CHECKLIST OF THE PLANTS OF THE GUIANAS

(Guyana, Surinam, French Guiana)

2nd Edition

J. Boggan, V. Funk, C. KelloffSmithsonian Institution

and

M. Hoff, G. Cremers, C. FeuilletORSTOM

Prepared under the auspices of the Centre for the Study of Biological DiversityUniversity of Guyana, Georgetown, Guyana

CHECKLIST OF THE PLANTS OF THE GUIANAS

(Guyana, Surinam, French Guiana)

2nd Edition

J. Boggan, V. Funk, C. KelloffBiological Diversity of the Guianas

Department of BotanyNational Museum of Natural History

Smithsonian InstitutionWashington, D.C. 20560 U.S.A.

and

M. Hoff, G. Cremers, C. FeuilletHerbarium, ORSTOM97323 Cayenne Cedex

French Guiana, FRANCE

1997

Prepared under the auspices of the Centre for the Study of Biological DiversityUniversity of Guyana, Georgetown, Guyana

CHECKLIST OF THE PLANTS OF GUIANA(Guyana, Surinam, French Guiana)2nd Edition

1st Edition: December 19922nd Edition: February 1997

Published by the Biological Diversity of the Guianas Program, Department ofBotany, National Museum of Natural History, Smithsonian Institution,Washington, D.C. 20560 U.S.A. This is publication number 30 in theSmithsonian's Biological Diversity of the Guianas Program publication series.

Copies available at no charge while supplies last.

Printed in the U.S.A.This publication has been printed on acid free paper

Cover illustration: Phenakospermum guyannense (L.C. Rich.) Endl. ex Miq. (Strelitziaceae), drawn by Alice Tangerini, Smithsonian Institution.

CONTENTS

INTRODUCTION 1 Geology 2 Vegetation types 2 Some current programs and projects in the Guianas 5 Explanation of checklist and editorial notes 5 Acknowledgements 7 References 8CHECKLIST 9 Chlorophyta-Characeae (stoneworts) 9 Hepatophyta (liverworts) 9 Anthocerophyta (hornworts) 15 Bryophyta (mosses) 15 Psilotophyta (psilopsids) 22 Lycophyta (lycophytes) 22 Pterophyta (ferns) 24 Spermatophyta (seed plants) 40 INDEX 211

APPENDIX A. Contributors and Reviewers 229 APPENDIX B. Contributors and reviewers for the revised edition 237

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PREFACE TO THE REVISED EDITION

by John Boggan

Since its publication in 1992, numerousadditions and corrections have been made to theChecklist of the Plants of the Guianas. Wehave incorporated these changes in a revisededition, along with some changes in the formatthat we hope will make the checklist easier touse. Additions have included newly-describedspecies, new records for the Guianas, and namesoverlooked or otherwise omitted from the firstedition. Synonymy and reference lists have beenexpanded. In several cases (e.g., Nyctaginaceae,Turneraceae), the species list for a family hasbeen completely revised. All the spermatophytes(seed plants) are arranged together alphabeticallyby family. The country distribution codes havebeen changed to GU, SU, and FG for Guyana,Surinam, and French Guiana, respectively, inanticipation of a checklist including the"Guayanan" portions of Venezuela and Brasil.

The treatment of the various families remainsuneven, with some families well-known (eitherwith recent monographic or floristic treatments,or with a specialist actively identifyingspecimens) and others (e.g., Dioscoreaceae,Smilacaceae) very poorly known. All availablepublications dealing with Guianan plants sincepublication of the first edition have beenconsulted, in some cases resulting in substantialchanges; in other cases, older publications notseen for the original checklist were consultedwith the result that certain families are in muchmore complete (e.g., Liliaceae s.l. but excludingSmilacaceae).

We have tried not to include any unpublishednames or combinations without indicating themas such (i.e., "sp. nov. ined." or "comb. nov.ined."). However, it has not been possible toverify every name found on herbariumspecimens or in the literature as validlypublished. The editors regret and take fullresponsibility for the use of any invalid names.

The first edition of the "Checklist of thePlants of the Guianas" was a collaborativeproject, with contributions by many researchersworldwide. Authors of the family lists wereidentified as "contributed by...." Otherwise,family lists were compiled by the editors, and insome cases these were reviewed by additionalspecialists (identified as "reviewed by....").Additions and corrections suggested by thereviewers were incorporated. Authorship of therevised edition is complicated by the fact thatadditions and corrections were made by theeditors, by most (but not all) of the originalcontributors and reviewers (listed in AppendixA), and by new contributors and reviewers(listed in Appendix B). The editors have tried toclearly indicate the authorship of each familylist, and hope we have given sufficient credit tothe contributors.

It is the nature of publications such as this tobe obsolete as soon as they are published.During the time between the completion ofediting and final publication, several new specieshave been published, and new country recordshave been discovered. In addition, numerousadditions and corrections were received from JanC. Lindeman (U) shortly before going to press.Unfortunately, there was not enough time toincorporate most of these in the revisedchecklist. The majority of those not alreadyreceived from other sources were synonymswhose current status could not be verified. Newcountry records indicated by Lindeman(primarily for Surinam) are indicated with aquestion mark.

This is publication number 30 in theSmithsonian's Biological Diversity of theGuianas Program publication series. Thispublication was supported by the BDG programand copies are available from the program at nocharge while supplies last.

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INTRODUCTION

by V.A. Funk

This checklist is a preliminary listing of theknown vascular plants (flowering plants,gnetophytes, cycads, conifers, ferns and "fernallies") and "bryophytes" (mosses, hornworts,liverworts) from the Guianas. Although ourknowledge of the algae is insufficient to enableus to produce a species list at this time, we haveincluded the Characeae (stoneworts) of themacroalgae because the information was easilyavailable. This checklist excludes lichens andfungi, which are usually classified togetherunder fungi and are not plants in the strict sense.A checklist of the lichens reported from theGuianas before 1987 has been published(Hekking & Sipman 1988) and hopefully, a listof all the fungal groups will be forthcoming.

This checklist is an attempt to list all of thevascular plants and bryophytes of the Guianas inan effort to encourage further research.Checklists have many uses. They can aid inidentifications, although new species and newrecords are always a possibility. In addition,checklists tell us how many organisms of acertain kind are in a particular area, thus givingus an idea of the species richness. Checklistscan act as an indicator of endemicity within anarea and provide information on the number ofintroduced species. They can be used in acomparative manner between and among areas.If they cover a natural area (one that has biotic orgeologic significance) they can be used toformulate questions about centers of origin andevolutionary history. In addition, a checklist is away to help standardize spelling and authors andto promote accurate labeling of specimens.Checklists, however, reflect our knowledge at aparticular time and our current taxonomicopinions and so their usefulness is tempered bythe knowledge that they are never entirelycorrect and that the list will be constantlychanging. For instance, we are sure that this listcontains incorrect synonomies, records based onmisidentifications, that additional species remain

to be added, and that many species have not yetbeen described or even discovered. However,this type of "snapshot" is valuable because itprovides a way of focusing and stimulating workin the area. In order to help us keep thischecklist as current and accurate as possible,please contact the Director, Biological Diversityof the Guianas Program, with any additions orcorrections.

The part of the northeastern shoulder ofSouth America along the Atlantic coastcommonly known as the "Guianas" consists ofthree political areas (Map 1) with a total area ofabout 500,000 sq. km. The westernmostpolitical entity is Guyana, formerly known asBritish Guiana, which has 800,000 inhabitantsand an area of 231,800 sq. km. To the east ofGuyana is Surinam, formerly known as DutchGuiana, with 400,000 inhabitants and 173,840sq. km. The easternmost is French Guiana, anoverseas department of France, with 120,000inhabitants and 88,240 sq. km. (areas fromMcConnell and Choubert 1975). In all three ofthese areas, the population is concentrated alongthe coast and near the respective capital cities.

The Guianas constitute only part of aphytogeographic or natural area called theGuiana Shield (see Geology section). To makethem a natural area would require includingVenezuela south and east of the Orinoco Riverand small parts of Colombia and Brazil. Duringthe next year the "Flora of the VenezuelanGuayana" project of the Missouri BotanicalGarden will produce a checklist for theVenezuelan Guayana. Proposed collaborationbetween the Smithsonian and Missouri willproduce a combined checklist covering the bulkof the Guiana Shield a natural area to study anddetermine if the flora of the Guiana shield is aunique evolutionary unit.

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For this checklist, we have recorded over9200 species from the three Guianas:approximately 6500 from Guyana, 5100 fromSurinam, and 5400 from French Guiana. Ofthese, only about 3% are introduced andnaturalized. The families with the largestnumber of species in the Guianas are theLeguminosae (Cronquist, 1981, divides thelegumes into three segregate families but theyare often treated as one) with more than 800species and the Orchidaceae with about 700.Other large families are the Rubiaceae and thePoaceae (grasses), both with more than 400species.

GEOLOGY

Underlying all of northeastern SouthAmerica is the Guiana Shield (Map 2), whichoccupies a broad area between the AtlanticOcean and the Orinoco and Amazon rivers. Inaddition to the three Guianas, the Guiana Shieldcovers a small part of eastern Colombia and theVenezuelan departments of Amazonas andBolivar to the west and parts of the Braziliandepartments of Roraima, Pará and Amapá to thesouth and east. Geologically, the Guiana Shieldis an ancient Precambrian land mass (4 billion -590 million years old) made up of variedformations of sedimentary and igneous originthat were metamorphosed and folded(McConnell and Choubert 1975). Overlying thecentral portion of the Shield are the Roraimasediments that, although lacking in fossils, arebelieved to have been laid down on the GuianaShield during the Cretaceous Period (140 - 68million years ago) probably as shallow marine orbrackish water deposits (Gansser 1954, Maguire1970). The Roraima formation consists of pink,yellow and white sandstones, red quartzites,green, black and red shales, conglomerates andboulder beds (Fanshawe 1952). At an earliertime, the Roraima formation must have beenconsiderably larger, perhaps extending south tothe Matto Grosso highlands in Brazil (Fanshawe1952, Gansser 1954). Erosion decreased the sizeof the formation and the remaining sedimentsextend west from the Kaieteur escarpment incentral Guyana, forming the Pakaraima

Mountains, and on through parts of Venezuelaand Colombia and south into northern Brazil(Leechman 1913). Within this area, erosion hascreated vertical-walled, more or less flat-toppedpeaks called "tepui". These table-like formationsare often virtually inaccessible and are sounusual that in a fictional account of a scientificexpedition, one of them was referred to as "TheLost World" (Doyle 1912). Their unique floraand fauna has resulted in the descriptor "lostworld" being applied to all of these tablemountains. The highest peaks of the Guianas arefound in the Roraima Formation in westernGuyana with Mt. Roraima reaching 2772 m., Mt.Ayanganna to 2134 m., and Mt. Wokumung to2042 m.

The Guiana Shield can be divided into fournatural geologic regions: 1) coastal plain; 2)central pediplains and rugged mountains ofPrecambrian metamorphic and granitic rocks; 3)high plateaus of the Pakaraima Mountains,formed by the tabular Roraima Formation; and4) southern pediplains descending to the RioNegro and the Amazon basin.

VEGETATION TYPES

The vegetation of the Guianas is determinedby the underlying geology, altitude, and annualamount of rainfall. All of the Guianasexperience at least one dry season a year andduring most years, two. As with most tropicalcountries, the rains sometimes come early orsometimes late, and occasionally a dry or rainyseason does not occur at all. In general, there issufficient rainfall for most of the Guiana area tobe forested. The estimated percentage of forestcover varies from 86% for Guyana (Haman andWood 1928) to 97.7% for French Guiana(Granville 1974). The Guianas have aremarkable diversity of organisms and their richflora and fauna remain largely unexplored andunexploited. They constitute one of the fewtropical areas worldwide that still has themajority of its forests intact. Estimates vary, butclearly only a small percentage of the Guianashas been deforested.

There are nine vegetation types in theGuianas according to Lindeman and Mori

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Map 1: The Guianas

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Map 2: The Guiana Shield (McConnell & Choubert 1975)

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(1989): 1. seasonal evergreen forest (orseasonal wet forest); 2. mangrove; 3. strandvegetation (beach vegetation); 4. marsh forests;5. swamp forests; 6. herbaceous swamps; 7.savannas; 8. montane vegetation; and 9.inselbergs (granitic outcrops). However,Lindeman and Mori's list focuses mainly onSurinam and French Guiana. Fanshawe (1952)lists 28 vegetation types in six categories forGuyana: 1. rain forest; 2. seasonal forest; 3. dryevergreen forest; 4. montane forest; 5. marshforest (includes savannas); and 6. swamp forest(includes mangrove and herbaceous swampstypes). Neither list covers all of the vegetationtypes with the Lindeman and Mori list lackingseveral forest types and the Fanshawe listmissing the strand and inselberg vegetation.Combining the two lists gives a broader, morecomplete range of vegetation types found in thisregion. One possible way to combine the twolists is to use the 28 types of Fanshawe's and addthe two missing categories from Lindeman andMori.

SOME CURRENT PROGRAMSAND PROJECTS IN THE

GUIANAS

Several programs and projects are nowunderway in the Guianas. A few that deal withfloras are listed below. The first twocollaborated to produce this checklist.

Biological Diversity of the Guianas (BDG) -Smithsonian Institution. The BDG started in1983 and now operates from the "Centre for theStudy of Biological Diversity" on the campus ofthe University of Guyana. The BDG programseeks to document and study the flora and faunaof the Guianas. Activities include collectingspecimens to be housed at the Centre andtraining students and staff of the University aswell as producing checklists, flora treatments,inventories, vegetation maps, and otherpublications such as a listing of the plants andanimals of Kaieteur Falls National Park inGuyana.

Herbarium of Cayenne - ORSTOM. Thegeneral herbarium was founded in 1965 by R. A.A. Oldeman to succeed the savanna (grass)

collection established by J. Hoock between 1955and 1965. From the beginning, the activity ofthe laboratory focused on the floristic andecological studies of the forests of FrenchGuiana. Activities in Cayenne include collectingspecimens in remote areas of the country,especially inselbergs, and producing treatmentsfor flora projects. In 1988, a checklist of theFlowering Plants and Pteridophytes of FrenchGuiana was published (Cremers et al. 1988).The herbarium of some 60,000 specimens isstored electronically (completed in 1988) in thedata bank AUBLET.

Flora of the Guianas. An internationalconsortium of nine botanical institutions formedin 1983 to produce a written account of theplants of the Guianas. The Flora project has itseditorial center at the University of Utrecht, TheNetherlands. Some of the contributors to thischecklist are also participating in the Flora of theGuianas project; however, this checklist is notpart of the Flora nor is it associated with it in anyway.

Flora of Central French Guiana. A jointproject by the New York Botanical Garden andORSTOM to produce a flora of 50,000 hectaresof rainforest near Saül, French Guiana.

Iwokrama International RainforestProgramme. Located in the center of Guyana,the principal aim of the Iwokrama Programme isto demonstrate, through scientific research,methods of sustainable utilization of the forestfor timber and other extractive resources whileconserving biological diversity.

Conservation International. Operates a"Guianas Regional Program" (consisting ofSurinam and Guyana) to assist these countries inconservation matters and to conductethnobotanical studies.

EXPLANATION OF CHECKLISTAND EDITORIAL NOTES

The checklist includes the vascular plantsand bryophytes of Guyana, Surinam and FrenchGuiana. Although the checklist does not covermost algae, the Characeae are included. Thelichens and fungi are not included. As thischecklist is preliminary, the quality and coverage

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of the families varies widely depending on theavailability of recent floristic and/ormonographic literature and the participation ofspecialists. However, contributing specialistssometimes have supplied data that could not beverified from the literature. At this point it isn'talways clear what is new and/or unpublishedinformation and what is based on literature theeditors have not seen.

Family circumscriptions of the floweringplants follow Cronquist (1981), except ourrecognition of the family Bonnetiaceae on therecommendation of A. Weitzman (personalcommunication). Pteridophytes follow theclassification in the United States NationalHerbarium (US), as arranged by D. Lellinger.Other groups are as arranged by their respectivecontributors.

Author abbreviations, when used, followStafleu and Cowan (1976-1988). "H.B.K." iscited throughout the checklist as "Kunth".

A few taxon names are included in the listthat we have been unable to verify. Some maybe new records, most are probablymisidentifications or synonyms and still othersare names from the literature that the editorscannot verify. The "unverified names" were notcontributed by the specialists.

Sources of Information: Several sourceswere used for the data presented in the checklist:1) the literature: see references at the end of eachfamily and in the General Reference section; 2)contributions by specialists, either as providedby an individual and therefore listed as"contributed by..." or a list compiled by theeditors was reviewed or proofread by a specialistin that group, in which case it is listed as"reviewed by...."; 3) databases: CentreTechnique Forestier Tropical - France,Herbarium of Cayenne - ORSTOM, BiologicalDiversity of the Guianas Program - SmithsonianInstitution; and 4) herbarium records, i.e.,specimens at: BRG, CAY, US, MO. Allabbreviations for herbaria follow Holmgren et al.(1990). Families listed without contributors orreviewers were compiled by the editors.

The bibliographies are not exhaustive. Onlyrecent monographic and floristic literature(generally after 1900) relevant to the covered

geographic area was surveyed (see referencesfollowing the introduction and with eachfamily). Family references have been added bythe editors as an aid to the user of the checklist.Families without corresponding specialists havemore extensive literature citations. If a familyhad a recent treatment (e.g., in the Flora of theGuianas (Görts-van Rijn 1985 - ), Flora ofSuriname (Pulle et al. 1932-1984), Botany of theGuayana Highland (Maguire and Collaborators1953-1989), Flora Neotropica, or familymonographs) then little additional work wasdone for the present list.

In a few cases (Cyperaceae, Myrtaceae, someparts of Orchidaceae), separate lists wereprovided by more than one specialist. In all suchcases, the lists were not 100% in agreement andthe list presented is an amalgamation of the two.In the species lists in general, and thesynonymies in particular (see below), it shouldbe noted that different systematists will havedifferent opinions on the status of particularnames. Taxa collected on Mt. Roraima have caused aproblem because this table-mountain is locatedon the borders of Guyana, Venezuela and Brazil.Specimens from this area may lack a countrydesignation or it may be incorrect or unknown,especially in historical collections. To preventconfusion, taxa known in the Guianas only fromMt. Roraima have been listed as "Mt. Roraima"rather than Guyana. However, as we continue toexplore the Pakaraima Mountains, some taxapreviously known only from Mt. Roraima mayalso be found in Guyana.

Included and excluded names: This checklistincludes indigenous taxa and naturalized taxa (sonoted when possible) as well as taxa fromneighboring areas that may be expected in theGuianas. Infraspecific taxa at the level ofsubspecies and variety are included at thediscretion of the contributors. Infraspecific taxaat the level of "forma" are excluded as well asnames of varietal status not currently consideredto be distinct from the "typical" variety.

We have excluded taxa known only incultivation in the Guianas. Although some ofthese may be found as escapees, they are

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generally not well-established. DeFilipps (1992)has listed many of the cultivated plants in hisrecent publication.

Synonymies and taxa not known in theGuianas: The synonymy is not meant to beexhaustive. We have included synonyms only ifthe synonymized name is found in recentliterature. Because some of the synonymies aretaken from the literature they may not allconform to the taxonomic concepts of thecontributing specialist for that family. In many cases, a species has been reportedfrom the Guianas either because of a broadspecies circumscription that has since beenrestricted (or as an infraspecific taxon that hassince been raised to species status), or due tomisidentifications or other errors. In either case,the name is listed at the end of the family withthe notation "not in Guianas".

ACKNOWLEDGMENTS

Once again, we thank the many specialistswho have contributed to this checklist. It wouldnot have been possible without their assistance.Their names are listed at the beginning of thefamily list with which they assisted as well as inAppendix A. Sally Adkins, Keitha Dattilo,Susan Grose, Marilyn Hansel, Tom Hollowell,Mark Strong, and Fernanda Zermoglio assistedwith the proofreading and final preparation ofthe manuscript. We also thank Robert A.DeFilipps and Jan C. Lindeman for theirnumerous comments on the entire checklist. Wehave received comments, corrections, andencouragement from many sources, and regretthat we can not name them all. In addition wethank D. B. Lellinger and L. E. Skog for beingespecially helpful with the overall project

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REFERENCES

Cremers, G., C. Feuillet, J.J. de Granville, & M.Hoff. 1988. Liste des espèces dePhanérogames et de Ptéridophytes deGuyane française d'après l'Herbier duCentre ORSTOM de Cayenne. ORSTOM,Cayenne.

Cronquist, A. 1981. An Integrated System ofClassification of Flowering Plants.Columbia University Press, New York.

DeFilipps, R.A. 1992. Ornamental GardenPlants of the Guianas: An historicalperspective of selected garden plants fromGuyana, Surinam and French Guiana.Department of Botany, SmithsonianInstitution, Washington, DC.

Doyle, A.C. 1912. The Lost World. PuffinBooks, London.

Fanshawe, D.B. 1952. The vegetation of BritishGuiana: A preliminary review. ImperialForestry Institute paper no. 29: 1-96.

Gansser, A. 1954. The Guiana Shield (SouthAmerica). Eclog. Gelo. Helv. 47: 77-117.

Görts-van Rijn, A.R.A. [ed.]. 1985- . Flora ofthe Guianas. Koeltz Scientific Books,Germany.

Granville, J.J. de. 1974. Paysages végétaux dela Guyane. Pages 77-78 in Regards sur laFrance. Editions S.P.E.I., Paris.

Haman, M. & B.R. Wood. 1928. The forests ofBritish Guiana. Trop. Woods 15: 2-13.

Hekking, W.H.A. & H.J.M. Sipman. 1988. Thelichens reported from the Guianas before1987. Willdenowia 17: 193-228.

Holmgren, P.K., N.H. Holmgren, & L.C.Barnett. 1990. Index Herbariorum Part I:The Herbaria of the World. InternationalAssociation for Plant Taxonomy.

Leechman, A. 1913. The British GuianaHandbook. "The Argosy" Co., Ltd.,Georgetown, British Guiana and Dulau &Co., London.

Lindeman, J.C. & S.A. Mori. 1989. TheGuianas. In D.G. Campbell and H.D.Hammond [ed.], Floristic Inventory of

Tropical Countries. New York BotanicalGarden, New York.

Maguire, B. 1970. On the flora of the GuayanaHighland. Biotropica 2: 85-100.

Maguire, B. & collaborators. 1953-1989.Botany of the Guayana Highland. Parts I-XIII. Mem. New York Bot. Gard.

McConnell, R.B. & B. Choubert. 1975. GuianaShield-Regional Survey. In R.W. Fairbridge[ed.], Encyclopedia of World RegionalGeology, Part I: Western Hemisphere.Dowden, Hutchinson, & Ross, Inc.,Stroudsburg, PA.

Pulle, A., J. Lanjouw, A.L. Stoffers, & J.C.Lindeman. 1932-1984. Flora of Suriname.J.H. deBussy, Amsterdam; E.J. Brill, Leiden.

Stafleu, F.A. & R.S. Cowan. 1976-1988.Taxonomic Literature (second edition).Bohn, Scheltema, & Holkema, Utrecht.