spikes 7–11 cm long ........................... P. perciliatum 99. Ciliate trichomes of leaf margins appressed-ascending, minute; mature fruiting spikes 3–5.5 cm long ............................................................ 100 100(99). Mature fruit 2.5–4 mm diameter; leaves long acuminate at apex; leaf base subequilateral or somewhat inequilateral, usually varying from acute to subobtuse or obtuse .......................................... P. adenandrum 100. Mature fruit 2 mm diameter; leaves shortly acuminate, acute, or obtusely acute; leaf base usually inequilateral and mainly rounded but varying to subacute and subequilateral ............................ P. avellanum 101(98). Leaves 3–4 times longer than wide, acute at the inequilateral or subequilateral base ..................................................... P. rupununianum 101. Leaves 2–3 times longer than wide, but if 3 times longer than wide, then the base mainly rounded or cordiform ................................. 102 102(101). Stems densely villous; ciliate trichomes of leaf margins conspicuous and to 1 mm long; spikes pendent; petiole villous ........... P. cililimbum 102. Stems minutely puberulous to glabrescent; ciliate trichomes of leaf margins < 0.5 mm long; spikes erect; petiole glabrous or sparsely appressed-pubescent ..................................................................... 103 103(102). Spikes 1–1.7 cm long; subherbaceous plant 0.3–0.5 m tall, frequently emitting aerial roots from lower nodes; leaf blades 5.5–13 cm long .............................................................................................. P. guianense 103. Spikes 6–10 cm long; subligneous plant 1–1.5 m tall, lacking aerial roots; leaf blades 12–21 cm long ................................... P. subduidaense 104(97). Leaf base mainly acute at each extremity ........................................ 105 104. Leaf base rounded, obtuse, or subcordiform on one or both sides of midvein .......................................................................................... 106 105(104). Stem as well as lower surface of leaf densely pubescent; fruit densely hirtellous ........................................................................ P. wachenheimii 105. Without the above combination of characters .................... P. papilliferum 106(104). Both surfaces of leaf blade conspicuously rugose or bullate, the lower surface deeply honeycombed (alveolate); at least upper internodes densely retrorsely appressed-pubescent; known only from summit of

714

P IPERACEAE

Sororopán-tepui, Bolívar ................................................. P. subalpinum 106. Without the above combination of characters .................................. 107 107(106). Fruit densely hirtellous or hispidulous; floral bracts densely pilose centrally on the summit, elsewhere densely long-fimbriate .............. ........................................................................................... P. kegelianum 107. Fruit glabrous, papillate, or at most sparsely papillate-puberulent; floral bracts glabrous or puberulent centrally on the summit, elsewhere fimbriate on margins .......................................................... 108 108(107). Upper surface of leaf, including midvein and lateral veins, completely glabrous ..................................................................... P. hostmannianum 108. Upper surface of leaf with at least midvein pubescent .......... P. dilatatum Piper adenandrum (Miq.) C. DC. in A. DC., Prodr. 16(1): 273. 1869. —Artanthe adenandra Miq., Syst. Piperac. 515. 1844. Shrub 1–1.5 m tall. Forested tepui slopes, ca. 400 m; Amazonas (Cerro Yapacana). Guyana, Suriname, French Guiana. Piper aduncum L., Sp. Pl. 29. 1753. —Steffensia adunca (L.) Kunth, Linnaea 13: 633. 1839. —Artanthe aduncum (L.) Miq., Comm. Phytogr. 49. 1840. Piper celtidifolium H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 50. 1815 [1816]. Shrub or small tree 1–10 m tall; spikes arched-descending or recurved. Thickets, open and disturbed places, wooded slopes, along streams. Central America, West Indies, South America; 3 varieties, all in the flora area.

west to San Carlos de Río Negro, Serranía de los Pijiguaos, between Solano and San Carlos de Río Negro, Victorino). Widespread elsewhere in Venezuela from near sea level to 1900 m; Central America, West Indies, South America. P. aduncum var. cordulatum (C. DC.) Yunck., Lilloa 27: 129. 1953. —Piper angustifolium var. cordulatum C. DC. in A. DC., Prodr. 16(1): 286. 1869. —Piper elongatum var. cordulatum (C. DC.) C. DC., Repert. Spec. Nov. Regni Veg. 9: 230. 1911. Piper elongatum var. brachyarthrum Trel., Contr. U.S. Nat. Herb. 26: 37. 1927. —Piper aduncum var. brachyarthrum (Trel.) Yunck., Ann. Missouri Bot. Gard. 37: 32. 1950. Ca. 100 m; Bolívar (middle Río Caura). Widespread elsewhere in Venezuela.

Key to the Varieties P. aduncum 1. Upper surface of leaves not scabrous ............................ var. garcia-barrigae 1. Upper surface of leaves ± scabrous ........ 2 2. Stems glabrous, glabrescent, or sparsely and shortly pubescent; lower surface of leaves glabrescent or pubescent ............. ...................................... var. aduncum 2. Stems ± moderately to densely villous-pubescent with loosely spreading trichomes; lower surface of leaves rather densely and softly villous ........................ .................................. var. cordulatum P. aduncum var. aduncum 100–1500 m; Bolívar (near Roraimatepui, Gran Sabana, between Santa Elena de Uairén and El Paujil, 32.5 km northwest of Santa Elena de Uairén), Amazonas (north-

P. aduncum var. garcia-barrigae Trel. & Yunck., Piperac. N. South Amer. 1: 251. 1950. Ca. 100 m; Amazonas (San Carlos de Río Negro, between Solano and San Carlos de Río Negro). Colombia, Bolivia, and probably elsewhere in South America. Piper aequale Vahl, Eclog. Amer. 1: 4, pl. 3. 1796. —Schilleria aequalis (Vahl) Kunth, Linnaea 13: 687. 1839. —Anisillo. Piper caripense H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 48. 1815 [1816]. —Artanthe caripensis (H.B.K.) Miq., Syst. Piperac. 522. 1844. Piper funckii C. DC. in A. DC., Prodr. 16(1): 310. 1869. Piper johnstonii C. DC., Proc. Amer. Acad. Arts 40: 685. 1905.

Piper 715

Piper santa-martanum C. DC., Annuaire Conserv. Jard. Bot. Genève 21: 231. 1920. —Piper aequale var. santamartanum (C. DC.) Trel. & Yunck., Piperac. N. South Amer. 1: 219. 1950. Piper zedigodiense Trel. ex L. Williams, Explor. Bot. Guayana Venezolana 177. 1942. Glabrous shrub 0.5–3 m tall. Wet lowland to lower montane forests, mostly 100–600 m; Delta Amacuro (near Sierra Imataca), Bolívar (Altiplanicie de Nuria, Auyán-tepui, km 106 south of El Dorado, headwaters of Río Paramichi, Salto Pará, Río Suapure, south of Cerro Venado), Amazonas (near Río Mawarinuma, Serranía Yutajé). Common elsewhere in Venezuela; Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil. Piper alatabaccum Trel. & Yunck., Piperac. N. South Amer. 1: 408, fig. 371. 1950. Glabrous shrub 1–2.5 m tall; fruit with 4 enlarged lobes at summit. Lower montane forests, 300–800 m; Bolívar (Amaruay-tepui, El Abismo, Río Chicanán), Amazonas (Cerro Aracamuni, Cerro Yapacana). Guyana, Suriname, French Guiana.

anonifolia Kunth, Linnaea 13: 619. 1839, —Piper anonifolium Kunth, Linnaea 13: 619. 1839, nom. invalid., as synonym, “anonaefolium.” —Artanthe anonifolia (Kunth) Miq., Syst. Piperac. 504. 1844. Piper citrifolium auct. non Lam.: sensu Trel. & Yunck., Piperac. N. South Amer. 1: 384. 1950. Shrub or low suffruticose plant 0.8–1.5 m tall, the stems with 2 lines of recurved or retrorse setae-like trichomes. Guyana, Suriname, French Guiana, Amazonian Brazil; 3 varieties, 2 of these in Venezuela and the flora area. Key to the Varieties of P. anonifolium 1. Lower surface of leaves, especially midrib and principal veins, hirtellous ................ ................................ var. parkerianum 1. Lower surface of leaves glabrous .............. ................................. var. anonifolium P. anonifolium var. anonifolium. —Anicillo manso. Deciduous or lowland forests, wet forested slopes, 100–1500 m; Delta Amacuro (Río Cuyubini), Bolívar (Auyán-tepui, between Tumeremo and Bochinchito, between Upata and San Félix). Distribution as in the species. ◆Fig. 608. Two forms have been recognized by Yuncker in this variety (Bol. Inst. Bot. (São Paulo) 3: 82. 1966): f. angustifolium with leaves 4 times longer than wide, 12–20 cm long and f. parvifolium with leaves 2.5–3 times longer than wide, 9–13 cm long.

Piper amalago L., Sp. Pl. 29. 1753. Piper amalago var. medium (Jacq.) Yunck., Brittonia 14: 189. 1962. —Piper medium Jacq., Collectanea 1: pl. 8. 1786 [1787]. Piper amalago var. medium f. ceanothifolium (H.B.K.) Steyerm., Fl. Venez. 2(2): 322. 1984. Piper ceanothifolium H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 56. 1815 [1816]. Widely distributed in Central America, West Indies, South America to southern Brazil, Argentina, Bolivia. Shrub 1–6 m tall; leaves palmately veined. Deciduous and riparian forests, 100– 500 m; Bolívar (Río Botanamo, 7 km west of El Manteco, Represa Guri). Venezuelan Coastal Cordillera, lower slopes of the Andes, Zulia (Sierra de Perijá); Central America, West Indies, northern South America to southern Brazil, Bolivia, Argentina. ◆Fig. 607.

P. anonifolium var. parkerianum (Miq.) Steyerm., Fl. Venez. 2(2): 328. 1984. —Artanthe parkeriana Miq. in Hook., London J. Bot. 4: 467. 1845. —Piper parkerianum (Miq.) C. DC. in A. DC., Prodr. 16(1). 373. 1869. Piper citrifolium var. parkerianum (Miq.) Trel. & Yunck., Piperac. N. South Amer. 1: 385. 1950. Lower montane forests, 400–1200 m; Bolívar (km 134 of El Dorado-Santa Elena road, near Río Chicanán). Guyana, Suriname, French Guiana.

Piper anonifolium (Kunth) C. DC. in A. DC., Prodr. 16(1): 298. 1869. —Steffensia

Piper arboreum Aubl., Hist. Pl. Guiane 23. 1775.

716

P IPERACEAE

Piper geniculatum Sw., Prodr. 15. 1788 Piper macrophyllum Sw., Prodr. 15. 1788. Piper nitidum Sw., Prodr. 15. 1788. Piper verrucosum Sw., Prodr. 15. 1788. Piper falcifolium Trel., Contr. U.S. Natl. Herb. 26: 25. 1927. —Piper arboreum var. falcifolium (Trel.) Yunck., Ann. Missouri Bot. Gard. 37: 65. 1950. Shrub or small tree; spikes slender, elongate. Common along streams in moist forests, 100–1300 m. Widespread elsewhere in Venezuela; Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil, Bolivia; 2 varieties, both in the flora area. Key to the Varieties of P. arboreum 1. Stems, petioles, peduncles, and leaves glabrous ...................... var. arboreum 1. Stems, petioles, and peduncles pubescent; midrib pubescent above and below; lower surface of leaf ± pubescent to glabrate ........................ var. hirtellum P. arboreum var. arboreum. —Anisillo, Cordoncillo, Guayuyu montañera. 100-1300 m; Delta Amacuro (near el Palmar), Bolívar (Altiplanicie de Nuria, Río Carrao, near Río Cunucunuma, drainage of Río Cuyuní, Cerro Duida, El Dorado, 20-35 km southwest of El Manteco, near El Paují, Represa Guri, west Icabarú, Sierra Imataca, Ptari-tepui, between Tumeremo to Bochinchito), Amazonas (Río Cunucunuma, upper Río Orinoco, southeast of Puerto Ayacucho, San Juan de Manipiare, south of Sierra Parima, Río Ventuari, Cerro Yapacana). Distribution as in species. ◆Fig. 612. P. arboreum var. hirtellum Yunck., Ann. Missouri Bot. Gard. 37: 64. 1950. Steffensia luschnathiana Kunth, Linnaea 13: 617. 1839. —Artanthe luschnathiana (Kunth) Miq., Syst. Piperac. 494. 1844. Artanthe obrumbata Miq., Linnaea 20: 142. 1847. —Piper obrumbatum (Miq.) C. DC. in A. DC., Prodr. 16(1): 265. 1869. Piper geniculatum var. latifolium C. DC. in A. DC., Prodr. 16(1): 265. 1869. —Piper arboreum var. latifolium (C. DC.) Yunck., Bol. Inst. Bot. (São Paulo) 3: 82. 1966. Piper geniculatum Sw. var. B [beta] C. DC. in Urb., Symb. Antill. 3: 179. 1902.

Riparian and wet forests, often overlying sandstone on tepuis or on igneous outcrops, 100–500(–1100) m; Bolívar (near Río Hacha, near Icabarú, Gran Sabana, Cerro Pitón), Amazonas (near Río Cunucunuma, Cerro Duida, Sierra de la Neblina, middle Río Pacimoni, 30 km east from Puerto Ayacucho, between Yavita and Pimichín). Widespread in Venezuela; Panama, Guyana, Suriname, French Guiana, Brazil. Piper augustum Rudge, Pl. Guian. 1: 10. 1805. —Piper angustum Rudge ex Spreng., Syst. Veg. 1: 108. 1825, nom. superfl. sphalm. —Artanthe augusta (Rudge) Miq., Syst. Piperac. 398. 1844. Wet forests and forested slopes, often overlying sandstone, 200–900 m; Bolívar (Gran Sabana), Amazonas (Cerro Yutajé). Sierra de Perijá; Panama, Colombia, Guyana, Suriname, French Guiana, northern Brazil. Piper avellanum (Miq.) C. DC. in A. DC., Prodr. 16(1): 302. 1869. —Artanthe avellana Miq., Syst. Piperac. 478. 1844. Piper statarium Trel. & Yunck., Piperac. N. South Amer. 1: 339, fig. 310. 1950. Piper tonoroanum Trel. & Yunck., Piperac. N. South Amer. 1: 328, fig. 295. 1950. Small suffruticose plant 0.6–1.5 m tall. Riparian and wet lowland forests, sometimes on steep slopes overlying sandstone, 100– 300 m; Bolívar (Río Tonoro tributary of Río Paragua), Amazonas (Cerro Duida, Río Mavaca, near Río Ocamo, 30-35 km southeast of Puerto Ayacucho, between Puerto Ayacucho and Samariapo near San Fernando de Atabapo, Trapichote, Caño Yagua between Yavita and Pimichín). Mérida; Guyana, Suriname, French Guiana. ◆Fig. 613. Piper baccans (Miq.) C. DC. in A. DC., Prodr. 16(1): 279. 1869. —Artanthe baccans Miq., Linnaea 20: 166. 1847. Venezuela, Brazil; 2 varieties, 1 of these in Venezuela. P. baccans var. sancarlosianum (C. DC.) Steyerm., Fl. Venez. 2(2): 349. 1984. —Piper sancarlosianum C. DC. in A. DC., Prodr. 16(1): 337. 1869. —Borrajón, Caneya gruya. Piper amazonasense Trel. & Yunck., Piperac. N. South Amer. 1: 392, fig. 352. 1950.

Piper 717

Suffruticose plant 0.8–2 m tall with densely pubescent stems; lower surface of leaves pubescent; spikes very short, pendulous. Wet forests in sandy, lateritic, or granitic terrain, ca. 100 m; Amazonas (near Piedra Cocuy, drainage of Río Negro, Río Maricapuro tributary south of Río Orinoco, San Carlos de Río Negro). Brazil. ◆ Fig. 614. Piper bartlingianum (Miq.) C. DC. in A. DC., Prodr. 16(1): 257. 1869. —Artanthe bartlingianum Miq., Syst. Piperac. 510. 1844. Piper bartlingianum var. parimanum (Trel.) Steyerm., Fl. Venez. 2(2): 352. 1984. —Piper parimanum Trel., Bull. Torrey Bot. Club 58: 354. 1931. Conspicuously nodose shrub 1–2.5 m tall with densely glandular-puberulent stems; fruits ovoid-tetragonal, obtusely angled, apically papillate. Rainforests, 100–600(–900) m, Bolívar (Río Ichún, Río Paragua), Amazonas (Caño Caname, Río Manaviche, Río Ocamo, between Río Ocamo and Río Matacuni, upper Orinoco, Río Padamo, 10 km from Puerto Ayacucho, below San Fernando de Atabapo, Cerro Yapacana). Guyana, Suriname, French Guiana, northern Brazil. ◆Fig. 618. Piper bolivaranum Yunck., Fieldiana, Bot. 28(1): 205. 1951. Piper deminutum Yunck., Mem. New York Bot. Gard. 9: 422. 1957. Dwarf, ligneous, glabrous plant 0.1–1.5 m tall; leaves equilateral, long-acuminate; fruits short, pendulous. Common on lower and middle slopes of tepuis and in wet forests at their base, 300–1700 m; Bolívar (Río Chicanán, Gran Sabana, near Icabarú, Macizo del Chimantá (Abácapa-tepui, Apacará-tepui, Chimantá-tepui, Toronó-tepui), Ptari-tepui, Río Tírica, Cerro Venamo). Adjacent Guyana. ◆Fig. 616. Piper cernuum Vell., Fl. Flumin. 26. 1825 [1829]. Steffensia eximia Kunth, Linnaea 13: 665. 1839. —Piper eximium Kunth, Linnaea 13: 665. 1839, as synonym. —Artanthe eximia (Kunth) Miq., Syst. Piperac. 393. 1844. —Piper obliquum var. eximium (Kunth) C. DC. in A. DC., Prodr. 16(1): 307. 1869.

Artanthe spectabilis Miq., Linnaea 20: 138. 1847. Piper cabellense C. DC. in A. DC., Prodr. 16(1): 306. 1869. Piper gigantifolium C. DC. in A. DC., Prodr. 16(1): 305. 1869. Shrub or tree 2.5–15 m tall with upper part of stems densely pubescent; leaves 30– 60 × 20–35 cm, deeply lobed; mature spikes 20–63 cm long, pendent. Venezuela, Colombia, Brazil; 4 varieties, all in Venezuela, 2 of these in the flora area. Key to the Varieties of P. cernuum 1. Peduncle 2–4(–6.5) cm long ....................... ....................................... var. cernuum 1. Peduncle 7–10 cm long .............................. ............................ var. perlongispicum P. cernuum var. cernuum 100–400 m; Amazonas (Cerro Duida, Cerro Yapacana, base of Cerro Yutajé). Distribution as in species. P. cernuum var. perlongispicum (Yunck.) Steyerm., Fl. Venez. 2(2): 372. 1984. —Piper perlongispicum Yunck., Bol. Soc. Venez. Ci. Nat. 23(101): 89, fig. 10. 1962. Wet forests near the Brazilian-Venezuelan border, 600–800 m; Bolívar (northeast of Serranía Marutaní). Endemic. Piper cililimbum Yunck., Bol. Inst. Bot. (São Paulo) 3: 90. 1966. —Canilla de grulla. Shrub 1.3–2.5 m tall; leaves rugose, conspicuously ciliate; spikes pendent. evergreen lowland to upland forests, 100–800 m; Bolívar (near San Carlos de Río Negro, San Ignacio), Amazonas (Río Cunucunuma, south of Cerro Yutajé, near San Carlos de Río Negro). Brazil. ◆Fig. 617. Piper consanguineum (Kunth) C. DC. in A. DC., Prodr. 16(1): 296. 1869. —Steffensia consanguinea Kunth, Linnaea 13: 623. 1839. —Piper consanguineum Kunth, Linnaea 13: 623. 1839, as synonym. —Artanthe consanguinea (Kunth) Miq., Syst. Piperac. 529. 1844. Piper surinamense C. DC. in A. DC., Prodr. 16(1): 296. 1869. Piper leprieuri Pulle, Enum. Vasc. Pl. Surinam 141. 1906.

718

P IPERACEAE

Dwarf, subherbaceous, sparsely branched plant 0.1–0.5 m tall; leaves rounded, subcordate, short; spikes erect, 1.5–2.5 cm long. Forests bordering savannas or openings, 300–800(–1000) m; Bolívar (Auyán-tepui, Río Caura, near Icabarú, near Serranía PiaZoi). Guyana, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 609.

Slender shrub 2–2.5 m tall, mainly glabrous throughout, lower surface of leaves sparsely gland-dotted; petioles short. Gallery forests, 900–1100 m; Bolívar (near Santa Elena de Uairén). Reported from Sucre, but no specimen has been collected since the time of Humboldt and Bonpland; Colombia, Ecuador, northern Brazil. ◆Fig. 625.

Piper coruscans H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 53. 1815 [1816]. Piper coruscans var. membranaceum (C. DC.) Steyerm., Fl. Venez. 2(2): 383. 1984. —Piper pseudochurumayu var. membranaceum C. DC. in A. DC., Prodr. 16(1): 288. 1869. Steffensia pseudochurumayu Kunth, Linnaea 13: 662. 1839. —Artanthe pseudochurumayu (Kunth) Miq., Syst. Piperac. 407. 1844. Artanthe amazonica Miq., Linnaea 20: 146. 1847. —Piper amazonicum (Miq.) C. DC. in A. DC., Prodr. 16(1): 272. 1869. Piper orenocanum C. DC., Bull. Herb. Boissier 6: 564. 1898. Piper wurdackii Yunck., Mem. New York Bot. Gard. 10(5): 41. 1964. Shrub or suffruticose plant 1–3 m tall; leaves large, broadly ovate or suborbicularovate, rounded or subcordiform at base; spikes erect, 8–15 cm long. Evergreen lowland to montane forests, near sea level to 1300 m; Delta Amacuro (near Caño Macareo, lower Río Orinoco, Santa Catalina, Tucupita), Bolívar (Río Caura), Amazonas (Sierra de la Neblina). Colombia, Ecuador, Northern and eastern Brazil. ◆Fig. 622.

Piper cuyunianum Steyerm., Fl. Venez. 2(2): 391, fig. 53. 1984. Suffruticose glabrous plant 0.3–0.4 m tall; spikes short, 2–5 cm long, apiculate. Wet forests below high sandstone bluffs, ca. 500 m; southeastern Bolívar (headwaters of Río Chicanán). Endemic.

Piper crassinervium H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 48. 1815 [1816]. Shrub or small tree 2–4 m tall; spikes erect 5.5–16.5 cm long. Montane forests, 900–1700 m; Bolívar (58 km west of Santa Elena de Uairén, Quebrada O-paru-má near Kavanayén), Amazonas (west of Serranía Yutajé). North-central and northeastern Venezuela; Costa Rica, Panama, Colombia Peru, Brazil. Piper cumanense H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 47. 1815 [1816]. Piper variegatum H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 47. 1815 [1816], non Pers. 1805.

Piper demeraranum (Miq.) C. DC. in A. DC., Prodr. 16(1): 298. 1869. —Artanthe demerarana Miq. in Hook., London J. Bot. 4: 464. 1845. —Anisillo, Nudillo, Pabajuto metsaja, Rabo de rata, Raiz de muela, Sekum-warei-yek. Piper rubescens C. DC. in A. DC., Prodr. 16(1): 300. 1869. —Piper demeraranum var. rubescens (C. DC.) Trel. & Yunck., Piperac. N. South Amer. 1: 380. 1950. Piper esmeraldanum Trel., Bull. Torrey Bot. Club. 58: 353. 1931. Piper cauranum Trel. in L. Williams, Explor. Bot. Guayana Venezolana 175. 1942. Subherbaceous to shrubby plant, 0.5–2.5 m tall, stems pubescent; leaves cordulate or auriculate at base, the lower lobe usually incurved. Moist lowland to montane forests, 50–1200 m; Delta Amacuro (east of Río Toro), Bolívar (Isla Anacoco, Auyán-tepui, middle Caura, near Río Cuchivero, north of Río Hacha, Macizo del Chimantá [Chimantátepui], Río Paragua, Río Parguaza, Cerro Pitón, below of Sororán-tepui, Río Toro), Amazonas (upper Caño Caname, Capibara, Río Cunucunuma, Río Guanía, Cerro Huachamacari, near Macuruco, Sierra Parima, between Puerto Ayacucho and Gavilán, Río Siapa near Cerro Aracamuni, between Samariapo and Puerto Ayacucho, Cerro Sipapo, Tamatama, Cerro Ualipano, Cerro Yapacana). Apure, northwest Venezuela; Trinidad, Guyana, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 610.

Piper 719

Piper dilatatum Rich., Act. Soc. Hist. Nat. Par. 105. 1792. Shrub 1–4 m tall; leaves membranous, minutely pubescent along upper midrib. Evergreen to semideciduous forests, near sea level to 600 m; Delta Amacuro (lower Río Orinoco, Río Cuyubiní), Bolívar (Altiplanicie de Nuria, near El Dorado, northwest of El Manteco, Río Caura, Río Paragua, Río Parguaza). Widespread elsewhere in Venezuela; Costa Rica, Panama, West Indies, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil. Four forms have been recognized from the flora area: f. dilatatifolium (Trel. & Yunck.) Steyerm., f. guayranum (C. DC.) Steyerm., f. taboganum (C. DC.) Steyerm., and the typical form. These are based on variations in leaf shape and pubescence differences. Piper divaricatum G. Mey., Prim. Fl. Esseq. 15: 1818. Piper crassum Vell. Conc., Fl. Flumin. 25. 1825 [1829]. Piper glabrilimbum C. DC., Verh. Bot. Ver. Brand. 47: 107. 1905. Mostly glabrous shrub 1.5–3 m tall; spikes pendent, 3.5–6 cm long. Evergreen lowland forest, near sea level to 100 m; Delta Amacuro (between Amacuro and the mouth of Deadwater Creek Moat). Guyana, Suriname, French Guiana, Brazil, Bolivia. This species is easily recognized by the abundance of glandular dots on the stems, petioles, and peduncles, and the lack of pubescence. Piper dunstervilleorum Steyerm., Fl. Venez. 2(2): 418, fig. 58. 1984. Subherbaceous glabrous plant 0.5–1 m tall; leaves lanceolate, narrowly elongate, 6 times longer than wide; peduncles filiform, 2.5–3 cm long, ± equaling the spike. Wet montane forests, ca. 1300–1400 m; eastern Bolívar (drainage of Río Cuyuní). Endemic. Piper foveolatum Kunth ex C. DC. in A. DC., Prodr. 16(1): 329. 1869. —Piper poiteanum Kunth, Linnaea 13: 606. 1839, nom invalid., pro syn. —Enckea dubia Kunth, Linnaea 13: 607. 1839, non Piper dubium Miq.

Climbing shrub to 2–4 m; leaves equilaterally subcordulate or cordiform, palmately veined with usually 2 pairs of basal veins on each side of midrib, and an additional pair above. Riparian forests, bases of sandstone bluffs, 100–1400 m; Bolívar (Río Chicanán), Amazonas (Cerro Aracamuni, Cerro Huachamacari). Guyana, Suriname, French Guiana. ◆Fig. 621. Piper francovilleanum C. DC., J. Bot. 4: 219. 1866. Piper perlongivillosum Yunck., Fieldiana, Bot. 28: 207. 1951. Suffruticose plant 0.3–1.2 m tall with elongate hispid trichomes on stems; stipules conspicuous, persistent, caudate; leaves rugose, conspicuously ciliate. Evergreen lowland to lower montane forests, 100–600(–1300) m; Bolívar (upper Río Paragua, Sierra Parima), Amazonas (Río Ararí, Piedra Arauicaua, Río Cunucunuma, Cerro Duida, upper Orinoco, upper Río Padamo, Río Siapa). Amazonian Brazil. Piper gentryi Steyerm., Ann. Missouri Bot. Gard. 74: 89. 1987. Shrub 1–2 m tall with glabrous stems and peduncles, leaves with 11–15 principal veins on each side; bracts conspicuously fimbriate. Evergreen lowland forests, 100–200 m; Bolívar (Sierra de la Neblina). Endemic. Piper guianense (Klotzsch) C. DC. in A. DC., Prodr. 16(1): 301. 1869. —Artanthe guianensis Klotzsch, J. Bot. (Hooker) 4: 322. 1842. Subherbaceous plant or soft-stemmed shrub 0.3–0.5 m tall, usually developing aerial roots from lower nodes, with very short, erect spikes 1.2–1.7 cm long. Riparian forests, ca. 100 m; Amazonas (near San Carlos de Río Negro, near Solano). Colombia, Guyana, Peru northern Brazil. ◆Fig. 630. Piper hermannii Trel. & Yunck., Piperac. N. South Amer. 1: 397, fig. 358. 1950. Glabrous, subherbaceous or slightly ligneous plant 1–3 m tall; leaves broadly oblongelliptic, the lower surface conspicuously gland-dotted; stems and petioles conspicuously gland-dotted; spikes erect or pendent, 1.5–3 cm long; fruits conspicuously verru-

720

P IPERACEAE

culose apically. Evergreen lowland forests, ca. 100 m; Amazonas (near San Carlos de Río Negro). Apure, Barinas; southeastern Colombia.

widespread species of Piper in the flora area. Such a broad delimitation is tentative, however, and further study may show that several distinct taxa are involved.

Piper hippocrepiforme Steyerm., Fl. Venez. 2(2): 440, fig. 63. 1984. Completely glabrous suffruticose plant 2 m tall; leaves lanceolate-elliptic; spikes 2.5– 3 cm long; bracts horseshoe-shaped. Dwarf moist evergreen forests around cascades, ca. 1400–1500 m; eastern Bolívar (slopes of Cerro Venamo, drainage of Río Cuyuní). Endemic. ◆Fig. 615.

Piper holtii Trel. & Yunck., Piperac. N. South Amer. 1: 356, fig. 323. 1950. Suffruticose plant 0.5–1 m tall with retrorsely pubescent stems; leaves rugose pinnately veined throughout length of blade; petioles vaginate to base of leaf blade; spikes 2–3 cm long, erect. Around Mauritia palm swamps and lowland forests on white sand, ca. 100 m; Amazonas (Capibara, Maroa, northeast of San Carlos de Río Negro, Cerro Yapacana). Endemic.

Piper hispidum Sw., Prodr. 15. 1788. —Cordoncillo. Piper hirsutum Sw., Fl. Ind. Occ. 1: 60. 1797. —Steffensia hirsuta (Sw.) Kunth, Linnaea 13: 640. 1839. —Artanthe hirsuta (Sw.) Miq., Syst. Piperac. 446. 1844. Piper hispidum H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 50. 1815 [1816], non Sw. 1788. Artanthe asperifolia (Ruiz & Pav.) Miq., Syst. Piperac. 441. 1844. —Piper asperifolium Ruiz & Pav., Flora 1: 37, pl. 56. 1798. Shrub 2–3(–5) m tall with hispidulous or appressed trichomes on stems; upper surface of leaf scabrous; spikes 6–13 cm long, erect or ascending. Along streams, thickets and borders of wet evergreen or semideciduous forests, near sea level to 900 m; Delta Amacuro (Río Acure, between Amacuro and mouth of Deadwater Creek Moat, confluence of Río Cuyubini and Río Amacuro, upper Río Cuyubini, lower Río Orinoco), Bolívar (Río Chicanán, Río Cuyubini, Represa Guri, Río Merewari, 32.5 km northeast of Santa Elena de Uairén, Río Venamo), Amazonas (Río Manapiare, Mavaca, Salto Tencua). Widespread elsewhere in Venezuela; Central America, West Indies, common in South America. Three forms have been recognized from the flora area: f. lanceolatum (Trel. & Yunck.) Steyerm., f. surinamense (Miq.) Steyerm., and the typical form. These are based on leaf sizes and shapes. As treated here, Piper hispidum is very broadly circumbscribed and consequently one of the commonest and most

Piper hostmannianum (Miq.) C. DC. in A. DC., Prodr. 16(1): 287. 1869. —Artanthe hostmanniana Miq. in Hook., London J. Bot. 4: 465. 1845. Piper hostmannianum var. ramiflorum C. DC. in A. DC., Prodr. 16(1): 287. 1869. —Artanthe ramiflora Miq. ex C. DC. in A. DC., Prodr. 16(1): 288. 1869, as synonym. Piper subcrassifolium Yunck., Bol. Soc. Venez. Ci. Nat. 23(101): 68, fig. 4. 1962. Piper rio-paraguanum Trel. ex V. Badillo in Pittier et al., Cat. Fl. Venez. 1: 243. 1945, nom. nud. Piper hostmannianum var. glabrirameum Trel. & Yunck., Piperac. N. South Amer. 1: 307. 1950. Suffruticose, shrub, or small tree, or sometimes somewhat climbing or subherbaceous, 1.5–4 m tall, varying in shape, size, and venation of leaves and pubescence of stems; leaves markedly lustrous on upper surface. Wet forests, sometimes in swampy soil of Mauritia palm swamps, 100–200 (–1300) m; Delta Amacuro (near Araguao, Caño Guiniquina), Bolívar (Río Aponguao, headwaters of Río Chicanán, Río Cuyuní, Hato Divina Pastora, between El Dorado and Santa Elena de Uairén, south of El Dorado, southwest of El Manteco, Río Paragua, Cerro Venamo), Amazonas (Río Cunucunuma, Mavaca, 35 km southeast of Puerto Ayacucho, Santa María de los Guaicas, Tamatama, Trapichote, upper Río Orinoco). Northeastern South America. ◆Fig. 631.

Piper 721

Piper jauaense Steyerm., Bol. Soc. Venez. Ci. Nat. 132: 319. 1976. Shrub or small tree 1.5–3 m tall with glabrous stems, leaves, and peduncles; leaves dark-dotted on both sides. Wet montane forests on tepuis, 1500–1900 m; Bolívar (Cerro Jaua), Amazonas (Cerro Aracamuni, Cerro Marahuaka). Endemic. ◆Fig. 626. Piper javitense H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 54. 1815 [1816]. —Artanthe javitensis (H.B.K.) Miq., Syst. Piperac. 401. 1844. Climbing shrub with densely tomentose upper internodes; leaves large, broadly acute, unequally lobed at base, the lower surface villous. Lowland riparian forests, ca. 100 m; Amazonas (along Río Temi, Yavita). Endemic.

Amazonas (Cerro Aracamuni). Endemic. ◆Fig. 636. Piper liesneri Steyerm., Fl. Venez. 2(2): 467, fig. 67. 1984. Suffruticose plant 0.3–0.6 m tall with densely retrorsely pubescent stems, short peduncles and spikes; leaves unequally cordiform-based, with manifestly reticulate venation. Evergreen lowland forests, ca. 100 m; Amazonas (near San Carlos de Río Negro). Endemic.

Piper kegelianum (Miq.) C. DC. in A. DC., Prodr. 16(1): 372. 1869. —Artanthe kegeliana Miq., Linnaea 22: 77. 1849. Piper agostinii Yunck., Bol. Soc. Venez. Ci. Nat. 23(101): 101. 1962. Shrub 1–2 m tall with the upper surface of leaf strongly rugose-reticulate; fruits hirtellous, gland-dotted above. Wet lowland forests, 50–100 m; Bolívar (Isla Anacoco, Río Cuyuní basin). Monagas; Guyana, Suriname, French Guiana, eastern Brazil. ◆Fig. 633.

Piper marginatum Jacq., Icon. Pl. Rar. 2(11): pl. 215. 1786–1793 [1792]. —Schilleria marginata (Jacq.) Kunth, Linnaea 13: 718. 1839. —Artanthe marginata (Jacq.) Miq., Syst. Piperac. 381. 1844. —Anisillo, Canilla de grulla, Cordoncillo, Cordoncillo negro, Guayuyo, Santa Maria. Piper caudatum Vahl, Eclog. Amer. 1: 3. 1796. —Schilleria caudata (Vahl) Kunth, Linnaea 13: 716. 1839. —Artanthe caudata (Vahl) Miq., Syst. Piperac. 380. 1844. Piper anisatum H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 58. 1815 [1816]. —Piper marginatum var. anisatum (H.B.K.) C. DC. in Urb., Symb. Antill. 3: 172. 1902. Soft-stemmed shrub 1–3 m tall with anise-like fragrance; leaves large, round-ovate or broadly ovate, palmately 9–11-veined; spikes 7–24 cm long, slender, arched or recurving. Central America, West Indies, South America to Brazil and Ecuador; 2 varieties, all in Venezuela, 1 of these in the flora area. This is one of the most common species of Piper in the flora area, varying in pubescence of the leaves and stems.

Piper lemaense Yunck., Bol. Soc. Venez. Ci. Nat. 23(101): 66, fig. 3. 1962. Glabrous suffruticose plant 0.5–2 m tall; leaves with 4 principal veins on each side of midrib, areolate tertiary venation, glandular-dotted on lower side; petioles vaginate to base of leaf blade. Wet montane forests, 300– 1200 m; Bolívar (upper Río Cuyuní, between El Dorado and La Gran Sabana, between El Dorado and Santa Elena de Uairén, La Escalera Sierra de Lema, Río Venamo),

P. marginatum var. marginatum Thickets and wet or dry forests, near sea level to 400(–1300) m; Delta Amacuro (Río Orocoima), Bolívar (Altiplanicie de Nuria, Isla Anacoco, Canaima, Río Cuyuní, north of Río Hacha, west of El Manteco, Salto Pará, La Paragua, Río Paragua, Río Venamo), Amazonas (below headwaters of Río Atabapo, Rí Parucito, near Puerto Ayacucho, between Puerto Ayacucho and Sanariapo, near San Fernando de Atabapo, near Santa

Piper julianii Callejas, nom. nov. Piper pavasense Steyerm., Fl. Venez. 2(2): 510, fig. 76. 1984, non Trel. 1929. Shrub 3 m tall; leaves large, unequally cordiform at base, the upper surface glanddotted. Evergreen lowland to lower montane forests, 200–300 m; Bolívar (Río Caura, Salto de Pará). Endemic.

722

P IPERACEAE

Bárbara del Orinoco, Tamatama, Yavita). ◆Fig. 632. Two forms have been recognized from the flora area: f. catalpifolium (H.B.K.) Steyerm. with one or both surfaces of leaves pubescent and the veins pubescent on lower surface or on both sides versus the typical form which has leaves that are usually glabrous. Piper morilloi Steyerm., Fl. Venez. 2(2): 489, fig. 71. 1984. Suffruticose plant 1.5 m tall with papillate-puberulent stems and peduncles; spikes 2.3–2.7 cm long; leaves pinnately veined throughout their length. Evergreen lowland forests, 100–200 m; Amazonas (near Solano). Endemic. ◆Fig. 635. Piper mosaicum Steyerm., Ann. Missouri Bot. Gard. 76: 964. 1989. Shrub 1 m tall; leaves rugulose and pinnately veined to upper 1/4, the upper surface gray-silvery-splotched; spikes 1.7 cm long. Dwarf tepui forests bordering meadows, 1500–1600 m; Amazonas (summit of Cerro Aracamuni). Endemic. Piper neblinanum Yunck., Mem. New York Bot. Gard. 10(5): 43. 1963. Small shrub 1–2 m tall; leaves pinnately veined throughout their length; spikes 1.5–3.1 cm long. Dwarf tepui slope forests, 700–1200 m; Bolívar, (Cerro Sarisariñama), Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 635. Piper nigrum L., Sp. Pl. 28. 1753. —Pimienta negra. Glabrous shrubby vine; spikes 5–10 cm long, pendulous. Evergreen lowland forest, near sea level to 50 m; Delta Amacuro (near Tucupita). Widely cultivated in lowland tropics of both hemispheres, native to India. The white and black pepper of commerce is produced from the fruits of this species. Piper obliquum Ruíz & Pav., Fl. Peruv. 1: 37, pl. 63. 1798. —Steffensia obliqua (Ruíz Lopez & Pavón) Kunth, Linnaea 13: 663. 1839. —Artanthe magnifica Miq., Syst. Piperac. 391. 1844, nom. illegit. Piper flagellare H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 54. 1815 [1816].

—Artanthe flagellaris (H.B.K.) Miq., Syst. Piperac. 393. 1844. Piper ruizianum D. Dietr., Syn. Pl. 1: 118. 1839. Artanthe superba Miq., Linnaea 20: 137, 1847. Piper oparumaense Yunck., Fieldiana, Bot. 28(1): 206. 1951. Piper aristeguietae Yunck., Acta Biol. Venez. 2: 320. 1958. Shrub or small tree 1.5–6 m tall, with glabrous or glabrescent upper internodes; leaves large, rugose, unequally cordiform at base, with relatively short lobes; spikes 30– 71 cm long, pendent. Evergreen lowland to montane forests, 100–1300 m; Bolívar (Auyán-tepui, upper Río Cuyuní, El Dorado, between El Dorado and Santa Elena de Uairén, Gran Sabana, northeast of Luepa, Uaipán-tepui, Cerro Venamo), Amazonas (south of Sierra Parima, Cerro Sipapo). Venezuelan Coastal Cordillera, Sierra de Perijá; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. The taxonomy of this species should be considered tentative, as it is likely that more than one taxon is involved in this complex. Piper otto-huberi Steyerm., Fl. Venez. 2(2): 503, fig. 74. 1984. Shrub 1–2 m tall; leaves black-dotted on both surfaces; spikes 1.5 cm long, pendent. Riparian forests, ca. 100–200 m; Amazonas. Endemic; 2 varieties, 2 in Venezuela, both in the flora area. Key to the Varieties of P. otto-huberi 1. Leaf margins ciliate; upper surface of leaf less conspicuously black-dotted than lower surface ............. var. otto-huberi 1. Leaf margins eciliate; upper surface of leaf conspicuously black-dotted .............. ...................................... var. eciliatum P. otto-huberi var. eciliatum Steyerm., Fl. Venez. 2(2): 504. 1984. Amazonas (Río Autana, Ocamo, Río Orinoco, near San Fernando de Atabapo, Santa Bárbara del Orinoco). Endemic. ◆Fig. 611. P. otto-huberi var. otto-huberi Amazonas (Río Orinoco and Río Ventuari basins). Endemic.

Piper 723

Piper ovatum Vahl, Eclog. Amer. 1: 3, pl. 1. 1797. —Ottonia ovata (Vahl) Trel., Proc. Amer. Philos. Soc. 75: 711. 1935. —Anisillo, Raíz de candela, Raíz de muela. Subherbaceous, slender plant 0.5–1 m tall; fruits pedicellate, tetragonous. Semideciduous forests, 200–500 m; Bolívar (near Upata). Northeastern Venezuela; Trinidad. ◆Fig. 620. Mashed roots of Piper ovatum are used to poison fish and to alleviate toothache. Piper papilliferum Steyerm., Fl. Venez. 2(2): 507, fig. 74. 1984. Dwarf shrub 1.5 m tall, the stems with a subreflexed pubescence mixed with papillate protuberances; upper surface of leaves papillate-puberulent; ovary and fruit papillate. Riparian forests, 100–200 m; Amazonas (Piaroa, Río Guayapo, near San Simón de Cocuy). Endemic. ◆Fig. 634. Piper parapeltobryon Trel., Bull. Torrey Bot. Club 58: 356. 1931. Suffruticose plant 1–1.5 m tall; spikes 0.6–1.5 cm long; leaves gland-dotted, the bases equilateral peduncles 1.7–3 cm long, filiform. Wet forested tepui slopes, 1400– 2000 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Parú). Colombia. Piper peltatum L., Sp. Pl. 30. 1753, “pelatum.” —Peperomia peltata (L.) A. Dietr., Sp. Pl. 1: 142. 1831. —Lepianthes peltatum (L.) Raf., Sylva Tellur. 85. 1838. —Heckeria peltata (L.) Kunth, Linnaea 13: 565. 1839. —Pothomorphe peltata (L.) Miq., Comm. Phytogr. 37. 1840. Piper pruinosum H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 59. 1815 [1816]. Piper speciosum H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 59. 1815 [1816]. Herbaceous or soft-stemmed shrub 0.3–3 m tall; leaves membranous, 13–37 cm long, palmately 10–14-veined; spikes 3–12, on a 1.7–6.2 cm long common peduncle. Open or disturbed areas, roadsides, thickets, secondary or cut-over forests, and along streams, 50–200 m; Delta Amacuro (Caño Guiniquina), Bolívar (middle Río Caura), Amazonas (upper Río Cunucunuma). Widespread elsewhere in Venezuela; common from Mexico to southern South America. ◆Fig. 638.

Piper perciliatum Trel. & Yunck., Piperac. N. South Amer. 1: 326, fig. 293. 1950. Shrub 0.5–1.5 m tall with densely retrorsely pubescent stems; leaves reticulate, conspicuously ciliate; spikes 7–11 cm long pendulous or inclined. Evergreen lowland forests on laterite and white sandy soils, ca. 100–200 m; Amazonas (Río Guanía, near Maroa, San Carlos de Río Negro). Adjacent Colombia, Amazonian Brazil. ◆Fig. 641. Piper perstipulare Steyerm., Fl. Venez. 2(2): 519, fig. 78. 1984. Shrub 1.5–4 m tall with a conspicuous ligulate-lanceolate stipular development 5–7 cm long; leaves large, broadly elliptic to ovate-elliptic; spikes 7–9.5 cm long, erect to ascending,. Wet forests, 100–1300 m; southeastern Bolívar (Río Aponguao, Río Cuyuní basin, between El Dorado and Santa Elena de Uairén, between El Dorado and La Gran Sabana, Río Yuruani, Cerro Venamo), Amazonas (Sierra de la Neblina). Guyana. ◆Fig. 624. Piper piscatorum Trel. & Yunck., Piperac. N. South Amer. 1: 395, fig. 356. 1950. — Anisillo, Chawa pandres, Manchaua, Nudito, Raíz de candela, Raíz de mato, Raíz de muela. Strongly nodose, slender-stemmed subshrub; spikes with pilosulous rachis; fruits exserted, ovoid-tetragonous, densely papillate at apex. Wet forests, usually along streams, 100–400 m; Bolívar (middle Río Caura, Cerro Cuchivero, Río Cuyuní, between El Dorado and Santa Elena de Uairén, south of El Manteco, Río Guayapo, Salto Pará, Río Parguaza, La Unión), Amazonas (near Río Parucito). Endemic. ◆Fig. 640. The whole plant, broken into pieces, is used as a fish poison and to alleviate toothache. Piper politii Yunck., Mem. New York Bot. Gard. 9: 322. 1957. Climbing or creeping plant; leaves ovate, conspicuously reticulate-veined, the upper surface shining; spikes 2.3–3 cm long. 1300– 2000 m; Bolívar, Amazonas. Endemic; 3 subspecies, all in the flora area. The three known subspecies may eventually be relegated to synonymy as one variable species. More field investigations of this complex are needed.

724

P IPERACEAE

Key to the Subspecies of P. politii 1. Stems, petioles, apical stipule, and veins of lower surface of leaf abundantly pubescent .................. subsp. sipapoense 1. Stems, petioles, apical stipule, and veins of lower surface of leaf glabrous, or petiole and lower midrib sometimes sparsely pilosulous .............................................. 2 2. Leaf with principal lateral veins 3 or 4 on each side of midrib, pinnately veined to the upper 1/3; petioles and lower midrib sparsely pilosulous; leaf blades widest below the middle; tertiary venation more prominent on lower surface ....................................... subsp. politii 2. Leaf with principal lateral veins mainly 4–6 on each side of midrib, pinnately veined to the upper 1/4; petioles and lower midrib glabrous; leaf blades widest near the middle; tertiary venation more prominent on upper surface, obsolete on lower surface ............................... ................................ subsp. toronoense P. politii subsp. politii Tepui slope forests, 1300–1700 m; Bolívar (Cerro Guaiquinima), Amazonas (Cerro Yutajé). Endemic. P. politii subsp. sipapoense Steyerm., Fl. Venez. 2(2): 532. 1984. Steep tepui slopes in forest, 1500 m; Amazonas (Río Grande, Cerro Marahuaka, Cerro Sipapo). Endemic. ◆Fig. 639. P. politii subsp. toronoense Steyerm., Fl. Venez. 2(2): 530. 1984. Uppermost forested tepui slopes and summits on sandstone, 1900–2000 m; Bolívar (Macizo del Chimantá [Torono-tepui]). Endemic. Piper pseudoacreanum Steyerm., Fl. Venez. 2(2): 539, fig. 81. 1984. Scandent shrub with the principal leaf veins arising from lower 1/4–1/3 of blade; spikes 2.5 cm long in fruit, the fruits pubescent at base. Evergreen lowland forests, ca. 100 m; Amazonas (San Carlos de Río Negro, Santa María de los Guaicas). Endemic. ◆Fig. 623.

Piper pseudoglabrescens Trel. & Yunck., Piperac. N. South Amer. 1: 228, fig. 193. 1950. Glabrous shrub 1–3 m tall; leaves large, elliptic-oblong, equilaterally rounded, obtuse, or subcordate at base; spikes ascending to erect, 2–3 cm long with a conspicuous apical projection, the flowers arranged in spiral rows. Evergreen lowland to montane forests, along streams or on rocky slopes, 100–1300 m; Delta Amacuro (Jotajana), Bolívar (El Casabe, Guayapo, Represa Guri, Río Nichare, Salto Pará, Las Pavas, Serranía Pia-Zoi, near Santa Elena de Uairén), Amazonas (base of Cerro Duida, Sierra Parima, Platanal). Endemic. ◆Fig. 644. Piper pubivaginatum Steyerm., Fl. Venez. 2(2): 549, fig. 83. 1984. Suffruticose or small shrub 1–2 m tall; leaves pinnately veined the entire length; stems and stipules densely tomentose; spikes 2 cm long. Thickets and forests on white sand, ca. 100 m; Amazonas (near San Carlos de Río Negro). Endemic. Piper reticulatum L., Sp. Pl. 29. 1753. —Anisillo, Canilla de muerto. Piper smilacifolium H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 56. 1815 [1816]. Piper latum H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 57. 1815 [1816]. Tree 4–15 m tall or shrub 2–3 m tall, often with aerial roots at base of stem; leaves palmately 5–7(9)-veined, reticulately and transversely veined; fruits 4-sided, papillate-puberulent, with a truncate, glabrous apical disk. Evergreen or semideciduous forests, 100–300 m; Delta Amacuro (near El Palmar), Bolívar (near Salto Pará, Serranía de Imataca). Venezuelan Coastal Cordillera, occasionally in the Andes at higher altitudes; Central America, West Indies, Colombia, Ecuador, Peru, Brazil. ◆Fig. 628. Piper rupununianum Trel. & Yunck., Piperac. N. South Amer. 1: 303, fig. 268. 1950. Shrub 1–2 m tall; leaves 14–22.5 × 4–7 cm, narrowly lanceolate-elliptic to narrowly oblong-elliptic. Evergreen lowland forests,

Piper 725

ca. 100 m; Amazonas (Piedra Cocuy). Guyana, probably adjacent Brazil. Piper sabanaense Yunck., Bol. Soc. Venez. Ci. Nat. 23(103): 304. 1963. Suffruticose plant 1–1.5 m tall with petioles shorter than peduncles; leaves scabrous, subcordate, densely ciliate. Mauritia palm swamps, 700–900 m; Bolívar (Divina Pastora, San Ignacio de Yuruaní, near Santa Elena de Uairén). Endemic. ◆Fig. 637. Piper steyermarkii Yunck., Fieldiana, Bot. 28: 207. 1951. —Se-cun-wa-rei-yek (Arekuna). Climbing epiphytic shrub, completely glabrous; leaves long elliptic, acuminate; anthers dehiscing apparently by 3 fissures. Wet forests at base of sandstone bluffs, ca. 1600 m; Bolívar (Ptari-tepui). Endemic. ◆Fig. 642. Piper subalpinum Yunck., Fieldiana, Bot. 28(1): 208. 1951. Soft-stemmed shrub 1–2 m tall with densely retrorsely pubescent stems; leaves conspicuously rugose, equilaterally obtuse or rounded at base. Forested upper slopes near tepui summit, 2100–2300 m; Bolívar (Sororopán-tepui). Endemic. Piper subduidaense Yunck., Mem. New York Bot. Gard. 10: 42. 1963. Soft-stemmed shrub 1–1.5 m tall; leaves densely ciliate, inequilaterally-based, the lower surface glandular-dotted. Wet forested tepui slopes, 1600–1700 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 629. Piper tamayoanum Steyerm., Fl. Venez. 2(2): 572. 1984. Mauritia palm swamps, 900–1000 m; Bolívar (base of Cerro Ceitá near Santa Elena de Uairén). Endemic. ◆Fig. 643. Piper tenue H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 56. 1815 [1816]. Slender shrub or subshrub 1–2.5 m tall sparsely pubescent with lines of trichomes along angles of stems and petioles; leaves palmately 5-veined; fruits obtusely 4-sided, verruculose and glabrous except for the

sparsely papillate apex. Evergreen lowland and semideciduous forests, near sea level to 300 m; Delta Amacuro (Río Acure, Brazo Imataca, Sacupana), Bolívar (near El Dorado). Low elevations elsewhere in Venezuela, except absent from Coastal Cordillera and the Andes; Colombia. Piper tepuiense Steyerm., Fl. Venez. 2(2): 575, fig. 90. 1984. Subherbaceous plant 2 m tall, stems puberulent on upper internodes; leaves densely puberulent above and below; spikes bristlyappearing, prolonged to a slender apex. Forested slopes on sandstone at base of high bluff, ca. 1300–1400 m; Bolívar (Macizo del Chimantá [Amurí-tepui]). Endemic. Piper toronotepuiense Steyerm., Fl. Venez. 2(2): 576, fig. 90. 1984. —Sekúnwaráy. Herbaceous or subherbaceous plant, 1.5– 2.5 m tall; leaves 25 × 13–18 cm, suborbicular-ovate, long-acuminate, both sides puberulent; petioles and spikes pubescent. Forested slopes on sandstone at base of high bluffs, ca. 1700 m; Bolívar (Macizo del Chimantá [Toronó-tepui]). Endemic. Piper tortivenulosum Yunck., Bol. Inst. Bot. (São Paulo) 3: 128, fig. 112. 1966. Shrub 1.2–2 m tall; leaves elliptic-lanceolate or lanceolate, acuminate, acute or cuneiform at base, pinnately veined throughout the length with 8–11 principal veins on each side of midrib, and with tertiary venation on lower surface forming a twisted-irregular network. Evergreen lowland forests, ca. 100 m; Amazonas (near San Carlos de Río Negro). Brazil. Two forms have been recognized from the flora area: f. glabrum (Yunck.) Steyerm. with the lower surface of leaves completely glabrous and the typical form with the lower surface of leaf sparsely short-puberulent. Piper trigonum C. DC., J. Bot. 4: 212. 1866. Piper lucaeanum var. magnifolium C. DC. in A. DC., Prodr. 16(1): 322. 1869. Piper arieianum C. DC., Anales Inst. Fís.Geogr. Nac. Costa Rica 9: 166. 1897.

726

P IPERACEAE

Piper saltuum C. DC. in Urb., Symb. Antill. 7: 185. 1912. Piper acutissimum Trel. Contr. U.S. Natl. Herb. 26: 25. 1927. Piper duidaense Trel., Bull. Torrey Bot. Club 38: 355. 1931. —Piper arieianum var. duidaense (Trel.) Steyerm., Fl. Venez. 2(2): 344. 1984. Piper duidaense var. eciliatum Trel. & Yunck., Piperac. N. South Amer. 1: 401. 1950. Piper chimantanum Yunck., Mem. New York Bot. Gard. 9(3): 423. 1957. —Piper arieianum var. chimantanum (Yunck.) Steyerm., Fl. Venez. 2(2): 344. 1984. Mostly glabrous shrub; leaves ciliate-margined. Tepui slope forests, 1200–1700 m; Bolívar (Macizo del Chimantá), Amazonas (Cerro Duida). Venezuelan Coastal Cordillera; Nicaragua, Costa Rica, Panama, Trinidad, Colombia. Piper tuberculatum Jacq., Icon. Pl. Rar. 2(16): pl. 211. 1786–1793 [1795]. —Cordoncillo erecto. Piper tuberculatum var. minus C. DC. in A. DC., Prodr. 16(1): 266. 1869. Piper geniculatum f. puberula C. DC., Verh. Bot. Vereins Prov. Brandenburg 47: 106. 1905. Piper tuberculatum var. scandens Trel. & Yunck., Piperac. N. South Amer. 1: 367. 1950. Tree or shrub 1–5 m tall with stems often tuberculate or verrucose; petioles with tuberculate appendages usually present; leaves relatively small, obtuse or rounded at apex and strongly unequal at base. Thickets, roadsides, deciduous and gallery forests, near sea level to 300 m; Delta Amacuro (Capure, Tucupita), Bolívar (Altiplanicie de Nuria, middle Río Caura, El Dorado, Río Paragua). Widespread elsewhere in Venezuela; southern Mexico, Central America, West Indies, South America south to Peru, Brazil, and Bolivia. This is one of the most common species of Piper in Venezuela. Piper umbellatum L., Sp. Pl. 30. 1753. —Peperomia umbellata (L.) Kunth, Syn.

Pl. 1: 124. 1822. —Lepianthes umbellatum (L.) Raf., Sylva Tellur. 84. 1838. —Heckeria umbellata (L.) Kunth, Linnaea 13: 569. 1839. —Pothomorphe umbellata (L.) Miq., Comm. Phytogr. 36. 1840. Subherbaceous or soft-stemmed shrub 0.3–2 m tall; leaves membranous, 14–40 cm long, palmately 12–15-veined; 2–6 spikes on a 1–4 cm long common peduncle. Thickets and semideciduous forests, 200–300 m; Bolívar (near Tumeremo). Mainly Coastal Cordillera, scattered in Venezuela; common in Mexico, Central America, West Indies, South America. Piper venamoense Steyerm., Fl. Venez. 2(2): 589, fig. 92. 1984. Suffruticose, nearly glabrous plant 0.6 m tall; leaves pinnately-veined only to the middle with 4 veins on each side, ciliate-margined, and abundantly glandular-dotted on both sides. Wet montane forests, 1400–1500 m; Bolívar (Cerro Venamo basin near border of Guyana). Endemic. ◆Fig. 627. Piper vitaceum Yunck., Bol. Inst. Bot. (São Paulo) 3: 76, fig. 66. 1966. Venezuela, Brazil; 3 varieties, 1 in Venezuela. The other 2 varieties, var. vitaceum and var. airaoense Yunck., occur in Brazil. P. vitaceum var. venezuelense Steyerm., Fl. Venez. 2(2): 594, 1984. Sprawling or climbing plant; leaves pinnately veined in the lower third. Wet montane forests, ca. 1200 m; Bolívar (Río Cuyuní basin, Sierra de Lema, Salto La Danta). Endemic. Piper wachenheimii Trel., Bull. Misc. Inform. 1933: 339. 1933. Suffruticose plant 1.5 m tall; stems densely hirtellous with spreading trichomes; leaves conspicuously reticulate, scabrous, minutely hispidulous on lower surface; fruits densely hirtellous. Riparian lowland forests, ca. 300 m; Bolívar (Río Acarabisi). Guyana, Suriname, French Guiana, northeastern Brazil.

Piper 727

Fig. 607. Piper amalago

Fig. 609. Piper consanguineum

Fig. 608. Piper anonifolium var. anonifolium

Fig. 610. Piper demeraranum

728

P IPERACEAE

Fig. 611. Piper otto-huberi var. eciliatum

Fig. 612. Piper arboreum var. arboreum

Fig. 613. Piper avellanum

Piper 729

Fig. 614. Piper baccans var. sancarlosianum

Fig. 616. Piper bolivaranum

Fig. 615. Piper hippocrepiforme

Fig. 617. Piper cililimbum

730

P IPERACEAE

Fig. 618. Piper bartlingianum

Fig. 620. Piper ovatum

Fig. 619. Piper neblinanum

Piper 731

Fig. 621. Piper foveolatum

Fig. 622. Piper coruscans

Fig. 623. Piper pseudoacreanum

732

P IPERACEAE

Fig. 624. Piper perstipulare

Fig. 626. Piper jauaense

Fig. 625. Piper cumanense

Fig. 627. Piper venamoense

Piper 733

Fig. 628. Piper reticulatum

Fig. 630. Piper guianense

Fig. 629. Piper subduidaense

734

P IPERACEAE

Fig. 631. Piper hostmannianum

Fig. 633. Piper kegelianum

Fig. 632. Piper marginatum var. marginatum

Piper 735

Fig. 634. Piper papilliferum

Fig. 636. Piper lemaense

Fig. 635. Piper morilloi

Fig. 637. Piper sabanaense

736

P IPERACEAE

Fig. 638. Piper peltatum

Piper 737

Fig. 639. Piper politi var. sipapoense

Fig. 641. Piper perciliatum

Fig. 640. Piper piscatorum

738

P IPERACEAE

Fig. 642. Piper steyermarkii

Fig. 643. Piper tamayoanum

Fig. 644. Piper pseudoglabrescens

Plantago 739

PLANTAGINACEAE by Gladys Rodríguez Herbs, annual or perennial, acaulescent or rarely subfruticose, with rhizome or stout rootstock. Leaves alternate or opposite, simple, generally linear or ovate-elliptic, sometimes reduced, generally forming a basal rosette, exstipulate, blade and petiole often not differentiated; margins entire to serrate-dentate; venation appearing parallel from the base of petiole. Inflorescence of solitary or axillary heads or spikes. Flowers actinomorphic, bisexual, rarely unisexual (plants monoecious or dioecious); bracts similar to sepals. Sepals 4, free or nearly so, imbricate, ridged, with scarious margins. Corolla greenish, sympetalous, 4-lobed, salverform, the lobes imbricate, scarious, spreading or reflexed. Stamens (1–)4, inserted in corolla tube or hypogynous, alternate with the lobes; anthers 2-locular, cordate to ovate, dehiscing longitudinally from the base, versatile. Ovary superior, 1–4-locular, sessile; ovules 2–many, semianatropous, placentation axile or basal; style solitary; stigma dry, elongate, sometimes pubescent. Fruit a membranous circumscissile capsule, or an indehiscent achene or bony nutlet, enclosed in the persistent calyx. Seeds 2–many, peltately attached, sometimes mucilaginous; embryo straight or ± curved; endosperm fleshy. Cosmopolitan (principally in the temperate zones); 3 genera and ca. 260 species, 1 species in the flora area. 1. PLANTAGO L., Sp. Pl. 112. 1753. Psyllium Tourn. ex A.L. Juss., Gen. Pl. 90. 1789. Plantaginella Fourn., Ann. Soc. Linn. Lyon sér. 2, 18: 140. 1869. Lagopus Fourn., Ann. Soc. Linn. Lyon sér. 2, 18: 140. 1869. Distribution as in family; ca. 250 species, 5 in Venezuela, 1 of these in the flora area. Plantago major L., Sp. Pl. 112. 1753. —Llantén. Herb 10–50 cm tall; blade and petiole ± differentiated; blades ovate-elliptic; corolla tube glabrate, the lobes ca. 1 mm long, reflexed, apex generally acute. Generally in disturbed areas, sometimes cultivated, ca. 500–1200 m; Delta Amacuro (Tucupita, cultivated), Bolívar (2 km east of El Paují, near Upata). Widespread elsewhere in Venezuela; cosmopolitan. ◆Fig. 645. Plantago major is sometimes cultivated as a medicinal plant and used as a diuretic and to treat blindness, fever, gastric and duodenal ulcers, inflammations, and cancer.

Fig. 645. Plantago major

740

P LUMBAGINACEAE

PLUMBAGINACEAE by Carlos A. Vargas Shrubs, lianas, or herbs. Leaves alternate, spirally arranged or absent, basal or cauline, elliptic, ovate, lanceolate, rarely subulate, membranous or leathery, simple, entire, petiolate, usually with characteristic scattered chalk-glands at or depressed below the surface, these exuding water and usually calcium salts, and also with raised mucilage glands in the axils and on the proximal part of the upper leaf surface, exstipulate. Inflorescences racemose, cymose, often spike-like, terminal or axillary, bracteate; bracts dry and membranous (sometimes forming on involucre), floral nodes bracteolate, bracteoles 2(1). Flowers actinomorphic, 5-merous, often heterostylous. Calyx persistent, forming a distinct, often conspicuously 5- or 10ribbed tube with membranous intervals, the lobes mostly dry and membranous or scarious, often showy and somewhat petaloid. Corolla sympetalous or seldom of essentially distinct, clawed petals, often persistent, the lobes convolute, imbricate, or contorted, rarely leathery (Aegialitis). Stamens 5, isomerous, opposite the corolla lobes; filaments free; anthers dorsifixed or basifixed. Ovary superior, 5-carpellate, 1locular with a single, apically lobed style; stigmas dry, papillate; ovule solitary and basal, on an elongate, slender funiculus. Fruit partially or totally enclosed by the persistent calyx, most commonly a dry achene, sometimes a tardily circumscissile capsule, or the capsule upwardly dehiscent by valves. Seed winged; endosperm copious or rarely absent; embryo straight. Cosmopolitan; ca. 730 species and 27 genera, 1 species in the flora area. The center of diversity and endemism is found in cold and arid montane habitats of central Asia, and many of the species occupy xerophytic or saline habitats. PLUMBAGO L., Sp. Pl. 151. 1753. Perennial herbs, subshrubs, or shrubs. Stems ribbed, often elongate and climbing. Leaves cauline, membranaceous, pinnately veined, tertiary venation highly reticulate and anastomosing, sometimes clasping and auriculate at the base. Inflorescence a spike or raceme, with flowers shortly pedicellate, pedicels bibracteolate. Flowers distylous. Calyx tubular, covered with sessile or stalked glands along the 5 ribs, lobes triangular, sinuses hyaline; corolla salverform, glabrous, long-exserted, aestivation contorted, lobes obovate, rounded or truncate, often mucronate. Stamens free from corolla, included or exserted. Style 1, included or exserted. Fruit a membranaceous, circumscissile capsule at the base, the caducous part often splitting toward the apex by 5 valves, sometimes the apex of fruit covered by perianth remnants, long-beaked by the persistent style base. Seeds oblong, funiculate, reddish to dark brown. Mostly pantropics; 24 species, 4 in Venezuela, 1 of these in the flora area. Plumbago zeylanica L., Sp. Pl. 151. 1753. Plumbago scandens L., Sp. Pl. ed. 2, 215. 1762. Suffrutescent herb, somewhat climbing, 0.5–1.5 m tall; leaves obviously petiolate, with

the base ± clasping, but not auriculate, 3–15 × 1–6 cm, leaves and stems appearing lepidote due to salt deposits; corolla white, 17–30 mm long, the lobes rounded and conspicuously mucronate; calyx sticky, stipitate-glandular

Plumbago 741

over entire length. Disturbed areas, shrublands, open sites, near sea level to 50 m; Delta Amacuro (Sacupana), Bolívar (Ciudad

Bolívar). Distrito Federal, Nueva Esparta, Zulia; widespread introduction in the New World, native to the Old World. ◆Fig. 646.

Fig. 646. Plumbago zeylanica

Appendix List of new names published in this volume

Blastemanthus gemmiflorus subsp. sprucei (Tiegh.) Sastre, comb. nov. . . . . . . . Caularthron bilamellatum f. indivisa (Bradf. ex Griseb.) Carnevali & I. Ramírez, comb. & stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Otoglossum scansor (Rchb.f.) Carnevali & I. Ramirez, comb. nov. . . . . . . . . . . . Piper julianii Callejas, nom. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prosthechea crassilabia (Poepp. & Endl.) Carnevali & I. Ramírez, comb. nov. . .

742

127 273 488 721 538

Index Compiled by George Thornburgh

Entries in Roman type = accepted names of taxa and vernacular names Entries in italics = synonyms Page numbers in bold = illustrations

—A— Acacallis, 229 cyanea, 230 fimbriata, 230, 231 Aceituna, 199 Aceituno, 199 Acianthera, 503 fockei, 515 garciae, 515 lanceana, 517 omissa, 519 prolifera, 519 wilsonii, 522 yauaperyensis, 522 Acineta, 230 alticola, 231, 231 Acoidium, 592 fuscum, 594 Acraea, 543 Acrocarpidium majus, 698 tenellum, 697 urocarpum, 698 Acronia, 503 Adenanthe, 125 bicarpellata, 125, 126 Adenarake, 125 macrocarpa, 126 muriculata, 126, 127 Adeneleutherophora, 311 graminifolia, 313

Adipe, 239 longicornis, 240 Aeranthus pachyrrhizus, 258 Aganisia, 232 alba, 491 brachypoda, 232 fimbriata, 230 lepida, 491 pulchella, 232, 233 venusta, 491 Agonandra, 198 brasiliensis, 198 subsp. brasiliensis, 199, 199 subsp. racemigera, 199 silvatica, 199 Agracia ácido, 81 Ahorca vena’o, 655 Ai-yek, 65 Ají de paloma, 135 Ajicito, 138 Ajonjolí, 668 Albajaca, 674 Amalago, 705 Amalia, 561 Amphyglottis, 325 Anacheilium, 536 Anamomis, 79 Anathallis, 503 barbulata, 513 brevipes, 513

743

maguirei, 518 sclerophylla, 521 Anaucé, 674 Ancistrocarpus, 674 maypurensis, 674 Ancistrothyrsus, 627 hirtellus, 627, 628 Angraecum fasciola, 257 lansbergii, 258 micranthum, 258 poeppigii, 258 Angreacum maculatum, 476 Anicillo manso, 715 Anisillo, 716, 718, 721, 723, 724 Anthereon, 503 mentosus, 518 Apatostelis, 579 alata, 583 garayi, 585 minimiflora, 586 Aptandra, 165, 166 benthamiana, 165 liriosmoides, 165 spruceana, 166 tubicina, 165 Aracamunia, 232 liesneri, 233, 234 Arepillo, 115 Arichai, 22

744

I NDEX

Aroi-waray-yek, 129 Artanthe, 705 adenandra, 714 aduncum, 714 amazonica, 718 anonifolia, 715 asperifolia, 720 augusta, 716 avellana, 716 baccans, 716 bartlingianum, 717 caripensis, 714 caudata, 721 consanguinea, 717 demerarana, 718 eximia, 717 flagellaris, 722 guianensis, 719 hirsuta, 720 hostmanniana, 720 javitensis, 721 kegeliana, 721 luschnathiana, 716 marginata, 721 obrumbata, 716 parkeriana, 715 pseudochurumayu, 718 ramiflora, 720 spectabilis, 717 superba, 722 Ashudu, 692 Aspasia, 234 variegata, 234, 235 Aspegrenia scirpoidea, 472 Aspidogyne, 234 confusa, 236 pumila, 236 robusta, 236 steyermarkii, 235, 236 Asta blanca, 168 Auliza, 325 Aulomyrcia, 54 albido-tomentosa, 59 amshoffiana, 59 bolivarensis, 59 chilensis, 59 citrifolia, 60 dumosa, 64 edulis, 64 hostmanniana, 59 inaequiloba var. paniculata, 62

karuaiensis, 44 obtusa, 62 paniculata, 62 parvifolia, 65 platyclada, 64 var. kaieteurensis, 64 ptariensis, 64 roraimae, 62 roraimensis, 62 rotundata, 65 salticola, 65 spruceana, 59 tomentosa, 67 triflora, 60 vinacea, 64 Aurosá, 677 —B— Badea, 653 Barbosella, 236 kegelii, 516 orbicularis, 237, 237 Baskervilla, 237 colombiana, 237, 238 Batemannia, 237 colleyi, 239 lepida, 238, 239 Beadlea, 291, 385 bifida, 386 cranichoides, 291 elata, 292 Bejuco ciruelo, 115 Bejuco parcha, 650, 655 Beloglottis, 385 americana, 385 bifida, 386 chiropterae, 386 sect. Helonoma, 385 Bifrenaria, 239, 392, 549 aurantiaca, 550 longicornis, 240, 240 maguirei, 372 minuta, 393 rudolfii, 393 steyermarkii, 241 venezuelana, 240, 241 wageneri, 592 Biophytum, 620 calophyllum, 621, 621 cardonaei, 624 casiquiarense, 621 passargei, 621

sp. A, 622, 624 sp. B, 623, 624 sp. C, 623, 624 Blastemanthus, 127 densiflorus, 127 gemmiflorus, 127 subsp. gemmiflorus, 127 subsp. sprucei, 127, 127 paniculatus, 127 sprucei, 127 Blepharocalyx, 4 eggersii, 4, 4 Bletia, 241 stenophylla, 241, 242 Boerhavia, 102 caribaea, 102 coccinea, 102, 103 diffusa, 102 paniculata, 102 Bolbophyllaria, 253 oerstedii, 255 Bollea, 241 cardonae, 243 hemixantha, 242, 243 Bonatea pauciflora, 382 pratensis, 381 Borax, 308 Borrajón, 716 Bougainvillea, 104 buttiana, 104 spectabilis, 103, 104 Brachionidium, 243 brevicaudatum, 244, 244 julianii, 244, 245 longicaudatum, 244, 245 neblinense, 245, 245 parvum, 245, 245 sect. Yolanda, 243 Brachynema, 166 axillare, 167, 167 Brachystele, 246 guayanensis, 246, 246 Braemia, 247 vittata, 247, 247 Brassavola, 248 angustata, 248 cucullata, 248, 249 martiana, 248, 249 Brassia angusta, 251 bidens, 251, 252 caudata, 251, 252

I NDEX

forgetiana, 251, 252 lanceana, 251, 252 lawrenceana, 251, 252 macrostachya, 251 neglecta, 251, 252 Buenas tardes, 110 Bulbophyllum, 253 correae, 255 dunstervillei, 254, 255 exaltatum, 254, 256 geraense, 254 meridense, 254 meristorhachis, 254 oerstedii, 255 pachyrrhachis, 255 roraimense, 255, 255 setigerum, 255, 256 Burro muerto, 172 —C— Cafecito, 177 Callejas, 721 Calpidia, 117 Calycampe angustifolia, 60 latifolia, 60 Calycolpus, 5 alternifolius, 6, 8 bolivarensis, 6 calophyllus, 6, 6 var. angustifolius, 6 goetheanus, 7, 8 roraimense, 8 roraimensis, 7, 8 Calycorectes, 8 enormis, 9, 9 macrocalyx, 26 yatuae, 28 Calyptranthes pullei var. immaculata, 13 Calyptranthes, 10 clusiifolia, 11 conduplicata, 11 crebra, 11 fasciculata, 11, 13 florifera, 12 forsteri, 11, 14 lucida, 12 macrophylla, 12, 15 multiflora, 12 nigrescens, 12, 14

obtusa, 45 pulchella, 12, 15 pullei, 13, 14 ruiziana, 13 sericea, 11 Camaridium, 426 album, 434 amazonicum, 435 equitans, 437 iguapensis, 438 imbricatum, 438 ochroleucum, 435 pendulum, 442 squamatum, 444 uncatum, 445 vandiforme, 437 Cámero, 653 Campomanesia, 16 aromatica, 16, 17 grandiflora, 16 Camptouratea leblondi, 138 sagotii, 138 Campylocentrum, 256 fasciola, 257, 259 hondurense, 257, 259 huebneri, 258 lansbergii, 258 micranthum, 258 nataliae, 258, 259 pachyrrhizum, 258, 259 panamense, 258 poeppigii, 258 steyermarkii, 257 uroplectron, 258 Caneya gruya, 716 Canilla de grulla, 717, 721 Canilla de muerto, 724 Canjilón guayabero, 16 Carurú, 677 Casabe, 106, 107, 114 Casabe marrón, 106 Casabe morado, 107 Casabito, 106 Castalia blanda, 120 Catasetum, 260 apertum, 265 barbatum, 262, 267 bergoldianum, 262, 268 bicallosum, 262, 264, 268 bicolor, 262, 268

745

callosum, 263, 268 chloranthum, 264 collare, 263, 267 comosum, 262 costatum, 263 cristatum, 263, 267 discolor, 263, 265 ×dunstervillei, 263, 266 ferox, 263, 268 gomezii, 263, 269 gongoroides, 262 longifolium, 263, 266 macrocarpum, 264, 267 maroaënse, 264 merchae, 264, 268 microglossum, 264, 267 parguazense, 264, 268 pileatum, 264, 268 planiceps, 264, 267 recurvatum, 264 rivularium, 264, 268 ×roseo-album, 264, 266 scurra, 281 splendens, 265 stenoglossum, 262 ×tapiriceps, 265, 267 warczewiczii, 281 ×wendlingeri, 265, 267 yavitaënse, 265, 268 Cathedra, 167 acuminata, 168, 168 caurensis, 168 crassifolia, 168 Cattleya, 269 jenmanii, 270, 270 lawrenceana, 270, 271 liliastrum, 572 superba, 270 violacea, 270, 271, 574 Caularthron, 271 bicornutum, 272, 272 bilamellatum, 272 Cazabe, 106, 107 Cazabito, 107 Centrogenium, 316 calcaratum, 316 setaceum, 316 Centroglossa peruviana, 290 Ceratachilus grandiflorus, 579 Ceratoccoca maypurensis, 674

746

I NDEX

Ceratochilus, 577 Cercouratea chaffanjonii, 135 ferruginea, 136 grosourdyi, 137 laxa, 141 Cespedesia, 128 bonplandii, 129 spathulata, 128, 129 sprucei, 129 Chaenanthe, 298 barkeri, 298 Chamelophyton, 503 kegelii, 516 Chãmore, 656 Chamuré, 645 Chaubardia, 273 surinamensis, 274 tigrina, 275 Chaubardiella, 274 tigrina, 275, 275 Chaunochiton, 169 angustifolium, 169, 170 loranthoides, 169, 170 mouriroides, 166 Chawa pandres, 723 Cheiradenia, 275 cuspidata, 276, 276 imthurnii, 276 Chelyorchis, 276 ampliata, 277, 277 Chionanthus, 186 caribea, 187 compactus, 187, 187 implicatus, 187 Chipachi, 677 Chipo-cuy-yek, 59 Chipoki, 60 Chipo-yek, 64 Chunchunillo blanco, 62 Chytraculia, 10 Cinco llagas, 652 Cladobium, 555 violaceum, 558 Clavo de pozo, 188 Cleistes, 278 huberi, 279, 280 lepida, 279 moritzii, 279 parviflora, 279 rosea, 279, 280 stricta, 279

tenuis, 279 triflora, 279, 280 unifoliata, 280 Clowesia, 281 warczewiczii, 281, 281 Cochleanthes, 282 flabelliformis, 282, 282 Cohniella, 592 cebolleta, 594 Comparettia, 283 falcata, 283, 284 Congrio, 109 Copeycito, 140 Copito negro, 170 Cordoncillo, 716, 720, 721 Cordoncillo erecto, 726 Cordoncillo negro, 721 Corona de rey, 177 Coryanthes, 284 albertinae, 285 cataniapoënsis, 285, 285 feildingii, 285 gomezii, 285, 285 macrantha, 285, 286 maculata, 287 pegiae, 286, 287 rutkisii, 286, 287 Cranichis, 287 alfredi, 287 diphylla, 287, 288 guatemalensis, 287 monophylla, 287 nigrescens, 287 oligantha, 535 ovatilabia, 287 stachyodes, 535 stictophylla, 287 Craniolaria, 668 annua, 668, 669 Crocodeilanthe, 503 Cryptarrhena, 288 kegelii, 289, 289 Cryptocentrum, 289 dunstervilleorum, 290, 290 gracillimum, 290 hoppii, 291 peruvianum, 290 subsp. peruvianum, 291 Cryptophoranthus, 503 pleurothallopsis, 482

Cuculina, 260 Cuentica roja, 677 Cumello, 115 Cumello banero, 115 Cupi fino, 172 Cupi hoja fina, 115 Curame, 60, 63 Curamita, 115 Curamito, 115 Curtido, 59 Curtidor, 59 Cuyuvi, 181 Cyathoglottis candida, 571 Cybelion, 394 Cyclopogon, 291, 385 bifidus, 386 cranichoides, 291, 292 elatus, 291 Cyclosia, 457 Cycnauken, 293 Cycnoches, 293 chlorochilon, 294 cucullata, 295 haagii, 294, 294 loddigesii, 295 lusiae, 295, 295 pescatorei, 416 thurstonorum, 294, 295 ventricosum var. chlorochilon, 294 versicolor, 294 Cymbidium andersonii, 297 cordigerum, 320 diurnum, 320 ochroleucum, 435 violaceum, 270 Cypripedium, 564 klotzschianum, 499 lindleyanum, 499 palmifolium, 566 Cyrtopodium, 296 andersonii, 297 broadwayi, 297 cristatum, 297, 297 eburnea, 401 flavescens, 297 fowliei, 297 graniticum, 297 parviflorum, 297 Cystochilum, 287

I NDEX

—D— Dau dau, 45 Decaisnea, 533 Dendrobium album, 434 cebolleta, 594 Diacrium, 271 amazonicum, 272 bicornutum, 272 bilamellatum, 272 venezuelanum, 272 Diadenium, 298 barkeri, 298, 298 Dialissa, 579 Dichaea, 299 ancoraelabia, 301 brachypoda, 301 camaridioides, 301 echinocarpa, 303 histrio, 304 hookeri, 302 hystricina, 302, 304 kegelii, 302, 303 latifolia, 302 var. longa, 302 longa, 302 morrisii, 302, 303 muricata, 302, 303 panamensis, 302 pendula, 303 var. swartzii, 303 picta, 303 rendlei var. rendlei, 305 var. trinitensis, 305 robusta, 303 splitgerberi, 303 swartzii, 303 tachirensis, 304 trinitensis, 304 trulla, 305 venezuelensis, 304, 305 Dichaeopsis, 299 Dicrypta, 426 bicolor, 437 discolor, 437 longifolia, 444 villosa, 445 Didactyle, 253 exaltata, 254 meridensis, 254

Dikylikostigma, 307 preusii, 308 Dilkea, 628 acuminata, 629, 629 johannesii var. parvifolia, 629 magnifica, 629 parviflora, 629 Dimerandra, 305 elegans, 306, 306 emarginata, 306 Diothonea imbricata, 387 Diplocrater acuminatu, 168 Dipteranthus, 307 peruvianus, 465 planifolius, 307, 307 Discyphus, 307 scopulariae, 308, 308 Don diego de noche, 110 Dryadella, 309, 309 lueriana, 309 Duboisia, 459 reymondii, 460 Duckeella, 309 adolphii, 311 alticola, 310, 310 pauciflora, 310, 311 Dukatu, 113 Dulacia, 170 adhaerens var. stenopoda, 172 candida, 172, 173 crassa, 172 cyanocarpa, 172 guianensis, 172, 174 inopiflora, 172, 173 macrophylla, 172, 173 redmondii, 172, 174 tepuiensis, 172, 174 Dumatsi, 63 Dunstervillea, 311 mirabilis, 311, 311 —E— Echinella aspasicensis, 513 lappiformis, 517 Echinosepala, 503 aspasicensis, 513

747

lappiformis, 517 Educadu, 114 Ekedek, 114 Elleanthus, 311 arpophyllostachys, 313 attenuatus, 313 caravata, 313 columnaris, 313 confusus, 313, 314 gracilis, 313 graminifolius, 313 malpighiiflorus, 315 norae, 315 puber, 492 pusillus, 313 sphaerocephalus, 314, 315 wageneri, 314, 315 Eloyella, 315 panamensis, 316, 316 Eltroplectris, 316 calcarata, 316, 317 Elvasia, 129 brevipedicellata, 129 canescens, 129, 130 caurensis, 130 elvasioides, 130, 130 hostmannia, 130 quinqueloba, 130, 130 sagoti, 130 steyermarkii, 146 Enckea, 705 dubia, 719 Encyclia, 317, 536 acuta, 320 amicta, 324 amicta, 320 auyantepuiensis, 319, 323 chloroleuca, 320 conchaechila, 320, 322 cordigera, 320 crassilabia, 538 diurna, 320 fragans subsp. aemula, 538 granitica, 321, 323 guianensis, 321 ivonae, 321 latipetala, 324 leucantha, 321, 322 oncidioides, 321 pachyantha, 321, 324 pamplonense, 540

748

I NDEX

[Encyclia] picta, 321 pilosa, 324, 322 pygmaea, 539 recurvata, 321 remotiflora, 324 tigrina, 539 vespa, 540 wageneri, 320 Epidendropsis, 325 vincentina, 345 violascens, 345 Epidendrum, 317, 325, 536 aemulum, 538 agathosmicum, 333 alsum, 333 althausenii, 333 amazonicoriifolium, 334, 349 amazonicum, 336 amictum, 321 anceps, 334 apuahuense, 334 atropurpureum, 320 attenuatum, 334 bathyschistum, 336 bicornutum, 272 bilamellatum, 272 calanthum, 334, 346 carnosum, 337 carpophorum, 335, 350 carthagenense, 594 caudatum, 251 caurense, 335 chimantense, 335 chloranthum, 320 chloroleucum, 320 churunense, 335 ciliare, 335 var. squamatum, 335 clavatum, 341 commelinispathum, 336 compressum, 336 conchaechilum, 320 concretum, 532 congestoides, 336 cordigerum, 320 coriifolium, 334 coronatum, 336 crassifolium, 342 crassilabium, 538 cristatum, 336, 348 cucullatum, 248

dendrobioides, 344 densiflorum, 337 sect. Diacrium, 271 dichaeoides, 337, 347 dichotomum, 342 diurnum, 319, 320, 321, 324 duckei, 337 durum, 333, 337, 344 echinocarpum, 303 ellipticum, 342 elongatum, 342 ernstii, 338 flabelliformis, 282 flexuosum, 337 fragans, 538 var. aemulum, 538 frigidum var. stenophyton, 338 globosum, 397 graniticum, 321 halatum, 321 hombersleyi, 337, 351 huebneri, 340, 341 ibaguense, 338, 352 var. schomburgkii, 339 var. confluens, 334 ibaguense × secundum, 338, 352 imatophyllum, 337 imbricatum, 415 imitans, 334 imthurnii, 338 incisum, 339 inconstans, 342 klotzscheanum, 338, 348 labiatum, 402 labiosum, 618 latifolium, 335 latipetalum, 321 laxum, 336 lechleri, 335 leucanthum, 321 leucochilum, 339 lindenii, 342 lineare, 396 longicolle, 339 macrocarpum, 339 magnicallosum, 339 micronocturnum, 339 microphyllum, 339, 346 miersii, 342 miserrimum, 340, 346

montigenum, 340 moyobambae, 336 myrmecophorum, 340, 351 nocturnum, 340 var. latifolium, 335 var. tumuc-humacience, 344 norae, 340 nuriense, 341 oncidioides, 319 ophioglossoides, 586 orchidiflorum, 341, 351 ottonis, 462 pabstii, 337 pachyanthum, 321 pachyphyton, 341 palmarum, 612 pamplonense, 540 paniculatum, 337 pilosum, 324 prostratum, 341 pseudoramosum, 341 purpurascens, 341, 347 pusillum, 360 pygmaeum, 539 ramosum, 341 rectopedunculatum, 340 f. denticulatum, 341 remotiflorum, 324 repens, 342 rigidum, 342 ruscifolium, 520 satyrioides, 394 saxatile, 342 schlechterianum, 336 schlimii, 342 schomburgkii, 339 var. confluens, 334 sculptum, 342 secundum, 334, 342, 352 selligerum, 321 serricardium, 345 sertorum, 343 smaragdinum, 343, 351 spilotum, 340 splendens, 343 stalkyi, 343, 346 stamfordianum, 343 strobiliferum, 344 strobiloides, 344 teretifolium, 398 tigrinum, 539

I NDEX

trichorhizum, 615 trigoniflorum, 586 tumuc-humaciense, 344 ulei, 344, 349 unguiculatum, 344 uniflorum, 333 urbanianum, 345 urichianum, 345, 352 utricularioides, 395 variegatum, 540 vespa, 540 vincentinum, 345 violascens, 345, 350 wageneri, 320 weddellii, 342 xanthinum, 342 yatapuense, 336 sp. A, 345 Epistephium, 353 cruegeri, 355 duckei, 354, 355 elatum, 354 ellipticum, 354 hernandii, 354 parviflorum, 354, 356 petiolatum, 354 sclerophyllum, 355 subrepens, 355, 356 Epithecia, 299 Eriopsis, 357 biloba, 358, 358 colombiana, 358 grandibulbosa, 358 rutidobulbon, 358 sceptrum, 358 sprucei, 358 Erycina, 359 glossomystax, 360 pusilla, 360, 360 Erythrodes, 234, 360, 409 arietina, 361, 361 confusa, 236 paleacea, 361 pumilus, 236 robusta, 236 santensis, 410 steyermarkii, 236 stigmatoptera, 410 Escobil, 674 Eugenia, 17 acetosans, 60 amblyosepala, 22, 29 anastomosans, 22, 29

baileyi, 22, 30 benthami, 27 biflora, 22, 32 bracteata, 59 cachoeirensis, 22, 33 callichroma, 23 caurensis, 22 chrysophyllum, 23, 31 citrifolia, 23 coffeifolia, 23, 32 conduplicata, 23 congestissima, 24 cribrata, 23, 34 cuaoensis, 24 cumini, 96 deflexa, 61 diplocampta, 24 dittocrepis, 24, 33 egensis, 24, 34 emarginata, 24, 35 fallax, 61 feijoi, 24, 36 ferreiraeana, 24 ferruginea, 44 flavescens, 24 var. guianensis, 24 floribunda, 82 florida, 25, 32 gomesiana, 25 guianensis, 62 inaequiloba, 62 jambos, 98 kaieteurensis, 25 kunthiana, 27 lambertiana, 25 var. lambertiana, 25, 35 ligustrina, 25 var. ligustrina, 25, 30 lingua, 22 llewelynii, 23 longiracemosa, 25 macrocalyx, 25 magna, 26 malaccensis, 98 marlierioides, 26 melinonis, 23 montana, 45 monticola, 26, 31 var. latifolia, 26 var. racemosa, 26 moritziana, 26 multiflora, 63 nitida, 62

749

ochra, 27 omissa, 26 ottonis, 27 pachystachya, 26, 37 patrisii, 26, 37 peregrina, 22 producta, 28 prosoneura, 25 protenta, 27, 40 protracta, 27, 82 pseudopsidium, 27 ptariensis, 22 pubescens, 27, 38 punicifolia, 27, 40 pyrifolia, 64 quitarensis, 65 ramiflora var. montana, 24 rivularis, 86 rondonensis, 79 roriamana, 28 roseiflora, 27, 38 schomburgkii, 25 steyermarkii, 79 subalterna, 27 tapacumensis, 27, 36 tenella, 84 tepuiensis, 28 tomentosa, 67 trinervia, 28 tumulescens, 28 turumiquirensis, 79 umbonata, 28, 38 vismeaefolia, 84 wentii, 26 yatuae, 28, 39 sp. A, 28 sp. B, 28, 31 Eulophia, 362 alta, 362, 362 Eulophidium, 476 maculatum, 476 Eurystyles, 363 cotyledon, 363, 363 Evelyna, 311 arpophyllostachys, 313 columnaris, 313 gracilis, 313 wageneri, 315 —F— Farasu, 62, 64

750

I NDEX

Fernandezia, 299 acuta, 414 graminifolia, 558 Fournieria, 128 Fractiunguis, 545 —G— Galeandra badia, 365, 366 beyrichii, 365 carnevaliana, 365 devoniana, 365, 366 duidensis, 365, 366 juncea, 367 lacustris, 365 macroplectra, 365, 366 magnicolumna, 365 minax, 365, 366 stangeana, 367 stylomisantha, 366, 367 Galeoglossum, 533 Galeottia, 367 burkei, 368, 368 jorisiana, 368, 368 Garayella hexandra, 516 Godoya gemmiflora, 127 spathulata, 129 Gomidesia, 54 bonnetiasylvestris, 59 lindeniana, 62 Gomphia aquatica, 134 castaneaefolia, 135 grandiflora, 137 polyantha, 139 Gomphichis, 369 adnata, 369 costaricensis, 369, 370 Gongora, 370 atropurpurea, 370, 371 macrantha, 285 pleiochroma, 371 Goodyera guayanensis, 246 Gregarito, 106 Gregorito, 107 Guanchezia, 371 maguirei, 372, 372 Guapira, 104 amacurensis, 106

ayacuchae, 106 bolivarensis, 106, 108 cuspidata, 106, 108 davidsei, 106 ferruginea, 106, 109 fragrans, 107 glabriflora, 107 guianensis, 107 marcano-bertii, 107 microphylla, 107 neblinensis, 107 rusbyana, 109 sancarlosiana, 109 sipapoana, 109 Guayabillo, 60, 82 Guayabillo blanco, 82 Guayabita, 92 Guayabito, 6, 63, 138 Guayabito de agua, 91 Guayabo, 91 dantero, 26 Guayabo blanco, 91 Guayabo de caño danero, 90 Guayabo montañero, 82 Guayabo rajado, 92 Guayabo sabanero, 91 Guayabo tortugero, 91 Guayuyo, 721 Guayuyu montañera, 716 Gyas, 241 —H— Habenaria, 372 amalfitana, 380 amambayensis, 376 arecunarum, 380 armata, 376 ayangannensis, 377 caldensis, 377, 378, 379 culmiformis, 379 demerarensis, 380 distans, 377 dusenii, 377 enthomantha, 376 ernestii, 377, 378 floribunda, 378 georgii, 381 guanchezii, 381 heptadactyla, 378, 384 huberi, 378, 384 leaoana, 382 lehmanniana, 379

leprieur var. heptadactyla, 378 leprieuri, 378, 379 longicauda, 379 maculosa, 380 mesodactyla, 379 monorrhiza, 380 moritzii, 376 nilssonii, 380 obtusa, 380 parviflora, 380 pauciflora, 382 petalodes, 378 pratensis, 381 var. parviflora, 381 pygmaea, 381, 383 quadrata, 381 quinqueseta, 381 repens, 381, 383 rodeiensis, 377, 378 roraimensis, 382 sartor, 379 schomburgkii, 382 setacea, 382 var. setacea, 382 speciosa, 380 sprucei, 382 trifida, 382 viridiaurea, 378 sp. A, 382 sp. B, 383 Ha-dá-sa, 64 Heckeria, 705 peltata, 723 umbellata, 726 Heimerlia, 117 Heimerliodendron, 117 Heisteria, 175 acuminata, 177, 178 acuta, 181 barbata, 177, 179 densifrons, 177 duckei, 177, 180 fatoensis, 181 flexuosa, 177 flexuosa, 181 guianensis, 177 insculpta, 177, 178 longipes, 177 maguirei, 177 maytenoides, 177, 180 microcarpa, 177 nitida, 177

I NDEX

ovata, 177, 180 pentandra, 177 scandens, 181 spruceana, 179, 181 tubicina, 166 Helonoma, 385 americana, 385, 386 bifida, 386 chiropterae, 386, 386 Heterotaxis, 426 Hexadesmia, 555 bifida, 557 boliviensis, 557 dunstervillei, 557 fusiformis, 558 sessilis, 559 Hexisea, 387 bidentata, 387 var. imbricata, 387 imbricata, 387, 388 reflexa, 546 Hierba de boca, 102 Hoehneella santos-nevesii, 274 Hoja de culebra, 658 Hormidium, 536 Hostmannia, 129 elvasioides, 130 sagoti, 130 Houlletia, 388 odoratissima, 389, 390 roraimensis, 389, 390 vittata, 247 Huebneria yauaperyensis, 342, 484 Huevillo de agua, 172 Humboldtia, 579 aspasicensis, 513 biflora, 601 dura, 600 fockei, 515 foliata, 600 memor, 601 orbicularis, 601 revoluta, 520 zephyrina, 523 Huntleya, 390 albido-fulva, 390 meleagris, 390, 391 Hybochilus huebneri, 530 Hylaeorchis, 392 petiolaris, 392, 393

—I— Ihchan-tó-pa-mën-tu-kinketipeñ, 106 Ionopsis, 394 paniculata, 395 pygmaea, 591 satyrioides, 394, 395 utricularioides, 395 Ipiya, 25 Iramatemoko, 87 Ishakyek, 107 Isochilus, 395 elegans, 306 linearis, 396, 396 Iwa-pichuna, 63 —J— Jacquiniella, 396 globosa, 397, 397 steyermarkii, 397, 398 teretifolia, 398, 398 Jada-jada, 114 Jadasa, 64 Jambosa vulgaris, 98 Jarasa, 62 Jasmín colorado, 110 Jasmín de cafetal, 110 Jasmín de tarde, 110 Jujuli, 62 Jussiaea, 188 affinis, 190 decurrens, 190 densiflora, 190 erecta, 190 foliobracteolata, 190 helminthorrhiza, 190 hyssopifolia, 191 inclinata, 191 latifolia, 191 leptocarpa, 191 var. genuina, 191 linifolia, 190, 191 lithospermifolia, 192 var. pubescens, 192 var. typica, 192 maypurensis, 191 mollis, 191 natans, 190 nervosa, 191 var. pubescens, 191

751

var. typica, 191 peruviana var. glaberrima, 191 var. typica, 191 peruviana, 191 quadrangularis, 192 rigida, 192 sedoides, 192 suffruticosa, 191 torulosa, 192 —K— Kanau, 60 Kanomi-dek, 91 Kaso’ kana, 657 Kaso’kana, 646 Kegeliella, 398 orientalis, 399, 399 Koellensteinia, 399 carraöensis, 400, 401 eburnea, 401 graminea, 400 hyacinthoides, 401 lilijae, 401 roraimae, 401 tricolor, 401 Kraenzlinella, 503 Krugia, 40 ferruginea, 44 Kumachi, 62 —L— Laelia marginata, 562 sect. Schomburgkia, 561 undulata, 563 Lagopus, 739 Lanium, 325 microphyllum, 339 Laurel, 130 Leaoa, 555 reedii, 559 Leitgebia, 150 colombiana, 152 gleasoniana, 152 guianensis, 152 imthurniana, 152 Leochilus, 402 labiatus, 402, 402 Lepanthes, 403 aithalos, 404

752

I NDEX

[Lepanthes] cercion, 404 duidensis, 404, 405 dussii, 404 exilis, 405 funerea, 515 helicocephala, 405, 405 marahuacensis, 405, 405 orbiculata, 601 pariaënsis, 406 pectinata, 406 unitrinervis, 405, 406 yauaperyensis, 518 Lepanthopsis, 406 floripecten, 407, 407 obliquipetala, 407, 408 pulchella, 408 vinacea, 408 Lepianthes, 705 peltatum, 723 umbellatum, 726 Leptorchis, kappleri, 412 Leucohyle, 408 mutica, 409, 409 Leuconymphaea amazonum, 120 blanda, 120 Ligeophila, 409 amazonica, 410 juruensis, 410 stigmatoptera, 410, 410 Limodorum altum, 362 diurnum, 320 lanceolatum, 551 pendulum, 303 Limoncillo, 81, 168, 177, 186 Linociera, 186 caribea, 187 Liparis, 411 elata, 413 elliptica, 412 jamaicensis, 412, 412 kappleri, 413 nervosa, 412 f. kappleri, 413 subsp. nervosa, 412, 413 subsp. A, 413 verticillata, 413 vexillifera, 412 Liriosma, 170

candida, 172 crassa, 172 guianensis, 172 inopiflora, 172 macrophylla, 172 micrantha, 172 tepuiensis, 172 Llantén, 739 Lockhartia, 413 acuta, 414 imbricata, 414, 415 lasseri, 414 latilabris, 415, 415 pallida, 414 weigelti, 415 Lophiaris, 592 carthagenensis, 594 fragrans, 596 lanceana, 596 nana, 596 Ludwigia, 188 affinis, 190, 193 decurrens, 190, 194 densiflora, 190, 192 erecta, 190 foliobracteolata, 190 helminthorrhiza, 190 hyssopifolia, 190, 195 inclinata, 191, 196 latifolia, 191, 194 leptocarpa, 191, 192 linifolia, 191 lithospermifolia, 192 natans, 190 nervosa, 191, 195 octovalvis, 191, 193 peruviana, 191, 192 quadrangularis, 192 rigida, 192 sedoides, 192, 196 suffruticosa, 191 torulosa, 192, 197 Lueddemannia, 415 pescatorei, 416, 416 Luxemburgia duidae, 148 longifolia, 148 Lycaste, 416 fulvescens, 417 lindeniana, 495 macrophylla, 417, 417 Lyroglossa, 418 grisebachii, 418, 418

Lysimnia, 248 —M— Macradenia, 419 brassavolae, 420, 420 lutescens, 420 modesta, 420 mutica, 409 rubescens, 420 Macroclinium, 420 mirabile, 421, 421 norae, 421 wullschlaegelianum, 422, 422 Ma-hana-hana, 114 Mahchó kaí mën, 62 Maikanpi-miyuyek, 12 Majako shodö, 26 Malaxis, 422 excavata, 422 maguirei, 422, 423 Manchaua, 723 Manniella americana, 385 Manteco, 106 Manzana rosa, 98 Mapurite, 674 Mapuu ke thotho, 659 Mapuu thotho, 651, 659 Maracuyá, 644 Marlierea, 40 acuminata, 44 bipennis, 43, 52 buxifolia, 43, 49 caesariata, 43, 48 cana, 43, 53 caudata, 43 convexivenia, 43 cuprea, 43 dussii, 95 ensiformis, 44 ferruginea, 44 foveolata, 44 guildingiana, 44, 47 insignis, 44 intonsa, 63 karuaiensis, 44 ligustrina, 44 lituatinervia, 45 maguirei, 45, 53 mcvaughii, 45

I NDEX

montana, 45, 50 obtusa, 45 pudica, 45, 48 rugosior, 45, 50 schomburgkiana, 45, 49 scytophylla, 46 spruceana, 46, 49 subcordata, 46 suborbicularis, 46, 51 subulata, 46 summa, 46, 51 uaupensis, 46 umbraticola, 46, 52 uniflora, 47, 53 ventuarensis, 47 sp. A, 47 Marlieriopsis, 4 eggersii, 4 Marsupiaria, 426 equitans, 437 mattogrossensis, 437 Masdevallia, 309, 423 brevis, 554 guayanensis, 424 guttullata, 424 huebneri, 607 manarana, 424, 425 norae, 424, 425 picturata, 425 sect. Rhombopetalae, 309 sect. Saltatrices, 309 sprucei, 425, 425 triaristella, 607 sect. Triaristellae, 606 triglochin, 607 wendlandiana, 425, 425 Maxillaria, 392, 426 acutifolia, 433 alba, 434 albiflora, 434 alpestris, 434, 446 alticola, 434 amazonica, 438, 441 anatamorum, 434 arachnitis, 440 attenuata, 438 aurea, 434, 447 auyantepuiensis, 435, 439 bolivarensis, 435, 448 brachybulbon, 435, 449 brevipedunculata, 440 brunnea, 436, 441 buchtienii, 444

camaridii, 435 chlorantha, 435 colemanii, 435, 446 colleyi, 616 condensata, 439 conferta, 436 connellii, 436, 450 coriacea, 436 cristata, 494 cryptobulbon, 436 desvauxiana, 436, 451 discolor, 437, 447 ×dunstervillei, 437 eburnea, 437 equitans, 437 foldatsiana, 437,447 graminea, 400 grandiflora, 437 grobyoides, 437, 451 guadalupensis, 438, 452 guareimensis, 438, 451 iguapensis, 438 imbricata, 438 kegelii, 438, 446 lactea, 434, 438 lactiflora, 438 lasallei, 438 longifolia, 444 longipetiolata, 439 loretoensis, 439, 448 luteoalba, 439, 450 macrophylla, 417 mapiriensis, 439 marmoliana, 440 mattogrossensis, 437 melina, 434 meridensis, 440 minuta, 393 nasuta, 440, 442 notylioglossa, 440 nuriensis, 440, 453 ochroleuca, 440 var. longipes, 440 pallidiflora, 616 parkeri, 440, 448 pauciflora, 441 pendens, 442 pendula, 442 perparva, 393 petiolaris, 392, 436 porrecta, 441, 449 proboscidea, 441 pterocarpa, 442

purpurea, 436 quelchii, 442, 453 ramosa, 442 reichenheimiana, 442 rigida, 442 ringens, 441 riopalenquensis, 439 rudolfii, 392 rufescens, 433, 443 rugosa, 443 santanae, 443, 452 schlechteri, 443 scorpioidea, 443 serrulata, 434 setigera, 443, 454 similans, 436 spilotantha, 438 splendens, 444 squamata, 444 steelei, 564 stenophylla, 444 superflua, 444 surinamensis, 436 tafallae, 442 taracuana, 438 tenuis, 445, 452 terumensis, 444 trinitatis, 436, 441 uncata, 444, 445 vandiformis, 437 verrucifera, 436 villosa, 445, 449, violaceopunctata, 445 xylobiiflora, 445, 454 Mayba-pio-keu-yek, 62 Mayepea, 186 caribea, 187 Medadenium labiosum, 618 lindeniae, 618 Medewadi, 114 Mendoncella, 367 burkei, 368 jorisiana, 368 Merewadi, 106 Mesadenella, 455, 455 angustisegmenta, 455, 455 Microchilus, 360 Microstylis, 422 Microtea, 672 debilis, 673, 674 maypurensis, 674

753

754

I NDEX

Mijarro, 106 Miltonia, 456 Miltoniopsis, 456 santanaei, 456, 456 Minquartia, 181 guianensis, 181, 182 Mirabilis, 109 jalapa, 110, 110 Mitranthes sartoriana, 92 Monachanthus, 260 discolor, 263 roseo-albus, 264 Morichalero, 114 Mormodes, 457 carnevaliana, 457, 458 cucumerina, 458, 458 orinocoensis, 458 vernixioidea, 458 subsp. autanensis, 458, 458 Motodi, 658 Motodipicada, 658 Murciélago, 649 Myanthus, 260 barbatus, 262 Myoxanthus, 459 aspasicensis, 513 subsp. arenicola, 512 lappiformis, 517 parvilabius, 460, 461 reymondii, 460 simplicicaulis, 460 speciosus, 460, 461 trachychlamys, 460 Myrcia, 54 aguitensis, 61 albido-tomentosa, 59 aliena, 59 amazonica, 59, 68 angustifolia, 60 bipennis, 43 bolivarensis, 59, 68 bonnetiasylvestris, 59, 69 bracteata, 59, 70 calycampa, 60 chilensis, 59 citrifolia, 60, 70 clausa, 60 clusiifolia, 60, 70 compta, 60 crispa, 60, 71, decorticans, 61, 72

deflexa, 61, 72 dichasialis, 61, 69 dumosa, 64 edulis, 64 ehrenbergiana, 61 exploratoris, 61, 69 fallax, 61, 73 fenzliana, 62 ferruginea, 44 gentryi, 62, 77 grandis, 62, 74 granulata, 86 guianensis, 62 var. guianensis, 62, 75 hirtellaefolia, 60 humboltiana, 61 inaequiloba, 62, 71 induta, 63 intonsa, 63 kegeliana, 61 kylistophylla, 63 laevis, 63 latifolia, 60 leptoclada, 59 liesneri, 63 lucida, 63, 75 var. attenuata, 63 magnoliifolia, 63, 74 minutiflora, 63, 75 multiflora, 63, 74 nubicola, 64 obtusa, 62 paivae, 64 pistrinalis, 64 planifolia, 65 platyclada, 64, 76 ptariensis, 64, 76 pyrifolia, 64 quitarensis, 65 revolutifolia, 65, 73 roraimae, 62 rotundata, 65 var. atrans, 65 var. rotundata, 65, 76 salticola, 65 schomburgkiana, 65 servata, 65, 71 sessiliflora, 66 sipapensis, 66 sororopanensis, 66, 77 splendens, 66, 78 spruceana, 61 subsessilis, 66

var. ovalis, 66 var. subcordata, 66 var. subsessilis, 66, 78 sylvatica, 66 tepuiensis, 67, 78 tomentosa, 67, 78 sp. A, 67 sp. B, 67 sp. C, 67 sp. D, 67 Myrcianthes, 79 fragrans, 79, 80 prodigiosa, 80, 80 rhopaloides, 80 umbellulifera, 87 Myrciaria, 81 bipennis, 43 caurensis, 81 cordata, 81 dubia, 81, 82 ehrenbergiana, 61 floribunda, 82, 83 involucrata, 86 lituatinervia, 45 nitida, 62 paraensis, 81 protracta, 82 puberulenta, 84 quitarensis, 65 tenella, 84 uliginosa, 82 verticillata, 82 vismeifolia, 83, 84 Myrobroma, 609 Myrtaceae, 1 Myrteola, 84 nummularia, 84, 85 Myrtus alternifolia, 6 biflora, 22 calophylla, 6 citrifolia, 60 clusiaefolia, 60 cumini, 96 deflexa, 61 emarginata, 24 erythroxyloides, 87 fragrans, 79 goetheana, 8 latifolia, 60 ligustrina, 25 monticola, 26 myricoides, 98

I NDEX

var. stenophylla, 98 myricoides var. roraimensis, 98 var. turumiquirensis, 98 nummularia, 85 pubescens, 27 punicifolia, 27 rhopaloides, 80 roraimensis, 98 salutaris, 92 splendens, 66 stenophylla, 98 sylvatica, 66 umbellulifera, 87 umbraticola, 46 —N— Najadaceae, 100 Najas, 100 wrightiana, 100, 101 Nanodes, 325 Neea, 110 amaruayensis, 113 bernardii, 113 bracteosa, 113 brevipedunculata, 113 cauliflora, 114 cedenensis, 114 clarkii, 114 davidsei, 114 floribunda, 114 grandis, 114 guaiquinimae, 114 huachamacarae, 114 ignicola, 114 liesneri, 114 mapourioides, 115 marahuacae, 115, 116 neblinensis, 115 obovata, 115, 116 ovalifolia, 115 parimensis, 115, 116 robusta, 115 sebastianii, 117 subglabrata, 117 tepuiensis, 117 tristis, 117 Neippergia, 230 Neobartlettia guianensis, 492 Neolehmannia, 325 barbeyana, 334

pabstii, 337 Neottia acaulis, 553 calcarata, 316 Nidema, 462 ottonis, 462, 462 Nirguo rebalsero, 677 Notylia, 463 amesii, 464 angustifolia, 464 aromatica, 464 fragrans, 464 laxa, 464 longispicata, 464 microchila, 464 mirabilis, 421 nana, 464 norae, 421 pentachne, 465 peruviana, 465 platyglossa, 465 rhombilabia, 465, 465 trullifera, 464 venezuelana, 464 wullschlaegeliana, 422 yauaperyensis, 465 Nudillo, 718 Nudito, 723 Nyctaginaceae, 101 Nymphaea, 119 potamophila, 121 amazonum, 120, 122 ampla, 121 blanda, 120 var. amazonum, 120 conardii, 120, 122 gardneriana, 120, 123 glandulifera, 120, 122 pulchella, 121, 121 rudgeana, 123, 124 var. amazonum, 120 Nymphaeaceae, 118 —O— Ocampoa, 287 Ochnaceae, 124 Ocoma, 199 Octomeria, 465 anomala, 470 apiculata, 470 brevifolia, 472 complanata, 473

755

connellii, 470 cordilabia, 470, 473 deltoglossa, 470 dentifera, 470 erosilabia, 470, 473 exigua, 470, 474 var elata, 470 filifolia, 470 flaviflora, 471 gemmula, 471, 474 heleneana, 471 integrilabia, 471 kestrochila, 470 lancipetala, 471 minor, 471 monticola, 471 nana, 471 parvifolia, 471 parvula, 471 rhizomatosa, 472 romerorum, 472, 474 scirpoidea, 472, 474 spathulata, 472 splendida, 472 steyermarkii, 472, 474 surinamensis, 472 taracuana, 472 yauaperyensis, 473 sp. A, 473 Odontoglossum, 474, 487 arminii, 488 sect. Aspasia, 234 brevifolium, 488 naevium, 475, 475 subgen. Otoglossum, 487 Oeceoclades, 476 maculata, 476, 476 Oenothera octovalvis, 191 Ojo de pavón, 12 Olacaceae, 162 Olax macrophylla, 172 Oleaceae, 186 Onagraceae, 188 Oncidium, 477, 479 ampliatum, 277 sect. Aphanobulbia subsect. Iridifolia, 359 baueri, 479 bernouillanum, 277 boothianum, 479 carthagenense, 594

756

I NDEX

[Oncidium] cebolleta, 594 sect. Cebolletae, 592 citrinum, 479 convolvulaceum, 488 dactyliferum, 479 sect. Glanduligera, 542 globuliferum, 488 glossomystax, 360 sect. Iridifolia, 359 kappleri, 479 lanceanum, 596 lunatum, 577 nanum, 596 nigratum, 479 sect. Oblongata, 276 orthostates, 480, 480 papilio, 543 pardalis, 480 pusillum, 360 sancti-pauli, 488 scansor, 488 sect. Serpentia, 487 sphacelatum, 480 uaipanense, 479 vagans, 480 sect. Varicosa subsect. Serpentia, 487 warmingii, 480, 481 Onychium flavescens, 532 Ophidion, 482 pleurothallopsis, 482, 482 Ophrys nervosa, 412, 413 Opiliaceae, 198 Orchidaceae Family Description, 203 Glossary, 205 History, 200 Illustrated Glossary, 206– 207 Key to the Subkeys, 208 Orchidofunckia, 288 Orchis monorrhiza, 380 quinqueseta, 381 stylomisantha, 367 Orleanesia, 482 amazonica, 484 maculata, 484 yauaperyensis, 483, 484 Ornithidium, 426

aureum, 435 chloroleucum, 436 confertum, 436 mapiriense, 439 serrulatum, 434 squamatum, 444 vestitum, 436 Ornithocephalus, 484 bicornis, 485 brachyceras, 485 falcatus, 485, 486 gladiatus, 486, 486 kruegeri, 486 planifolius, 307 Oroshomi thotho, 645, 648, 653 Otoglossum, 487 arminii, 488, 489 globuliferum, 488 scansor, 488 Otopetalum, 503 Otostylis, 489 alba, 491 brachystalix, 490, 491 lepida, 491 paludosa, 491 venusta, 491 Ottonia ovata, 723 Ouratea, 131 angulata, 134 aquatica, 134 arbobrevicalyx, 134 articulata, 135 asisae, 135 attenuata, 135 ayacuchae, 135 brevicalyx, 135 brevipedicellata, 135 castaneifolia, 135 chaffanjonii, 135 clarkii, 135 coccinea, 135, 138 coccinea, 137 crenata, 139 croizatii, 136 culminicola, 136 davidsii, 136 deminuta, 136 discophora subsp. pervenulosa, 136 discophora, 136, 142 duidae, 136

engleri, 136 evoluta, 136 ferruginea, 136, 143 fusiformis, 137 grandiflora, 137 grosourdyi, 137 guaiquinimensis, 137 guianensis, 137, 140 guriensis, 137 heterobracteata, 137 huberi, 137 inconformis, 141 inundata, 141 lajaensis, 137 leblondi, 138, 143 liesneri, 138 longifolia, 137 longistyla, 138 macurucoensis, 141 maguirei, 138 maigualidae, 138 marahuacensis, 138 medinae, 138 megaphylla, 138 multibracteata, 138 obovata, 138 odora, 141 oligantha, 138 orisina, 139 ornata, 139 papillata, 139 paratatei, 139 paruensis, 139 pisiformis, 139 pittieri, 137 polyantha, 139, 142 propingua, 139 pseudoguildingii, 139 ptaritepuiensis, 139 pulverulenta, 140 pumila, 134 var. heterodoxa, 134 ramosissima, 140, 142 rigida, 140 roraimae, 140, 144 rorida, 140 rotundipetala, 140 sagotii, 138 saldariagae, 140 sipapoensis, 140, 144 soderstromii, 140 spruceana, 141, 143 squamata, 141

I NDEX

steyermarkii, 141, 145 subamplexicaulis, 141 superba, 141 tatei, 141 thyrsoidea, 141 vasivae, 137 venezuelensis, 141 wurdackiana, 141 yapacanae, 141, 144 Oxalidaceae, 619 Oxalis, 624 barrelieri, 624, 625 borjensis, 625 calophylla, 621 frutescens, 625 subsp. borjensis, 625 subsp. frutescens, 625 juruensis, 625 pentantha, 625 —P— Pabajuto metsaja, 718 Paduari-yek, 27 Palmorchis, 491 guianensis, 492, 493 puber, 492, 493 pubescens, 492, 493 Palo chaparro, 109 Palo chaparro banero, 109 Palo culebra, 115 Palo de cachicamo, 114 Palo de culebra, 113, 114, 115 Palo de culebra, 115 Palo de cuyubi, 181 Palo de mono, 106, 107 Palo de panema, 177 Palo de rábano, 127 Palo perro de agua, 115 Paphinia, 494 cristata, 494 dunstervillei, 494 lindeniana, 495, 495 Paphiopedilum sect. Phragmopedilum, 498 Papiliopsis, 542 Parcha, 641, 643, 644, 649, 650, 651, 653, 656, 658 Parcha de culebra, 641, 652, 653

Parcha de culebra de agua, 658 Parcha de huesito, 649 Parcha de ratón, 656 Parcha diente de culebra, 655 Parcha grenadina, 653 Parcha montañera, 655, 658 Parchita, 644, 652 negra, 640 Parchita de monte, 652 Pariente cuajo, 671 Pasaña, 110 Pasionaria, 652 Pasita, 177 Pasita hoja fina, 177 Passiflora, 630 acuminata, 640 adenopoda, 640 ambigua, 640, 660 amicorum, 641 auriculata, 641, 661 bomareifolia, 658 capparidifolia, 641, 660 cardonae, 642 cauliflora, 642 cirrhiflora, 643 coccinea, 643 costata, 643 edulis, 644, 662 Flower Diagram, 632 foetida, 644 var. foetida, 645, 663 var. gossypiifolia, 645 var. hispida, 645 var. moritziana, 645 var. orinocensis, 645 fuchsiiflora, 645 garckei, 646, 663 glandulosa, 646, 661 gleasonii, 647 gossypifolia, 645 gracilis, 647 guazumaefolia, 647 hispida, 645 holtii, 647 laurifolia, 648 lonchophora, 654 longiracemosa, 648, 662 maguirei, 649 maliformis, 649, 664 misera, 649 moritiziana, 645

757

multiformis, 650 nitida, 650, 667 nuriensis, 651 ovata, 651 pallida, 657 pedata, 651 subsp. stipularis, 651 var. stipularis, 651 phaeocaula, 651 picturata, 652 pulchella, 652, 665 pyrrhantha, 652 quadrangularis, 653 quadriglandulosa, 653, 663 retipetala, 654, 662 riparia, 654 rubra, 655 sclerophylla, 655, 661 securiclata, 655, 666 seemannii, 656, 664 serrulata, 656, 664 spinosa, 657 suberosa, 657 tuberosa, 657 variolata, 658, 666 vespertilio, 658, 665 sp. A, 659 sp. B, 659 sp. C, 659 Passifloraceae, 625 Paují, 87 Pawaruyek, 44 Pebajú, 677 Pedaliaceae, 667 Pelexia, 495, 552 aphylla, 553 callifera, 496, 496 cranichoides, 291 laxa, 496 maculata, 496 setacea, 316 Pendanga, 26 Pen-danga, 23 Pendango, 26 Pentaspatella, 150 ramosa, 153 Peperidia, 705 Peperomia, 681 acuminata, 690 alata, 690 var. angustifolia, 690 alpina, 690, 704

758

I NDEX

[Peperomia] angustata, 690, 704 atabapoensis, 696 bartlettii, 697 blanda, 691 var. langsdorfii, 691 boomii, 693 celiae, 691 chimantana, 693 choroniana var. puberulenta, 693 ciliata, 691 cladara, 691, 701 f. ciliata, 691 f. perglabra, 691 cryptotricha, 698 cuneata, 695 decipiens, 691 delascioi, 691 dendrophila, 691 diffusa, 694 distachya, 691 f. geminispica, 691 duidana, 699 elongata var. guianensis, 692 var. piliramea, 692 emarginata, 695 emarginella, 692, 701 var. glabrior, 692 enantiostachya, 692 epilobioides, 696 ernstiana, 698 foveolata, 691 fundacionensis, 692 galioides, 692 geminispica, 691 gentryi, 692 glabella, 693 var. glabella, 693 var. nervulosa, 693 guaiquinimana, 693, 699 haematolepis, 693 hederacea, 698 hernandiifolia, 693, 700 horrescens, 698 ionophylla, 697 jamesoniana, 693, 704 lanceolato-peltata, 693 lancifolia, 694, 699 subsp. erasmiaeformis, 694 longemucronata, 690

f. longirostrata, 690 longirostrata, 690 loxensis, 694 macaroana, 691 macrostachya, 694 var. macrostachya, 694, 704 var. nematostachya, 694 maculosa, 694, 700 magnoliaefolia var. emarginulata, 695 magnoliifolia, 694 major, 698 marahuacensis, 695 maypurensis, 695, 701 melanostigma var. nervulosa, 693 microreticulata, 690 moritzii, 696 myosuroides, 694 neblinana, 695 nematostachya, 694 nilssonii, 690 nummularifolia, 697 obtusifolia, 695, 702 var. cuneata, 695 var. emarginulata, 695 var. emarginata, 695 omnicola, 697 var. oblanceolata, 697 ornata, 695 ouabianae, 695, 700 pellucida, 695, 701 peltata, 723 pennellii, 691 pernambucensis, 696, 703 pseudojamesoniana, 693 purpurascens, 697 purpurinervis, 696, 704 pyrifolia, 690 quadrangularis, 696, 700 quaesita, 696, 699 reflexa, 698 f. americana, 698 var. americana, 698 roraimana, 697 rotundata, 696 var. rotundata, 696 var. trinervula, 696 rotundifolia, 697, 700 f. ovata, 697 var. ovata, 697 saginans, 696

salvaje, 691 serpens, 697, 702 silvestris, 695 spruceana, 697 striata, 697, 701 tafelbergensis, 695 tenella, 697 var. tenella, 698 var. tyleri, 698, 702 tetraphylla, 698 var. americana, 698 trinervula, 696 var. suboppositifolia, 696 tyleri, 698 uaupesensis, 698 umbellata, 726 urocarpa, 698, 702 venezueliana, 698 venusta, 698, 703 victoriana, 690 var. marginata, 690 viridispica, 690 wurdackii, 695 yatuensis, 698, 699 yutajensis, 699, 703 Peridiscaceae, 670 Peridiscus, 671 lucidus, 671, 671 Perissocarpa, 145 steyermarkii, 145, 146 umbellifera, 145, 146 Peristeria, 497 aspersa, 497, 498 cerina, 497 guttata, 497 pendula, 498 Pescuezo de pava, 677 Petalocentrum, 567 Petiveria, 674 alliacea, 674, 675 foetida, 674 Philacra, 146 auriculata, 147, 148 duidae, 147, 148 longifolia, 148 steyermarkii, 147, 148 Phloeophila, 503 Phragmipedium, 498 kaieteurum, 499 klotzschianum, 499, 499 lindleyanum, 499, 499 Phyllocalyx, 17

I NDEX

Phymatidium panamense, 316 Physinga, 325 prostrata, 341 Physosiphon, 503 obliquipetalus, 408 Physurus, 360 arietinus, 361 juruensis, 410 paleaceus, 361 pumilus, 236 santensis, 410 stigmatopterus, 410 Phytolacca, 674 rivinoides, 676, 677 rugosa, 677 thyrsiflora, 677 Phytolaccaceae, 672 Piitirijrida, 25 Pilumna, 596 Pimienta negra, 722 Pinelia, 500 alticola, 500 Pinelianthe, 500 alticola, 500, 500 Piper, 705 acutissimum, 726 adenandrum, 714 aduncum, 714 var. aduncum, 714 var. brachyarthrum, 714 var. cordulatum, 714 var. garcia-barrigae, 714 aequale, 714 var. santa-martanum, 715 agostinii, 721 alatabaccum, 715 alpinum, 690 amalago, 715, 727 var. medium, 715 amazonasense, 716 amazonicum, 718 angustatum, 690 angustifolium var. cordulatum, 714 angustum, 716 anisatum, 721 anonifolium, 715 var. anonifolium, 715, 727 var. parkerianum, 715

arboreum, 715 var. arboreum, 716, 728 var. falcifolium, 716 var. hirtellum, 716 var. latifolium, 716 arieianum, 725 var. chimantanum, 726 arieianum var. duidaense, 726 aristeguietae, 722 asperifolium, 720 augustum, 716 avellanum, 716, 728 baccans, 716 var. sancarlosianum, 716, 729 bartlingianum, 717, 730 var. parimanum, 717 blandum, 691 bolivaranum, 717, 729 cabellense, 717 caripense, 714 caudatum, 721 cauranum, 718 ceanothifolium, 715 celtidifolium, 714 cernuum, 717 var. cernuum, 717 var. perlongispicum, 717 chimantanum, 726 cililimbum, 717, 729 citrifolium, 715 var. parkerianum, 715 consanguineum, 717, 727 consanguineum, 717 coruscans, 718, 731 var. membranaceum, 718 crassinervium, 718 crassum, 719 cumanense, 718, 732 cuyunianum, 718 demeraranum, 718, 727 var. rubescens, 718 deminutum, 717 dilatatum, 719 distachyon, 691 divaricatum, 719 duidaense, 726 var. eciliatum, 726 dunstervilleorum, 719 elongatum

759

var. brachyarthrum, 714 var. cordulatum, 714 emarginatum, 695 emarginellum, 692 esmeraldanum, 718 eximium, 717 falcifolium, 716 flagellare, 722 foveolatum, 719, 731 francovilleanum, 719 funckii, 714 geniculatum, 716 f. puberula, 726 var. latifolium, 716 gentryi, 719 gigantifolium, 717 glabellum, 693 glabrilimbum, 719 guianense, 719, 733 hermannii, 719 hernandiifolium, 693 hippocrepiforme, 720, 729 hirsutum, 720 hispidum, 720 holtii, 720 hostmannianum, 720, 734 var. glabrirameum, 720 var. ramiflorum, 720 jauaense, 721, 732 javitense, 721 johnstonii, 714 julianii, 721 kegelianum, 721, 734 latum, 724 lemaense, 721, 735 leprieuri, 717 liesneri, 721 loxense, 694 lucaeanum var. magnifolium, 725 macrophyllum, 716 macrostachyon, 694 maculosum, 694 magnoliifolium, 694 marginatum, 721 var. anisatum, 721 var. marginatum, 721, 734 maypurense, 695 medium, 715 morilloi, 722 mosaicum, 722 myosuroides, 694

760

I NDEX

[Piper] neblinanum, 722, 730 nigrum, 722 nitidum, 716 nummularifolium, 697 obliquum, 722 var. eximium, 717 obrumbatum, 716 obtusifolium, 695 oparumaense, 722 orenocanum, 718 otto-huberi, 722 var. eciliatum, 722, 728 var. otto-huberi, 722 ovatum, 723, 730 papilliferum, 723, 735 parapeltobryon, 723 parimanum, 717 parkerianum, 715 pavasense, 721 pellucidum, 695 peltatum, 723, 736 perciliatum, 723, 737 perlongispicum, 717 perlongivillosum, 719 perstipulare, 723, 732 piscatorum, 723, 737 poiteanum, 719 politii, 723 subsp. politii, 724 subsp. sipapoense, 724, 737 subsp. toronoense, 724 pruinosum, 723 pseudoacreanum, 724, 731 pseudochurumayu var. membranaceum, 718 pseudoglabrescens, 724, 738 pubivaginatum, 724 quadrangulare, 696 reflexum, 698 reticulatum, 724, 733 rio-paraguanum, 720 rotundifolium, 697 rubescens, 718 ruizianum, 722 rupununianum, 724 sabanaense, 725, 735 saltuum, 726 sancarlosianum, 716

serpens, 697 smilacifolium, 724 speciosum, 723 statarium, 716 steyermarkii, 725, 738 subalpinum, 725 subcrassifolium, 720 subduidaense, 725, 733 surinamense, 717 tamayoanum, 725, 738 tenellum, 697 tenue, 725 tepuiense, 725 tetraphyllum, 698 tonoroanum, 716 toronotepuiense, 725 tortivenulosum, 725 trigonum, 725 tuberculatum, 726 var. minus, 726 var. scandens, 726 umbellatum, 726 variegatum, 718 venamoense, 726, 732 verrucosum, 716 vitaceum, 726 var. venezuelense, 726 wachenheimii, 726 wurdackii, 718 zedigodiense, 715 Piperaceae, 681 Piriko-so-yek, 8 Pisonia, 117 aculeata, 118, 118 cuspidata, 106 eucalyptifolia, 107 ferruginea, 106 fragrans, 107 microphylla, 107 rusbyana, 109 Pittierella, 289 Plantaginaceae, 739 Plantaginella, 739 Plantago, 739 major, 739, 739 Platystele, 500 johnstonii, 501, 501 lancilabris, 502 ovalifolia, 501 oxyglossa, 501 Plectrophora, 502 cultrifolia, 502 iridifolia, 502, 502

Pleurothallis, 470, 503, 513, 515, 598, 600 acutissima, 521, 522 amygdalina, 600 anomala, 600 aondae, 512 apiculata, 470 archidiaconi, 512 arenicola, 512, 524 aristata, 512 aspasicensis, 513 barbata, 513 barberiana, 512 barbulata, 513 biflora, 601 sect. Bipaleolatae, 598 sect. Brachystachyae subsect. Lepanthiformes, 598 brevipes, 513 brevis, 601 breviscapa, 515 broadwayi, 600 subsp. anomala, 600 callifera, 513 cedralensis, 600 ciliaris, 601 ciliata, 517 var. elongata, 517 ciliolata, 515 coffeicola, 520 connata, 408 consimilis, 515, 523 corazonica, 600 coriacardia, 513 corniculata, 514, 525 deborana, 514, 523 delascioi, 517 diffusiflora, 521 discophylla, 514 dura, 600 egleri, 600 elegans, 518 elvirana, 514, 527 erebatensis, 514 erinacea, 514 exasperata, 460 fimbriata, 518 floribunda, 514 floripecten, 407 fockei, 515 foliata, 600 funerea, 515

I NDEX

galeata, 515 garciae, 515 geographica, 515 gnomonifera, 601 granitica, 515, 528 grobyi, 516, 527 guadalupensis, 600 hexandra, 516, 523 hilariana, 516 hitchcockii, 516 holstii, 516, 524 huebneri, 517 humilis, 516 imraei, 516 intricata, 600 ionantha, 517 johnstonii, 501 kegelii, 516, 517 kerrii, 517, 523 lanceana, 517, 525 lancifera, 601 lancilabris var. oxyglossa, 501 lappiformis, 517, 526 sect. Lepanthiformes, 598 lepanthopsis, 600 linguifera, 513 longisepala, 517 loranthophylla, 517 maguirei, 518 memor, 600 mentosa, 518 minima, 518 minutipetala, 601 miqueliana, 518, 526 monocardia, 512 morilloi, 518, 528 moritzii, 518, 524 myrmecophila, 515 nanifolia, 519 obliquipetala, 408 omissa, 519 orbicularis, 601 ovalifolia, 501 oxyglossa, 501 papillosa, 516 parvifolia, 519 parvilabia, 460 pemonum, 519, 523 penduliflora, 521 pertenuis, 522 phoenicoptera, 519 picta, 519

plumosa, 517 pluriracemosa, 514 prolifera, 519 pruinosa, 520 pulchella, 522 pumila, 602 punctata, 517 revoluta, 520, 521 reymondii, 460 rhombipetala, 520 ringens, 516 roraimensis, 603 ruscifolia, 520 samacensis, 520, 526 sandaliorum, 520 sarcosepala, 520 scandens var. simplicicaulis, 460 sclerophylla, 521, 528 seriata, 521 sertularioides var. trinitensis, 522 setigera, 523 simplicicaulis, 460 speciosus, 460 spiculifera, 521, 522, 527 steinbuchiae, 521 stenocardium, 521 stenopetala, 521 suspensa, 521 tepuiensis, 519, 522, 525 trachyclamys, 460 trachytheca, 601 tricarinata, 515 tridentata, 522 trinitensis, 522, 527 tubulosa, 547 uncinata, 512, 513 vaginata, 516 velaticaulis, 514, 522 velatipes, 522 viridula, 547 vitatta, 520 vittariifolia, 522, 526 wendlandiana, 520 williamsii, 600 wilsonii, 522 yauaperyensis, 518, 522 zephyrina, 523 sp. A, 523 Plinia, 85 fruticosa, 95 involucrata, 86, 86

761

pinnata, 86 rivularis, 86 sp. A, 87 sp. B, 87 Plumbaginaceae, 740 Plumbago, 740 scandens, 740 zeylanica, 740, 741 Poecilandra, 148 pumila, 148, 149 retusa, 148 var. retusa, 149, 150 var. sclerophylla, 149, 150 sclerophylla, 150 Pogonia lepida, 279 macrophylla, 541 moritzii, 279 parviflora, 279 physuraefolia, 541 rosea, 279 stricta, 279 surinamensis, 606 tenuis, 279 triflora, 280 unifoliata, 280 Poko mën, 621 Polycycnis, 529 surinamensis, 529, 529 vittata, 247 Polyotidium, 530 huebneri, 530, 530 Polystachya, 531 amazonica, 532 cavanayensis, 532 concreta, 532, 532 flavescens, 532 foliosa, 532 stenophylla, 532 Pomagás, 98 Pomarosa, 98 Ponera amethystina, 560 felskyi, 558 leucantha, 558 Ponthieva, 532 diptera, 533 ovatilabia, 533, 534 Poró, 643 Pothomorphe, 705 peltata, 723 umbellata, 726

762

I NDEX

Prescottia, 533 auyantepuiensis, 535 carnosa, 535 oligantha, 534, 535 orchioides, 535 stachyodes, 535 tepuyensis, 536 Promenaea graminea, 400 Prosthechea, 536 aemula, 538, 539 calamaria, 538 crassilabia, 538 jauana, 539, 539 pamplonensis, 540 pygmaea, 539 tigrina, 539 vespa, 540 Proyek, 643 Pseudanamomis, 87 umbellulifera, 87, 88 Pseudoeriopsis, 357 schomburgkii, 358 Pseudoeurystyles, 363 Pseudomaxillaria, 426 chloroleuca, 436 Psidium, 88 acutangulum, 90, 93 aquaticum, 92 aromaticum, 16 australe, 90 cinereum, 90 densicomum, 91, 94 dubium, 81 glaucescens, 91 grandiflorum, 16 guajava, 91 guildingianum, 44 guineense, 91, 93 guyanense, 91 laruotteanum, 91 maribense, 91, 94 ovatifolium, 91 parviflorum, 92 persoonii, 90 quinquedentatum, 91 salutare, 92, 92 sartorianum, 92, 93 striatulum, 92 submetrale, 91 Psilochilus, 540 dusenianus, 541, 541 macrophyllus, 541

Psychopsiella, 542 Psychopsis, 542 papilio, 543, 543 picta, 543 Psygmorchis, 359 glossomystax, 360 pusilla, 360 Psyllium, 739 Pterichis, 543 galeata, 544, 544 —Q— Quekettia, 544, 590 microscopica, 545, 545 pygmaea, 591 —R— Rabo de rata, 718 Raek-i, 87 Raíz de candela, 723 Raíz de mato, 723 Raiz de muela, 718 Raíz de muela, 723 Reichenbachanthus, 545 modestus, 546 reflexus, 546, 546 Restrepia kegelii, 516 Restrepiella, 546 viridula, 547 Restrepiopsis, 546 tubulosa, 547, 547 viridula, 547 Rhaptostylum, 175 acuminatum, 177 pentandrum, 177 Rhynchadenia, 419 Rhynchopera, 503 punctata, 517 Rivina, 677 humilis, 677 var. humilis, 677, 678 octandra, 680 Rockia, 117 Rodriguezia, 547 batemanii, 548 candida, 548 lanceolata, 548, 549 leeana, 549 secunda, 548 Roraimanthus, 150

imthurniana, 152 Rudolfiella, 549 aurantiaca, 550, 550 —S— Sacoila, 550 lanceolata, 551, 551 Salvaje, 691 Santa Maria, 721 Sarcoglottis, 552 acaulis, 553 aphylla, 553 grandiflora, 553 metallica, 553 picta, 553 simplex, 553 stergiosii, 553, 553 Satyrium elatum, 291 orchioides, 551 Sauvagesia, 150 adima, 152 aliciae, 151 subsp. aliciae, 151, 154 amoena, 151, 155 angustifolia, 151 deficiens, 153 deflexifolia, 151, 155 duckei, 153 duidae, 151, 152 elata, 152 subsp. occidentalis, 152 erecta, 157 subsp. erecta, 152, 154 var. rubiginosa, 153 erecta, 152 erioclada, 152 var. grandiflora, 152 falcisepala, 152 fruticosa, 152 guianensis, 152 subsp. gleasoniana, 152 subsp. guaiquinimensis, 152 subsp. guianensis, 152, 156 subsp. sipapoensis, 152 imthurniana, 152, 155 subsp. chimantensis, 152 inconspicua, 157 kappleri, 157

I NDEX

laxa, 153 linearifolia, 153 subsp. venezuelensis, 153, 156 longifolia, 153 longipes, 153 marahuacensis, 152 miniata, 153 nudicaulis, 153, 154 ramosa, 153, 154 ramosissima, 153, 156 roraimensis, 153 rosacea, 152 rubiginosa, 153 serpyllifolia, 157 smithiana, 153 sprengelii, 155, 157 striata, 151 surinamensis, 153 tenella, 155, 157 Scaphosepalum, 554 breve, 554, 555 verrucosum, 554 Scaphyglottis, 555 amethystina, 560 bifida, 557 boliviensis, 557 cuneata, 559 dunstervillei, 557, 559 felskyi, 558 fusiformis, 558 graminifolia, 558 grandiflora, 558 huebneri, 557 leucantha, 558 michelangeliorum, 558 modesta, 558 prolifera, 559 propinqua, 559 reedii, 559 reflexa, 546 sessilis, 559 sickii, 559, 559 stellata, 560 tafallae, 442 violacea, 558 wercklei var. major, 558 Scelochilus, 560 ottonis, 561 paraguaensis, 561, 561 Schilleria, 705 aequalis, 714

caudata, 721 marginata, 721 Schlechterella, 549 aurantiaca, 550 Schoepfia, 182 brasiliensis, 183, 184 clarkii, 184 lucida, 183, 184 tepuiensis, 183, 184 Schomburgkia, 561 crispa, 562 heidii, 562, 563 marginata, 562 rosea, 562 undulata, 563 weberbaueriana, 562 Scuticaria, 564 steelei, 564, 565 Se-cun-wa-rei-yek, 725 Seguieria, 678 aculeata, 678, 679 cordata, 680 macrophylla, 679, 680 Sekum-warei-yek, 718 Sekún-waráy, 725 Selenipedium, 564 kaieteurum, 499 palmifolium, 566 steyermarkii, 565, 566 Serapias caravata, 313 Serrastylis modesta, 420 Sesamum, 668 indicum, 668 orientale, 668, 670 Shakrakarimi, 643, 653 Sierrito, 134 Sievekingia, 566 jenmanii, 566, 567 Sigmatostalix, 567 amazonica, 568 huebneri, 568, 568 Siphoneugena, 94 densiflora, 95 var. salicifolia, 95 var. tepuiensis, 95 dussii, 95, 95 var. salicifolia, 95 Sobralia, 568, 573 biflora, 573 candida, 571 cattleya, 571

763

dichotoma, 571 elisabethae, 572 fimbriata, 571, 574 fragrans, 571 granitica, 572 infundibuligera, 572, 575 liliastrum, 572, 575 var. rosea, 572 macrophylla, 572 oliva-estevae, 572, 576 sessilis, 573, 574, 574 speciosa, 573 stenophylla, 573, 575 suaveolens, 573 valida, 573, 576 violacea, 573, 574 yauaperyensis, 573, 574 Solenidium, 577 lunatum, 577, 577 Spathiger rigidus, 342 Specklinia, 503 aristata, 512 grobyi, 516 miqueliana, 518 orbicularis, 601 parvifolia, 519 picta, 519 samacensis, 520 seriata, 521 spiculifera, 521 vittariifolia, 522 Spiranthes, 291, 307, 316, 363, 385, 418, 455, 495, 550, 552 aphylla, 553 bifida, 386 callifera, 496 cranichoides, 291 elata, 291 grisebachii, 418 lanceolata, 551 laxa, 496 maculata, 496 metallica, 553 orchioides, 551 scopulariae, 308 simplex, 553 Stanhopea, 577 aurea, 579 candida, 578, 579 eburnea, 579 grandiflora, 579

764

I NDEX

[Stanhopea] randii, 579 venusta, 579 wardii, 579 Stanhopeastrum, 577 Steffensia, 705 adunca, 714 anonifolia, 715 consanguinea, 717 eximia, 717 hirsuta, 720 luschnathiana, 716 obliqua, 722 pseudochurumayu, 718 Stelis, 579, 583, 584, 585 alata, 583 aprica, 587 argentata, 584 aviceps, 584 bangii, 584 braccata, 586 cucullata, 584 cupuligera, 584 effusa, 585, 587 fendleri, 585 floribunda, 514 foliosa, 532 fraterna, 585 galeata, 515 garayi, 585 grossilabris, 587 guianensis, 585 imraei, 516 intermedia, 585 latisepala, 585, 588 maxima, 585 minimiflora, 586 moritzii, 518 obliquipetala, 408 obovata, 586 ophioglossoides, 586, 588 ovalifolia, 501 papaquerensis, 585 pusilla, 586 pygmaea, 586, 588 santiagoensis, 586 schomburgkii, 587 tenuilabris, 583 tolimensis, 587 trichorrachis, 586 tridentata, 587 trigoniflora, 586 tristyla, 584

velaticaulis, 522 vulgaris, 514 zonata, 587, 588 Stellilabium, 589 alticolum, 589, 589 Stenia, 590 pallida, 590, 590 tigrina, 275 Stenocalyx, 17 patrisii, 26 Stenocoryne, 239 longicornis, 240 Stenopolon, 590 Stenoptera, 363, 369 adnata, 369 costaricensis, 369 roehlii, 363 Stenorrhynchos, 363, 550 lanceolatum, 551 laxus, 496 Stictophyllorchis, 590 pygmaea, 591, 591 Stictophyllum, 590 pygmaeum, 591 Stilifolium, 592 Sturmia elliptica, 412 kappleri, 412 Synanthes, 363 Syzygium, 96 cumini, 96, 97 jambos, 97, 98 malaccense, 97, 98 —T— Tacsonia spinosa, 657 Taribara-yek, 60 Terciopelo, 9 Tetragamestus, 555 modestus, 558 Teuscheria, 591 venezuelana, 592 wageneri, 592, 592 Thouinea, 186 Todaroa, 256 Torrubia, 104 cepalantha, 106 cuspidata, 106 ferruginea, 106 flagrans, 107 microphylla, 107

rusbyana, 109 Tostón, 102 Trachelosiphon, 363 Trébol, 624 Trébol sabanero, 624 Triaristella, 606 triglochin, 607 Triaristellina, 606 triglochin, 607 Trichocentrum, 592 carthagenense, 594, 595 cebolleta, 594 cornucopiae, 594 fuscum, 594, 595 hartii, 594 lanceanum, 596 nanum, 595, 596 Trichopilia, 408, 596 aenigma, 597 albida, 597 fragrans, 597 mutica, 409 oicophylax, 597, 597 Trichosalpinx, 598 cedralensis, 600 dura, 600, 602 egleri, 600, 602 foliata, 600 greenwoodiana, 601 intricata, 600 memor, 600 minutipetala, 601 nageliana, 601 obliquipetala, 408 orbicularis, 601 oxychilos, 601 patula, 602 pumila, 602, 602 roraimensis, 602, 603 steyermarkii, 603 zephyrina, 523 Trichostigma, 680 octandrum, 680, 680 Trichovaselia, 129 canescens, 129 Trigonidium, 603 acuminatum, 604, 605 obtusum, 604, 605 tenue, 605 Trinitaria, 104 Tripa de pollo, 625 Triphora, 605 surinamensis, 606, 606

I NDEX

Trisetella, 606 huebneri, 607 triglochin, 607, 607 Trizeuxis, 607 falcata, 608, 608 pygmaea, 591 Tu‘ kinke me‘n, 86 Tue-becok, 177 tuHaemaereka hu, 645 Tulexis, 248 Tupurú ke tipeñ mën, 62 Turek, 82 Tyleria, 157 apiculata, 158 aristata, 158 breweriana, 158 floribunda, 158, 159 grandiflora, 158, 159 linearis, 158 pendula, 158 phelpsiana, 158 silvana, 158 spathulata, 158, 159 spectabilis, 158 terrae-humilis, 158 tremuloidea, 160, 160 —U— Ugni, 98 angustifolia, 98 myricoides, 98, 99 f. bifaria, 98 f. grossa, 99 f. oligandra, 99 f. stenophylla, 98 roraimensis, 98 stenophylla, 98 vaccinioides, 98 Uleiorchis, 608 cogniauxiana

f. maior, 609 liesneri, 609, 609 ulaei, 609, 609 Urana-u-yek, 181 Uraroene, 59 Uraurek-boni-yek, 28 Uropedium, 498 —V— Vainillón, 612 Vanilla, 609 ensifolia, 611 gardneri, 610 inodora, 610 mexicana, 610 odorata, 610, 611 palmarum, 612 penicillata, 611, 612 planifolia, 611, 612 pompona, 612 wrightii, 612 Vanillophora, 609 Vargasiella, 612 venezuelana, 613, 613 Vaselia, 129 quinqueloba, 130 Veyretia, 552 aphylla, 553 simplex, 553 Villouratea spiciformis, 140 —W— Wallacea, 160 insignis, 161 subsp. insignis, 161 multiflora, 161, 161 Wullschlaegelia, 613 calcarata, 614, 614 ulaei, 609

—X— Xerorchis, 614 amazonica, 615, 615 trichorhiza, 615 Ximenia, 184 americana, 185 var. americana, 185, 186 Xylobium, 616 colleyi, 616, 617 pallidiflorum, 616 steyermarkii, 241 —Y— Ya’ra worokopinyo, 23 Yaguaza, 168 Yamú, 96 Yerba salvaje, 694 Yolanda, 243 Yucú, 62 —Z— Zygopetalum brachystalix, 491 burkei, 368 jorisianum, 368 lindeniae, 618 paludosum, 491 tatei, 618 tricolor, 401 venustum, 491 Zygosepalum, 617 angustilabium, 618 labiosum, 618, 618 lindeniae, 618, 619 tatei, 618, 619 var. angustilabium, 618

765