The vegetation of the Antilles - Howard 1973 by JPZ.

VEGETATION-AND VEG-ETATIONAL HISTORY OF NORTH ERN LATIN AMERICA PAPERS PRESENTED AS PART OF A SYM POSIUM, " VEGETATION AND VEGETAT IONAL HISTORY IN NORTH ERN LAT IN AMERICA", AT TH E AMERI CAN INSTIT UTE OF BIOLOGICAL SCIENCES MEETJN G,S. BLOOMINGTON, IN D. (U.S.A.), 1970

edited by

ALAN GRAHAM Departmelll of Biological Sciences, Ken/ State University, Kent, Oflio (U.S .A.)

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ELSEVIER SCIENTIFIC PUBLISHING COMPANY Amsterdam - London- New York

. 1973

ELSEVIER SC IENTIFIC I'UllLISHING COMPANY 335 JAN VAN GA LENSTRAAT P.O. BOX 2 11 , AMSTERDAM , THE NETHERLANDS AMERICAN ELSEVIER PU BLISHING COMPANY, INC. 52 YANDERJ31LT AVENUE NEW YORK, NEW YORK 10017

Library of Congress Card Number: 72-87955 ISBN: 0-444-41056-2 With 91 illustrations and 53 tables CopyrightÂŠ 1973 by Elsevier Scientific Publishi.n g Company, Amsterdam All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior written permission of the publisher, Elsevier Scientific Publishing Company, Jan van Galenstraat 335, Amsterdam

PI\INTED IN TH E NETHERLANDS

Preface

The papers in th is vol ume were presented as part of a symposium, "Vegetatio n 11nd Vegetational H istory in Northern Lati n America", at the American Institute of Biological Sciences meeti ngs, Bloomington, I ndiana (1970). The authors and editor wish ~o express appreciation to the Ecological Society of America, Paleobotanical Section of the Bo tanical Society of America, and the American Society of Plant Taxonomists for sponso ring the symposium. This volume follows an earlier one on Floristics and Pafeojforistics of Asia and Eastem North America-a collection of papers presented in conjunction with the XI Intemational Bo ta nica l Congress, Seat.tle. The phrases "Floristics and Paleofloristics" and "Vegetation and Vegetational History" reflect an organ izational concept followed in both symposia and reflected in these subsequent publicati ons. This concept, briefly stated, is that reliable pa leobotanical data together with taxonomically sou nd floristic studies provide the most reasonable approach to studying the origin and evoh:1tion of modern plant communities. Recent references to the vegetation of major geographic regions (e.g., the Guayama Highland, Mexican dry regions) reflect a commonappre ciationfortheco mplexity of the communities. An analogous view o f individual plant biosystems (viz. , species or species groups) eventually evolved into a philosophical concept of plant systematics succinctly designated in 1945 as the New Systematics- complex biosystems require a multiplicity of approaches for comprehendin g their origin and evolution and for establishing a meaningful taxonomy. A lthough the relation between individual plant biosystems and comm unities of plants is a debated aspect of ecological theory, both are acknowledged as complex at their particul ar quantitative level. If understanding the biology of individual plants or groups of similar plants (viz., species) requires a multiplicity of approaches and techniques (the New Systematics), a compa.rable view sho uld extend to the st udy of communities, i.e., a New Floristics. In fact such an approach has gradually developed in ecological-floristic studies of Latin American communities even tho ugh the underlying modus operandi has not been formally designated as it has in syst ematic botany. A comparison of MacArthur and Wilson's (1967) mathematical models for island communities, Odum et al.'s (1970) studies on the effect of radiation o n Puerto Rican rainforests, and H oward et al.'s (1968 et seq.) ceo-fl oristic studies in Puerto Rico are witness to the diversity of methods used to investigate the biology of tropical plant communities. This volume concerns a comparatively neglected facet of the multi-dimensional method of studying community evolution in northern Latin America- the geologic history and development of the biotas. The thesis for these discussions is that interpretations based on reliable paleobota nical data and on taxonomically sound floristic data are com-

PR EFACE

plementary, a nd co llective ly provide a logical a pproach to the study of community evolution. Reaso ns fo r the slowness in integ rating paleobota nical da ta i n t~ floristic stud ies in La tin America arc several. Until recen tly there was an insullicientnum ber of studies to provide a n adequa te history of vegetation about the Caribbean Basin. As previously noted (G raham and Jarzen, 1969), there have been "more Tertiary paleobotani cal studies made in Ihe western Âˇu nited States than in the rest olthe Western Hemisphere south of the U ni ted States. Indeed there have been more revisions of foss il fl oras of the Colum bia Pla teau than combi ned Tertia ry paleobotanicn l studies in the countries of Guatema la, El Sa lv~tdo r, Honduras, Nicaragua, Costa Rica, Pana ma, the Guianas, Ecuador, Paraguay, Uruguay and Peru" (p. 309). Another deterrent to more effective integration of paleobotanical and modern flo ristic data concerns the reliability of identifica tions based on fragmented fossi l remains. Taxonomists interested in the fossi l record of Carya, fo r example, would find that specimens now designated as C. bendirei (similar to the modern C. ovata) have been described as fou r different species of Salix, two species of Carya, Rhus and Pnmus, and as Magnolia, Cassia, Celastrus, Aesculus, Hicoria, Juglans, Ptelea, Quercus, and Arbutus. T he expla nation for such taxonomic uncertainty is in part the same as that used to explain the tri-generic status of certain tree ferns as Hemitelia, Cyathea (sensu Holtum) and C11emidaria (sensu T ryon), or the assignment of the Lecythidaceae by fo ur recent authors to four different Orders of the plant kingdom. The taxonom ic history of the pa leobotanical example given spans 50 years of time, and in th is interva l considerable advance ment has bee n made in techniq ue and in the availability of comparative materia ls. More meaningfu l assessments of the current potential for coordinated paleobotanical and modern flo ristic studies are ava ilable. In the older literature there are reports of fossi l Com pos ita~ in deposits as old as Cretaceous. Such a record is incongruous considering the advanced evolutionary position of the fam ily. Pollen of the family is distinct, however, and recent extensive palynological studies demonstrate the fami ly began its evolutionary development in the latest Oligocene and does notbecome.common in fossil deposits until the Lower Miocene. A simi lar parallel exists between the recently revised paleobotan ical record of the Lythraceae and taxonomic opinion based on the comparative morphology of modern forms (G raham and Graham, 1971). Among those genera with an adequate fossil reco r~ , the presumed primitive gen us Lagerstroemia ranges from Recent to Early Eocene, while the presumed advanced member of the fam ily, Cuphea, extends back only to the Miocene. Among data from presently unpublished studies there is evidence for relative stability of the tropical flora during long spans of Tertiary time; a progressive southern migration of arboreal temperate elements into the Latin American biota, with Juglans, Alnus, and Myrica reaching northern South America only during latest Pliocene and Pleistocene times; and the existence during the Tertiary in the Isthmian region of a series of islands and channels serving simultaneously as a barrier to the

PR EFACE

VII

migration of large terrest rial vertebrates :md certa in r lants, while facilitating Atlant.ic-Pacific interclu.t nge or marine invertebrates, and providing a migration rou te fo r other plant species vi;1 island stepping-stones. These paleobotan ical data, when eventually refined and documented, will be of usc in interpreting the origin and development of modern Lat in American biotas. T he reconstruction of paleo-e nvironments, paleophysiography, pathways of migration, and former continuity between currently allopatric species and communities depends in tu rn upon reliable ecological, taxonorhic, and floristic data. Collectively these approaches arc beginning to provide a tentative preliminary model for community evolution about the Caribbean Basin. This volume is offered as a statement o f the current status of such studies. ALAN GRAHAM (Kent, Ohio)

References Graham, A. a nd Graham, S. A., 1971. T he geologic his10ry of Ihe Lythraccac. Brittonia, 23:335-346. Graham, A. and Jarzcn, D. M., !969. Stud ie s in neotropical paleobotany, I . The Oligocene comm u nities of Puerto Rico. Ann. Mo. Bot . Card., 56: 308-357. Howard, R. A., 1968. T he ecolog)' of an elfi n forest in Pucno R ico, I. Introd uction and composition studies. J. A mold Arborelllm, 49: 38 1- 4 18. MacArthur, R. H. and Wilson, E. 0 ., 1967. The Theory of Island Biogeography. PrincCIOil Univ. Press, Princeton, N.J., 203 pp. Odum, H . T. (Editor), 1970. A Tropical Rainforest: A Study of frradiation a11d Ecology al Ef Verde, Puerlo Rico. Div. Tech. Inform., U.S. Atomic Energy Comm, Oak Ridge, Tenn., 1066 pp.

Contents Preface . . . . . . . . . . . . . . . . . .

Chapter 1. The Vegetation of the Antilles RICHARD A. HOWARD Summary . . . . . . . . l ntroduct iOJl . . . . . . . Vegetation types in the Antilles Coastal formations . Lowland formations Montane forma tions Qrigins of the existing vegetation of the Antilles Relationships and distribution of the Antillean vegetation Disjunct distribution patterns in the Antilles . . . . . Distribut ion patterns wit hin the islands . . Long-distance 1elationships and disjunct distribution patterns Conclusions References . .

5 8 12 16 17

29 31 34 37 37

Chapter 2. A-Paleoclimatic Interpretat ion of the Eocene Floras of Southeastern

North America DAVID

D ILCHER

Summary . . . . . Introduction . . . Individual components of the fl ora Identification to modern taxa Evolution of the fossil taxa . Relation of range and ecological tolerance. Community structure . . . . . . . . . . . . Stability through time . . . . . . . . . Separation into equivalent modern communities Foliar physiognomy . . . . Leaf-margin analysis . . . . . . . . Leaf-size analysis . . . . . . . . . Interpretation of foliar physiognomy . Application to this nora Additional evidence Wood . . . . . . . Pollen . . . . . . . Paleoclimatic interpretations References . .

40 41 4t 42

43 44 44 45

46 48

50 53 54 55 55 55 56 57

Chapter 3. G eographical Relationships ofthe Flora of Mexican Dry Regions J. RZEDOWSKI Suillmary . Introduction

61 61

f X

CONTENTS

CONT ENTS

Relationships between diOcrent arid zones of Mexico Relationships with other arid zones of the world Relationships with zones of non-a rid climate . Conclusions References . . . . . . . . . . . : . â&#x20AC;˘ .

63 1;6'

70 71

Clrapter 4. Ecology of the Vegeta tion of Veracruz ARTURO G6MEZ-POMI'A

Summary . . . . Introduction . . . Physical condi lions Topography Climate Soils . Vegetation . . Forests Selvas . High sclvas . Low selvas . Grasslands . Arid or semi-arid vegetation Secondary vegetation References . . . . . . . . . . .

73 73 73 73 75 85

89 94 105 106 118

128 135

145 147

Chapter 5. The Phytogeography and Vegetation ofChiapas (Mexico)

149

149 150 154

155 155 158 161 163 164

Chapter 6. The Vegetation of Panama: a Review D UNCAN M. PORTER

Summary . Introduction Climate Soils Man Vegetation .

Vegetation maps . . . Koppen macroclima tes Holdridge Life Zones Human innucnccs . . . Darien forest types . _ . A summary of plant comnumiti<.,-s Incipient Forest . . . _ Early Secondary Forest Late Secondary Forest Liltoral Forest . . . . Thorn Forest _ . . . . Deciduous Seasonal Forest . Savanna .. . . . . _ . S<tltwatcr Swamp Forest . Saltwater Riparian Forest. Brackish Swamp Forest _ llrackish Riparian Forest Freshwater Marsh . . . Freshwater Swamp Forest Frcshw<tter Riparian Forest . Monsoon Forest . . . . Evergreen Seasonal Forest Premontane Rain Forest Cloud Forest Elfin Forest Acknowledgements ~ References . . . .

17 1

171 172 173 174 175

175 177 179

182 183 184 185 187 187 187

189 189 190 192

193 197

197 198 199 199 200

Chapter 7. Phytogeographic History of the Isthmus of Panama during the Past

DENNIS E. B REEDLOVE

Summary . . . . . . . Introduction . . . . . . Phytogeography of Chiapas . Vegetational formations ofChiapas Description of the formations . Optimum forma tions. . . . Seasonal format ions . . . . Wet-land forest formations . Treeless formations References . . . . . . . . . . .

167 167 169 170

170 171

12,000 Years (A History of Vegetation, Clima te, and Sea-level Change) ALEXANDRA S. B ARTLETT AN D ELSO S. BARGHOORN Summary . . . . . . . . . . Introduction . . . . . . . . . Northern South America . Southern Central America Panama . . . . . . . Geological history of Panama .. Pre-Quaternary history . . Late Quaternary and Recent history Physiography' and climate of Panama . Climate of the Canal Zone The present vegetation of Panama Preparation ofsamples . . . . Preparation techniques . Preservation of samples The pollen diagrams . . . . . Theoretical considerations The Gatun Basin diagrams The modern pollen rain . Climatic change in Panama . . . Identification and nomenclature of the fossil pollen .

203 204 .-

204

206 207 208

208 209

211 211

212 216 219

219 2 19 2 19

222 225 228

229

XII

CONTENTS

Nomenclature . . . . . . Vegetation 35,500 B. I'. Vegeta tion 11 ,300-9,600 B.P. Vegetatio n 9,600-7,300 ·1l.P.. Vegetation 7,300-4,200 B. P.. Vegetation 4,200 B.P. to the present Annotated list of species identified as po llen in Panama sediments . Monocotyledonac Dicotyledonae . . . . Other groups . . . . . Sea-level change- the literature Introduction . . . . . Early work Problems in interpreta tion of former sea leve l Sea levcl 35,000-18,000 D.P. . . . . . . . Climatic evidence from continental deposits . Evidence from oceanic cores Sea Jevel24,000-18,000 D.P. . . Sea leveJ· JB,000-10,000 D.P.. . Sea leveiiO,OOO B.P. to present Eustatic sea-level change in Panama . Introduction . . . . . . . . Mangrove pollen as a sea-level indicator. Sea-level data from the Gatun Basin Sea level 35,500 B. P. . . . . . . . .- . Sea level from4,200 B.P. to present . Acknowledgem ents References . . . . . . . . . . . . . .

CONTENTS

23 1 233 235 "239 241 245 249 249 255 268 268 268 269 269 27 1 274

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ALAN GRAHAM

301 301 303 303 308 310 312

Chapter 9. Literature on Vegetationa l History in Latin America ALAN GRAH AM

Summary . . . . Introduction . . . General bibliographic references . Biogeography and his torical geology . Paleobotany of Mexico . . . . . . .

324 325 328 358

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

36 1

277 277

Chapter 8. History of the Arborescent Temperate Element in the Northern L atin American Biota

Summary . . . . . . . . . . . . . . .......... . Introduction . . . . . . . . . . . . . . . . . . Origin of the eastern United S ta tes-eastern Mexican floristic relatio nship. Earlier studies. . . . . . . . . . . . . . . . . . . . . . . New data from the Miocene Pa rajc Solo Formation, Veracruz, Mexico Progressive southward introduction of the arborescent temperate element into the Latin American biota. Refere nces . . . . . . . . . . . . . . . . • .

Pa1cobolany of Cenlral America . Palcobolany of I he Antilles . . . Paleobolany of South America . Selected references o n I he pollen morphology of predominantly neotropica1 taxa .

276 276 282 282 283 290 29 1 293 293 294

315 315 316 316 319

XIII

Clwp/er I

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The Vegetation of the Antilles

RICHARD A . HOWARD Arnold Arboretum, Jamaica Plain, Ma.<s. (U.S.A.)

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Summary The Caribbean islands form an archipelago of over I ,000 islands of considerable range in size, alt it ude, soil types and environmental niches. l'he larger islands oft he Greater Anti lles have the largest number of species a nd the grea test endemism numerically. Especially on Cuba and Hispaniola there are geographic areas of high speciation and these arc largely_areas of mesophytic to xerophytic environments. Although there are many small endemic gen~ra restricted t o one or a few islands, only two genera with twenty s pecies o r more appear to be li mited to the Antilles. One fa mily, Picrodendraceae, is restricted to the Greater Antilles. Patterns of d isjunct distribution are ab undan t and involve the Greater Antilles a nd the lesser Antilles; islands of the Greater Antilles and Central America, and the Greater A nti lles and South America. Examples are ci ted of the relationships of the Antillean flora to Africa and to Asia.

Introduction The islands of the West Indies, the Ant illes, form an archipelago, that is, a series of small land masses scattered in a sheet o f water. In the southern part of t heir distribu tion their orientation north and south separates the Caribbean Sea from the Atlantic Ocean. Exposed land surfaces cover a distance of I ,700 miles from Barbados on the east to t he western tip of Cuba, while the distance from Grenada in the Lesser Antilles to the northern tip of the Bahamas is I ,200 miles. Nearly 1,000 named and vegetated islands comprise the Antilles, ra nging in size from Cuba, with 44,220 sq. miles and a vascula r flora of over 6,000 species, to i:;lets or small rocks of a few square yards and a flora o f a dozen species. Although isolated from each other at sea level, the islands are related below the surface in three prominent sweeping curves revealed· as sea mounts by under-sea exploratio n. The Bahama Islands appear as a continuation of the North American coastal shelf. The deepest point in the Atlantic Ocean, 27,500 ft., occurs in the Puerto Rico trench between the southern exten sion of the Bahama Bank and the Antillean arc. Lines of sea mounts extend south and west from the Greater Antilles, but present information is only speculative as to ·whether these existed in the past as continuous

, 2

R. A. HOWARD

T H E VEGETATION OF THE ANTILLES

land bridges for biological migrations. However, Grenada on the south is sepa rated from the South America n coastal shelf by a narrowj rench and by a sea-level distance of eighty miles. Western Cuba and the Yucatan peninsula are but I J 5 miles apart at sea level. The diversity of the Ant illes is seen in such factors as the altit ude of the islands, the temperature range, the soil types and the units of vegetation on each island. Hispaniola has the grea test rai1ge in altitude where the Enriquillo Basin is 100 ft. below sea level and the highest mountain is reported to be 9,700 ft. (Fig. I). Volcanic peaks in the Lesser Antilles range from 3,000 ft. to approximately 5,000 ft. The major part of

9,'100 8 ,300

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1'vlean average 1empemture

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December

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Havana Nassau

25. 1 25.6 26. 1 24.9 25. 1

1609 1275 1979 11 57 118 1

Grenada (560 m) Martinique (725 m) Guadeloupe (510 m) Puerto Rico ( 101 5 m) .

24.0 23.2 21.3 18.6

4 165 5.273 2630 4533

Dominica

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TJ\ ULE J CLIMATE WITH IN l'II E \VfST INDIES

receives that quantity Gf water, the vegetation of the area suffers a water deficiency which is shown in progressive stages of wilting, leaf fall and eventual death of the plant. Many areas of the Antilles receive less than 40 inches of annual rainfali. Jn fact, there was no rain reco rded fo r a period of 3 years by personnel of the Sea Board Oil Company workin g in the Enriquillo Valley in Hispaniola in the 1950's. Rainfall exceeding 200 inches annual precipitation has been suggested for several areas, but documentation is in freq uent. Many areas of the Antilles show periods of 6 or 7 months of reduced rainfall occurring as two dry periods when part of the vegetation may lose its leaves and show mass flowering. In these areas irrigation is necessa ry for

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3,200~

2,800 0

Fig. !. Altitudinal data on the islands of the Antilles. Diagonal lines indicate areas of less t han 1,000 ft . D otted areas exceed 3,000 ft. Highest mountains are indicated by appropriate numbers in feet.

the land surface within the archipelago is below 1,000 ft. in altitude. Although there are vegetational changes with altitude, there is no timber line within the Antilles and no high-altitude tundra or paramo. The average mean temperatures of the area, presented in Table I, are taken mostly from reports of government weather stations at elevations of less than 100m. Temperature range throughout the year is not great, nor is the daily variation in temperature. Day-length range is nearly two hours greater in Nassau in the north than it is in Trinidad just south of the area under consideration. With reference to the rates of evaporation within the area of the Antilles, the figures of 40 and 48 inches of rainfall have been cited as significant in determining the type and distribution of vegetation in the tropics. According to Beard (1949), the critical rainfall figure for the West Indies is 4 inches per month. Unless an area

~ SILICEOUS

J~11f:;l SALT~CYPS\n-1

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VOLCANIC-SOU!'RI&RE

ALUMINOUS-LATERITIC

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Fig.2. T he distribution of special soil types within the Antilles.

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R. A. HOWARD

T H E VEGETATION OF TH E ANTI L LES

commercial production of sugar cane or bananas. Other areas show no months of rain fall deficiency of less than4 inches, as exemplified by a lowland weather station on Dom inica or mountain stations in Puerto Rico and Martinique. The Antilles show a vari::ty of soil types which offer ecological niches or interesting areas for the plant collector to examine. The vegetat ion on the various soil types is often distinctive in composition or habit and is frequently high in endemic species. The accompanying map, Fig.2, indicates the location of siliceous savannas in western Cuba and on the north coast of Puerto Rico. The central area of Cuba has an interrupted serpent ine savanna dominated in various areas by shrubs or by palms. Areas or gypsum and salt concentration s, often in bands from an exposed syncl inous geological structure, occur in Hispaniola. A belt of alu minous lateritic soil extends through most of the island of Jamaica and occurs also in the southern peninsula of Hispan iola. Areas of volcanic activity occur in the Lesser Anti lles, with eruptive activity in this century on Martinique and St. Vincent. Fumeroles, sulfur vents and boili ng la kes with varying degrees of activity are encountered from Grenada to Montserrat. Outcrops of I imesto ne as sedimenta~y rock or elevated coral reefs are abundant and intrusive areas of igneous rock could be added to the_ chart as additional specialized habitats.

Montane formations lower montane rain forest montane rain forest or cloud forest montane thicket high mountai·n forest elfin woodland or mossy fo rest paramo tundra

Optimum/ormation rain forest Seasonal formations evergreen seasonal forest semi-evergreen seasonal forest deciduous seasonal fores t thorn woodland cactus scrub

deSert

Dry evergreen formations dry rain forest dry evergreen forest dry evergreen woodland and littoral woodland dry evergreen thicket and littoral thicket evergreen bushland and littoral hedge rock pavement vegetation

Sll'amp formations swamp forest and mangrove forest swamp wood Ia nd. swamp thicket herbaceous swamp

Stehle (1945, 1946) considered only the forest types of vegetation in the Lesser Antilles and presented a classification dependent on altitudi nal distribution, ra infall and soil type as the follo wing system . MangrOI'I: marine riparian

Mesophytic calcareous volcanic

Xerophytic · spiny deciduous sandy calcareous volcanic

Hygi'Ophytic marshy humid

Vegetation types in the Antilles

The history of botanical interest in the Antilles is a long one, but it has been devoted primarily to the studies of individual islands from the time of Columbus to the na~onalistic studies of island groups of more recent years. No comprehensive flo ra exists for the Antilles nor is one in preparation. As a result various systems of classification have been proposed for the vegetation of the Antilles or for individ ual islands. An extensive bibliography is given by Stehle (1945, 1946), although the proposals most wi_dely quoted are those of Beard (1944, 1949, 1955). Beard ha s proposed, with Clementsian terminology, a three-part vegetation classifica tion based on: (a) a floristic grouping-the climax association; (b) a physiognomic grouping-the forma tion; and (c) a habitat grouping- the formation-series. The following twenty-eight categories he recognizes are applied to all of tropical America, but not all are represented in the Antillean archipelago.

Seasonul-su·amp formations seasonal-swamp forest seasonal-swamp woodland seasonal-swamp thicket savanna

1 )

•

Altitudinal alluvial volcanic culminating

In order to consider the composition, distri bution and relationships of the Antillean vegetation it is· desi rable to discuss and il lustrate some of the vegetational types. The following modified classification will be followed wh ich utilizes the geograph ic location as well as the ecological fac tors of the area . Coaslal formations beach strand rock pavement mangrove Lowlandformations thorn scrub savanna marsh or swamp alluvium

Montana formations wet or dry forests on li mestone montane sclerophyll palm brakes tree fern communities pine forests cloud for~ts volcanic and soufriere communities crater lakes elfin thickets

Coastal formations

The beaches of the Caribbean area are attractions for the tourist and the botanist alike (Plate I, 1). Undisturbed beach areas are increasingly difficult to find. Sandy beaches, however, are oflimi ted extent on the islands of the Antilles when compared with the rocky or coralline coastal areas. Beaches may be composed of white sand of

R. A. HOWARD PLATE I

I. 2. 3. 4. 5. 6.

Sandy beach and strand vegetation o n Beata Is land, Hist>aniola. Alluvial bay with Avicennia, Balis and Sesuvium, Puerto Rico. Dry-thorn scrub vegetation, Jamaica. Savanna area with Prosopis, Antigua. A limestone cliff or mogote with thorn scrub vegetation, Jamaica. Enriquillo Basin, Hispaniola. Lake Enriquillo with the surface below sea level is in the upper right hand corner . Edges of the syncline are seen with alternating zones of gypsum a nd salt rock outcrops. Thorn scru b vegetation includes areas of pure stands of cacti.

T HE VEGETATION OF T H E ANTILLES

coralline o rigin or black sand of vo lca nic origin . Some islands rnay have on ly one kind of sand beach, while others have li mited amounts of each kind. A spit o n the southern end· o f Gre nada is black sand on one coast and whi te sand on the other. Beach p lants mos t commonly encountered are Canal'a(ia (Legum inosae), Cenclrrus, Distichifis (Gramincae), Euphorbia (E uphorbiaccac), ipomoea (Convolvulaceae), Ses111·ium (Aizoaccac) and Uniofa (Gramincac). Both t he _genera and the species involved arc o f wide geographic distribut io n, and no endemic taxa occur in this zone (Sauer, 1967). Inland from the.coastal beach, dunes may be bu ilt up consisting solely of sand or may be of sand d eposi ted on a rocky substratum (Plate 1, 1). T he sea grape, Coccoloba ut•ifera, is the classic componeni of such strand areas, although t his species is not as universally d istributed as most textbooks claim. Also present in the strand vegetation are Bontia dap/moides (Myoporaceae), Caesa/piuia or Guilandina spp. (Legumi nosae), Clrrysobafanus icaco (Chrysobalanaceae), Erit/raflisfruticosa (Rubiaceae), Hippomane manciuefla (Euphorbiaceae), Met opium toxiferum (Anacardiaceae), Scaet•ofa plumieri (Goodei1iaceac), Suriaua maritima (Su rianaceae or Simarou baceae), Toumejortia gnaplzafodes (Bo raginaceae). These are mostly examples o f single species of wide distri bution. Nearly all present taxonom ic or morphologic problems regmding their classificat ion or relat i on~11i ps. The rock-pavement substratun1 for a recognizable type of vegetation rnay occur ~ coastally on uplifted coral reefs as benches; dominate entire islands of such origin as in the Bahamas; or occur in isolated outcrops in inla nd areas. Such benches and elevated reefs are usually of soft rock, porous and weathered into sink holes of various sizes. They may consist of·extremely hard rock. On the soft rock matrix the vegetation is commonly rooted ove1 a considerable a rea. Succulent herbaceous species of Sesut'ium (Aizoaceae), Portulaca (Portulacaceae), A chyranthes (Amaranthaceae) and Peetis (Composi tae) are abundant. Where s h rubs predominate, the genera Rachical/is, Ernodea (Rubiaceae), Lantana (Verbenaceae), Jacquinia (Theophrastaceae) and Strumpfia (Rubiaceac) may be well rep resented by ind ivid uals of one species or several species. Spectacular coastal benches occu r at the easte111 end of Cuba aro und Punta Maisi, and a series of uplifted coastal benches are found on the so uthern coast of Hispaniola extendin g inland to considerable altitude. Globular growth fo r ms of cacti (Mefocactus) are consp icuous while spiny shrub .vegetation may consist of a large number of endemic genera or unusual genera as Phylostylon (Ulmaceae), tree forms of Cnidoscolus and Victorino (Euphorbiaceae) and clad ophyllo us species of Phyllanthus (Euphorbiaceae). More common s hru bs include Catesbaea (Rubiaceae), Co/ubrina (Rhamnaceae), Cordia (Boraginaceae), Dodonaea (Sapindaceae) Eugenia (Myrtaceae) Randia (Rubiaceae), Reynosia (Rhamnaceae) and Tabebuia(Bignoniaceae). The rock pavement areas may consis t of a very hard type oflimestone rock which erodes into sharp ed ges and which has a metallic ring when struck or when pieces are moved about (Howard and Briggs, 1953). The area has such a characteristic form that the rock is commonly known as d ogtooth limestone. Erosion may form sink holes of

R. A. HOWARD

considerable depth and the plants commonly have a deep tap root. Here one finds such genera as Comocladia (Anacardiaceac), Cynmantiles (Euphorbiaecac), Cuef/arda (Rubiaceac) and Swietenia (Me\iaceae). The mangrove vegetation type found in limited coastal atcas or in extensive swamp for mat ions has been subdivided by some workers into fresh-water or riparian and marine units. Four kinds of mangrove, as single species of Rhizopilora (Rhizophoracea~), Lagwrcu/aria and Conocarpus (Combretaceac) and A t•icemria (Verbenaceae) occur in the Antilles. Rhizopilora is the most abundant of the four occurring in pu re stands on the ocean side, but on the inland margin may be associated with such genera as Caesafpinia (Leguminosae), Dafbergia and Piscidia (Leguminosae), Ximeni; (Oiacaceae) and Acrosticllllm (Pte1 idaceac) as well as the other mangroves. The riparian mangrove formation has been characterized by Pterocarpus (Leguminosae), but this is at best of a limited occurrence. Hibiscus (Malvaceae) and Mac!taerium (Drepanocarpus) of the Leguminosae may also form pure stands or part of a mixture.

Lowland format ions A group of lowland formations may be recognized which can extend to or from the coastal areas. The most common and extecsive of these is a thorn scrub format ion in which the sharp-pointed or recurved hooks may be modified stems, stipules, leaves, inftorescent axes or emergences. Although this unit has been subdivided by Beard ( 1944, 1949, I955) as seasonal-formations and to a Jesser extent by Stehle (I 945, 1946) as xerophytic types, four units are of interest. The most common of the lowland formations is the thorn scrub, considered by many botanists to be the most diagnostic formation in the West Indies since it dominates areas of low rainfall and is most difficult to penetrate and most easy to remember (Plate T, 3). Abundant are Leguminosae as Acacia, Brya, Calliandra, Haematoxylon, Piscidia, Pithecef/obium, Pictetia; Cactaceae as Cereus, and Opuntia and their segregates; Rubiaceae as Guel/arda, Psychotria and Rondeletia ; Euphorbiaceae as A eidolon, Croton, Jatropha and Phyflantlrus; as well as Tabebuia (Bignoniaceae), Bw·sera (Burseraceae) a nd Raut•o/jia and Tabernaemontana (Apocynaceae). Individual plants of this vegetation in coastal areas may be modified into grotesque windswept aberrant forms. In the Dominican Republic and Haiti, a thorn scrub occurs in areas below sea level in the Enriquillo Valley-Cui de Sac area giving way to succulent sea coast vegetation at the shores of the lake {Plate T, 6). It also occurs on alternating salt rock and gypsum outcrops on the slopes of the abutting mountain ranges. Endemism is apt to be high in such areas, and the plants seem to be characterized by small flowers which appear infrequently. Beard attributes the occurrence of savannas (grasslands with scattered shrub growth) in tropical America to alternating periods of water-logging due to poor drainage and to dessication producing conditions too severe for the growth of trees

T HE VEGETATION OF THE ANTILL ES

(Plater, 4). The savanna he classifies as one of a series of seasonal-swamp formations. Stehle docs not recognize a savanna formation , presumably because he considers primarily the vegetation types of the islands of Guadeloupe, Dominica and Martinique. Savannas are infreque nt in the Lesser Antilles and the Grand Savanna of Dominica is the only good example of a Sporobo/us grassland with occasional shru bs of Haematoxylon (Lcgum inosae) and Byrsonima (M alpighiaceae). Savannas on siliceous soils are abui1dant on the Isle of Pines and Pinar del Rio provinces of Cuba and of limited occurrence in the area of Laguna Tortuguero on the north coast of Puerto Rico. These areas, however, may occasionally be so wet as to constitute classification as a swamp. A serpentine savanna occurs in central Cuba with the best examples in the Las Villas (Santa Clara) province (Plate II, 3). Shrubs may be more abundant and the composition of the vegetation more diversified with species of Byrsonima (Malpighiaceae), Jacaranda (Bignoniaceae), Rondeletia (Rubiaceae), and Erytlrroxylum (Erythroxylaceae) showing reduced leaf size. Unusual leaf forms also occur in genera of the Malpighiaceae as well as in species of Aristoloclria (Aristolochiaceae) and Passiflora (Passifloraceae). Other areas of serpentine soils may show a dominance of palms and the speciation of Copernicia in Cuba on serpentine soils is a classic example (Leon, 1946; Leon and Alain, 1951). ~ Areas of persistent moisture can be classi fied as swamp or marsh formations. The largest marsh or swamp in the West Indies is the Cienaga Zapata on the south coa>t of Cuba. An area of this swamp known as the Bahia de Cochinos or the Bay of Pigs has an historical political connotation. The saw grass C/adium (Cyperaceae) dom inates this area, yet ge nera of interesting distributional patterns also occur in the swamp. In open water are species of Va/lisneria (Hydrocharitaceae), Nymplraea and Nuplwr (Nymphaeaceae) and Po/ygonum (Polygonaceae) with northern rather than tropical affinities. In drier areas are found Solidago (Compositae), Cerardia (Scrophulariaceae) and Vicia (Legumi nosae), usually not regarded as tropical genera. Hammock vegetation in the Cienaga Zapata consists of Saba/, Coccothrinax and Acoelorrhaphe _(Palmae), but also Fraxinus (Oleaceae). Swamps are also found occupying large areas on the southern portion of the eastern end of Hispaniola in the vicinity of Higuey. Swamps at higher elevations may be the result of altered d rainage patterns such as the Mason River savanna in Jamaica (Proctor, 1970) and Sabana Nueva above San J uan de Ia Maguana in the Dominican Republic. Peculiar small hillside marshes or savannas occur in the Luquillo Mountains of eastern Puerto Rico. The occurrence of northern genera or those with montane South American affinities make such areas of particular interest. Ascyrum (Guttiferae), Drosera (Droseraceae), Lobelia (Campanulaceae), Potentifla (Rosaceae), Sisyrinclrium (lridaceae), Steffaria (Caryophyllaceae), Utricufaria (Lentibulariaceae), Viola (Violaceae) and Xyris (Xyridaceae) are often found in such swamps. The category of all uvial formations in the Antilles is a useful one and can include such areas as coastal embayments or river flood plains on many islands, the rich

R. A. HOWAR D PLATE I I

J. Mountains of the Cordillera Central of Hispaniola. A pine forest occurs on the ridges, with a broad-leafed vegetation in the valleys. 2. A pine forest in the Sierra de Moa, Cuba. Red lateritic soils support a flora rich in endemic species. 3. An area of serpentine soil with a savanna and shrub vegetation in the Las Villas province, Cuba. 4. A crater lake, the Grand Etang, in Grenada. A montane broad-leafed wet forest. The margin of the lake has a fl oating bog vegetation . 5. An abandoned c ultivated field in Ja maica has developed a shrub vegetation almost entirely of introduced weed species. 6. TI1e Botanic Garden of St. Vincent, the first botanic garden in the West Indies.

TH E VEGETATION OF TH E ANTILLES

I>LATE lll

1. An elfin summit forest in the Luquillo Mountains, Puerto Rico. Area of Prestoea 111ontana (Palmae)

in the upper left corner occurs on a landslide area. 2. Interior of elfin summit forest showing the density of the stems and the abundance of leafy liverworts and other epiphytes. 3. The Cockpit country of Jamaica. A limesto ne mass with steep-sided valleys. Dry limestone fores t type. 4. A mined out pit of bauxite soils, Jamaica. 5. Gage's Soufriere on Montserrat. An example of vegetation loss due to the heat and fumes of active fumeroles. 6. The slo pes of the Soufriere Mountain on Guadeloupe. Period ic emissions from summit fumeroles have killed the woody vegetation. Philodendron giganteum (Araceae) survives the fumes o r invades the affected area.

R . A. HOWARD

tobacco lands of Pinar del Rio in Cuba, portions or the Enriquillo Basin in Hispaniola and the bauxite areas of Hispaniola and Jamaica. The flat lands of Pin ar del Rio asso~iated with the mogotes of the Viiiales area support a vegetation with species or genera of disj unct distribution such as Befaria, Kalmia and Pieris or the Ericaccae, Laclmocau!on (Eriocaulaccae), Lechea (Cistaceae) and Quercus (Fagaceae). The endemic cycad Microcycas is also found in this area. The soils are generally rega rded as a derivative of the limestone mogotes. The occurrence of bauxitic ores in soils of Hispaniola and Jamaica have proven to be of economic significance to the respective cour.tl ;-.~ (?l:: te Ill, 4). The pockets of ba uxite soils occur in depressions in the base limestone rock of Late Eocene or Oligocene age and occur from elevations of 3,000-5,000 ft. The ore as mined is about 20 % ferruginous iron oxide, 50% alumina and less than 3 % silica. The origi n of the ore is still a matter of speculation fo r the base li mestone rock has less than 0. 1 % bauxite, hence a fa ir estimate of over 3,000 ft. oflimestone rock wou ld be required to be eroded and form the amount of bauxite occurring in some of the pits. A few endemic species of flowering plants are recognized on the bauxite soils in Hispaniola, but none has been discovered in Jamaica. The physiologica l effect of the alul]linum ions in the soil on the vegetation, however, has permitted a general classification of the species in the area of bauxite ~oil s as aluminum intolerant, aluminum tolerant and aluminum accumulating species (Howard and Proctor, 1957). Sand cays have developed in coastal areas on many of the Caribbean island s, enclosing bays of shallow water wh ich have been exploited as salt ponds historically as well as currently (Plate T, 2). Such alluvial areas, when unexploited or drying out, natu rally support a halophytic vegetation of succulent herbs as Achyranthes (Amaranthaceae), Atriplex (Chenopodiaceae), Heliotropium (Boraginaceae), Salicornia and Suaeda (Chenopod iaceae) and shrubs or trees as Annona (Annonaceae), Al'icennia (Verbenaceae), Batis (Bataceae), Capparis (Capparaceae), Dalbergia (Leguminosae), Pithecellobium (Leguminosae), and Thespesia (Malvaceae) .

TH E VEGETATION OF THE ANTILLES

fo rmation occurs in the Cockpit country of Jamaica where the erosion holes dominate and the limestone matrix is continuous. The palm genus Gaussia is fou nd on the mogotes of Cuba anti occurs elsewhere only in western Puerto Rico on similar limestone crags. Spathe!ia, a monocarpic member of the Ru taceac, is present on the mogotes of Cuba, in the Cockpit country of Jamaica and on the limestone islands of t he Bahamas. Several species of plants from Venezuela have been transferred recen tly to this genus, but the habi tat of the species is not descri bed. The best examples of stem gigantism in the Anti lles are Bombax emarginatum, found on the mogotes of Cuba. In general species of Andira (Legum inosae), Bucicla and Buchenavia (Combretaeeae), Dipholis (Sapotaceae), Oxandra (Annonaceae) , Sapium (E uphorbiaceae) and Zanthoxylum (Rutaceae) may predominate on the limestone outcrops. The dry limestone mountain forests of the Grenadines in the Lesser Antilles are composed pri marily of species of Bauhinia (Legum inosae) , Bourreria (Boragi naceae), Casearia (Flacourtiaceae), Coccoloba (Polygonaceae), Hymenaea and Lonchocarpus (Leguminosae), Spondias (Anacard iaceae) and Tabebuia (Bignon iaceae). A montane sclerophyll ype of vegetation, occurring in the Trinidad Mountains PLATE IV

Montane fo rmations The mountain vegetation of the Antilles can be subdivided as Beard has done, although a more simple approach is all that is required for descriptive purposes of this paper. Interesting limestone-based forests composed of primarily deciduous species with seasonal flowering characteristics are found on the mogote or haystack mountains of Cuba and Puerto Rico, the Cockpit country and the John Crow Mountains in Jamaica and on the Samana peninsula of Hispaniola (Plate III, 3; Plate I, 5). Endemis~ is apt to be high in these areas. Limestone-based forests of species of wider distribution are found on many islands of the Lesser Antilles. The origin of the mogotes as an erosional feature is clear. The mogotes a re usually steep-sided and composed of a porous structure and possess many erosion caverns. The mogotes are isolated and the valleys continuous in Cuba and Puerto Rico, but perhaps an earlier stage in their

J. The open aspect of a montane forest with trees of Dacryodes, S wartzia, PiJ/recellobium and Ta-

launza, St. Lucia. 2. C limbing plants of the Cyclanthaceae and Araceae dominate a montane forest on Gros Piton, St. Lucia.

R.A. HOWARD

T il E VEGETATION O F THE ANTILLES

and portio ns of the Sierra Macstra ranges of Cuba, predomi nates in Puerto Rico and o n most of the islands of the Lesser Antilles (Plate IV, I, 2) ..:rhe vegetation may occm in layers and Beard's classificatio n is based on the number of tiers of vegetation. The plant s comprising the vegetation are considered as evergreen, without coordinated leaf fall, and fl owering is mostly con tinuous rather than conspicuously seasonal. Genera arc numerous within this vegetati_on, and Stehle lists 175 genera with 385 species in the islands he considers. These are represented as 14% Burseraceae, 10% Lauraceae, 9% Elaeocarpaceae and 7% Myrtaceae. Asprey and Robbins ( I953) indicate the dominants in this forest in Jamaica to be species of Alcllomea (Euphorbiaeeae), Calopllyllum (Guttiferae), Ficus (Moraceae) and Psidium (Myrtaceae). ln the Lesser Antilles the dominant s are a few species of Dacryodes (Burseraceae), Sloanea (Eiaeocarpaceae), Sterculia (Sterculiaceac) and Licania (Chrysobalanaceac) (Hodge, 1954), altho ugh the genera Ormosia, Pitllecellobium and Swartzia (Leguminosae), Simarouba (Simaroubaceae) and Pouteria (Sapotaceae) are well-represented by individuals of single species. · Wi thin the broad-leafed montane sclerophyll vegetation the forest may be d ominated in places by palms. When these occur in conspicuous stands, as on the slopes of El Yunque in the Luquillo Mountains of eastern Puerto Rico, they indicate a n unstable soil area or rock slide zone. Beard has considered tl1e palm a successional development in areas affected by hurricanes, but Bannister ( 1970) disagrees as the resul t of her studies in Puerto Rico. My own as yet unpublished work on a plane crash cite in the Luquillo M ountains reveals palm seedlings as an early invader of open areas, with the fru its of Prestoea montana being distributed by birds and rodents. There is no ready explanation, though, fo r the pure stands of tree ferns, such as Cyatbea sp. often fo und in the Antilles. These are most common in Puerto Rico and occur frequently in the Leeward l slands of the Lesser Antilles. G enerally tree ferns of the Antilles are numerous, but scattered and many species occur only as subcanopy plants. The pine fo rests of the Antilles might be considered as a variation of the montane sclerophyll. The genus Pinus occurs in North America, but does not reach South America. fn the West Indies it is found only in Cuba and Hispaniola (Plate II, 1). I ts introd uction as a timber tree for afforestation has been successful on the other island s only when soil innoculants of mycorrhiza have been m ade. Pure stands of pine occur at low elevations in western Cuba and at higher elevations in the Oriente province of eastern C uba. The Cordillera Central of Hispaniola and the ranges of the southern peninsula possess pine forests on the ridges at higher to summit elevations with a broad-leafed sclerophyll fo rest in many areas in the river valleys. On the north coast of Cuba, near Moa, the pine forest occurs on lateritic iron-rich soils (Plate IJ, 2). Pine forests are good collecting areas for the botanist seeking unusual plant species, and the percentage of endemic species found in pine forests exceeds that oflowland forests. In Hispaniola alone the pine itself is parasitized by Arceutobium bicarinatum (Loranthaceae). This mistletoe has been considered an endemic species restricted to Hispaniola, but the genus is known from western North America as well as Asia. Recently

the same species has been recognized g rowing on Pinus oocarpa in Honduras in Cen tral America. A mo ng the shrubs found in the pine forests arc endemic species of Rubus (Rosaceae), Sah•ia (Labiatae) and Gesneria (Gcsneriaceae), but unusual genera in the Eupho rbiaceae and Mclastomataccae arc frequen t. The vegetatio n is reduced in stature and increases in density at the summi ts of the volcan ic peaks of the Antilles. Since many of these peaks are cloaked in clouds much of the time, the humidity is high and the woody vegetation is commonly laden with epiphytes (Plate Hl , I, 2). This type o f vegetation has been callccl·a mossy forest, an elfin fores t or a montane thicket. Detailed studies of such an area have been pu blished for o nly one such fo rmation (Howard, 1968, 1969) where it is ind icated that the grow th habits of the plants are unus ual in the abundance of a long-shoot- shortshoot and d ichotomous apical growth and of a sy!'n podial branching habit; the abundance of aerial o r advent itious ro ots; the low ferti lity o f the flowers and the red uced rate of metabolism and transpiration within the plants. The transition to lower-alt itude forest types may be gradual or relat ively abrupt. The principal genera in s uch areas may be Clus.ia (Guttiferae), Didymopanax and Oreopanax (Araliaceae), /nga (Leguminosae), Hibiscus (Malvaceae), /lex (Aqui foliaceae), Freziera (Theaceae), Riclreria (Euphorbiaceae), Weinmannia (Cunoniaceae), Tabebuia (Bignoniaceae), Ocotea ( Lauraceae), Psychotria (Rubiaceae) and Charianthus (Melastomataceae). The species p resent are widespread in their distribution in comparable environments and endemics are few. In the Lesser Anti lles volcanic activity has occurred in recent times, and there are currently active fumeroles ( Howard, 1962; Plate IIJ, 5). Regrowth of the vegetation is slow following cataclysmic volcanic activity, and the stages in this development can be found on peaks of adjacent island s. Species of Lobelia (Campanulaceae), Clrariant!rus and Tibouchina (Melastomataceae), Pitcairnea (Bromeliaceae), Cladium (Cyperaceae) and Freziera (Theaceae) appear to be the early invaders. Such fo rmations of scattered or low-stature plants are not comparable to tundra or paramos vegetation as has been suggested. The destruction and loss of vegetation in the lee o f active fumeroles can be observed on many islands. Species of the Araceae, Philodendron and Anthurium, Lycopodium ( Lycopodiaceae) and Pteridium (Dennstaediaceae) are the most resistent to the sulfur fumes or the first to invade a quiescent a rea (Plate HI, 6). A crater lake of about 17 acres and of considerable dept h is found on St. Vincent's Soufriere Mountain. Shallow crater la kes, some ephemeral, are on islands from Grenada to St. Kitts. Neither crater lakes nor fumeroles are found in the Greater Antilles. Eleocharis and Rhynchospora (Cyperaceae), and Nepsera (Melastomataceae) are the principal invading genera of the shallow margins of crater lakes with Montrichardia (Araceae), Miconia, Psychotria as the early invading shrubby genera. The marginal vegetation of the crater lakes often exists as a quaking bog (Plate II, 4). These then are some of the ecological niches on the islands of the West Indies in which plants exist. The total flora of the Antilles is conservatively estimated to be 8,000 species of flowering plants. Unhappily, the vegetation has been studied by many

R . A. HOWARD

THE VEGETATION OF THE ANTILLES

taxon omist s commo nly regarded a s "splitlers", eit her on the basis of their own j udgement of what is an individual taxo n or in the fact that their research is based on the flow of a single island wh ich t hey isola te from the adjacent island s or other areas in their limited considera tio ns. The monograph ic approac h at the generic or fa milial level to the vegetation of northern Latin America, including the Antilles, is needed. P robably not more than I0 % of the genera found in the area a rc represented by monogra phs published in the last fifty years. Thus there is an inherent difficulty of a weak laxo nomic base in mak ing broad sta tements or drawing conclusions on the distributio n of laxa within I he Anti lles . Nevertheless, this is the objective of this paper.

ment is necessa ry in considering the nat ive clements of the Antilles-tn a-discussion of the geographic ranges o f distri bution. In t he total flora of the Anti lles seven f;1mil ies of flowerin g plants predom inate in the number of genera, and there is onl y a slightly difTcrent composition when one considers the number or species as ind icated in Ihe followi ng ta ble. LARGE FAMILIES OF Tllf! WEST INDIAN FLORA

in order of the number of genera

iu order of the mt•nber of species

Composittic Gramincac Lcguminosac Rubiaccac Orchidaccac "Euphorbiaccac Mclastomataccac

O rigins of the existing vcgctalion of the Antilles T he total fl ora of the West Indies involves a sizeable com plemen t of weeds and extraterritorial cultivated plants, often now well-establ ished, which should be excluded from geographic considerations. Pre-Columbian movements of people in the Antillean area are known th rough many p teroglyphs and through a study of artifacts of kitchen middens ( Kirby, 1971). Neither of these sources of evidence of fo rmer occupation, however, yield much, i!' a ny, co nvincing evidence of the plant materials used and therefore p robably introd uced from one area to another. However, the occurrence of certain plants in t he Antilles may be the result of deliberate or accidental pre-Columbian introductio n. The post-Columbian introduction of cultivated plants now naturalized, often as weed s, is better documen ted (Plate Ir, 5). The records of Columbus' voyages indicale the Etfropean plants deliberately carried to be e~tablished as cultivated plants. The introduction of co mmercia l crops such as bananas, sugar cane and coffee can also be documented. The first botanical garden in the West Ind ies was established on St. Vincent in 1765 ( Plate 1[, 6) fo llowi ng the interest expressed by the colonial planters for additional plants of fo reign origin to improve the agricultural econ omy of this tro p ical area (Guildi ng; 1825; Howard , 1953). The voyages of Captain William Bligh for t he breadfruit bro ugh t plants from Europe to the Southern Hemisphere and from the South Pacific islands to fi rst St. Vincent and then Jamaica ( Howard, 1954). The French used the island of M a urit ius as a transplant point fo r East l ndian plants desti ned for the French co lonies and botanic gardens in the Antilles (Jacquin, 1797). An interest in establi shing plantations of rubber-producing species Jed to the introduction o f such plan ts from Central America, South America, Africa, Madagascar and the Pacific tropics (Howa rd and Powell, 1963). Many of the genera and species involved are established and not distinctive in the existing vegetation of the Antilles. The ackee (Biighia sapida) is associated now with the tra nsportation of slaves across the Atlantic Ocean from A frica and it is now well-established in Jamaica. If fruit trees and ornamental plants cou ld be transported, it is not difficult to believe that seed s of grasses and weedy plants could have been introduced just as easily. Thus some judge-

Gramincac Lcguminosac Rubiaccac Composilae Euphorbiaccac Orchidaccac Myrlaccac

Relationships and distribution of the Antillean vegetation The native genera of the AntiHes are mo stly small, appearing to average less than five species per genus. There are ten large genera with sixty or more species, and, as Table II indicates, Cuba a nd H ispaniola tend to have the larger number of species of all the islands. T he number of species in these genera ta pers o fT in the smaller islands a nd in the Lesser Antilles as a group a nd often does so in continental La tin America. Thete TABLE II DISTI\IIlUT ION Of WEST IND IAN G ENf!RA WITH MORE THAN

..!2 -~

.!;!

.§ g

"':-"

Croton Eugenia Miconia Panicum Paspalum Peperomia Phyl/anthus Pilea Psychotria Rondeletia

45 12t 46 71 61 45 65 65 67 60

66 62 64 47 39 60 24 100 37 26

16 45 24 28 26 38 22 45 51 28

13 25 17 31 24 21

II. Piper 12. Pleurotha/lis

30 38

33 27

14 27

I. 2. 3. 4. 5. 6. 7.

8. 9. I 0.

60 SPE CIES

'"'

.;;: ~ ·

.!,!

..!2

"'~

'"' "''"'

()

\:>

22-

16 17 3

51 28 21 23 50 20 20 17 6

13 20 36 39 39 71 II 17 49 5

20 18 65 38 32 176 10 25 52 9

32 63 69 39 57 69 18 26 28 25

10 7

27 18

96 55

371 138

96 68

'--l

R. A. HOWARD

T I-l E VEG ETATIO N OF T HE ANTILLES

are so me genera well-represented or developed in the Ant illes which have but a few s pecies occ urring ou tside or the Antilles. Other genera such as Ptj!l!r and Pleurothal/is a rc extre mely well-devel oped in Central or Sou th America and are less well-represented within the Anti llc~. T he genera which comprise the flora of the West Ind ies reveal ma ny patterns of distribution rang ing from patH rop ical genera to isla nd endem ics and those tbat show disjunct ranges (Fig.3) . These patterns will be cited with comments and documented with examples of genera selected to s how a particular geographic pattern. T he ex-

States; B = the Bahamas; C = Cuba; H = Hispaniola, including Haiti and the Dominica n Repu blic; J =" Jamaica; PR = P uerto R ico; LA = Lesser A ntilles, incl udi ng the Br itish Virgin Islands, the Leeward and Windward Islands south to and incl uding Grenada; CA = Cent ral America, including the tropical portions of Mexico and the Yucatan peninsu la; SA = South America, including T rin idad and Tobago as well as the Dutch islands or Aru ba , Bonaire and Curacao. No .attempt will be made in this paper to distinguish ranges from the Antilles into South America, although ranges to the cast and south from Venezuela occu r as do those west and sou th ft om Venezuela alo ng the A ndes. Both of these ranges may have ex ten~ i o ns into Brazi l or co untries fa r ther to the south. (!) Pan-Caribbean distribution (Table 111). A group of genera found on all o f the islands of the Greater Antilles, the majority of the islands of the Lesser Antilles, and represented in tropical Mexico and Central America and in northern South A merica. (2) A Western Cont inental unit (T able IV; Fig.4). A group of genera wh ich are well represented in Central America and may have a range extending into the Greater

Fig.3. The patterns of geographical distribution and relationship of genera of the West Indies.

amples given do not represent all of the possible examp les or an exhaustive survey of the West Indian genera. Species could also be selected to show th e same patterns. To the best of my ability I have avoided examples from families or of genera in which the existing taxonomic treatments are unsatisfactory. If a genus has been attributed to a geographic area with some question by some author and the record cannot be documented with an herbarium sp ecimen or the commen t of a monographer, I have ch osen to disregard the citation. The foll owing abbreviations will be used in the examples which follow. NA = North America; US = United States, although primarily the southeastern United

Fig.4. The major patterns of generic distribution within Latin America. Vertical lines indicate a Western Continental unit which extends eastward across the Greater Antilles usually terminating in the Virgin Islands (arrows), but with examples extending into Guadeloupe. Diagonal broken lines indicate the Southern Continental unit which extends nort hward in the Lesser Ant illes.

R. A. HOWA RD

Antilles as well as so uth and cast on the South America n continent. This unit is best shown as a lis t of those genera which arc missing in the Lesser Antilles. The extension of these genera across the Greater Antilles may involve Cuba, Hispaniola and Puerto Rico as a pathway and may include or exclude Jamaica. Usually the American and British Vi rgin Isla nds represent a geographic break or terminus of the generic range. Genera which do not appear to occur in Puerto Rico as an extension of the Western Continental ra!lgc or those that may continue into the Leeward 1slancls or Guadeloupe wi lt be listed separa tely as Table V and Table VI respectively. Examples of disjunct distribution between the islands oft he Greater Antilles and South America will be conside red as Tables XXVI - XX IX. Examples of genera wh ich occur in the Antilles and Central Ameri ca, but are wanting in South America form Table VII. (3) A Southern Continental uni t (Table Vllf; Fig.4). A group of genera which are well represented in South Americl) and appear to have a range extension northwa rd through the Lesser Antilles. A few genera may reach Puerto Rico, but these get~era have not been reported fro m Hispaniola, Cuba or Jamaica. (4) The Antillean unit. There are relati vely few genera which appear to be restricted solely to the islands of the Greater Antilles. The two genera _Wa!lenia (Myrsinaceae) and Calycogonium (Melastomataceae) (Table IX) with twenty or more species serve as examples of large genera in the West Indian floRI found only in the Grea ter Antilles. Table X lists genera better developed in the Antilles with a few species in South America, but which appear to be lacking in Central America. (5) Greater Antillean units. The size and complexity of the flora that has developed within the Greater Antilles can be shown best by the number of endemic genera now recognized ar'd their distributi on on one or more islands (Fig.5). Alain (1962) has estimated that Cuba has a flora of 5,800 species with approximately 50% of these endemic to that island. Included in the flora of Cuba are 41 endemic genera cited in Table X I. Although..figures of the total number of species for other islands are not read ily available in modern reliable statistics, estimates can be made of the approximate percentage of endemism and of the number of endemic genera. Hispaniola has specific endemism of about 33 %, and the flora includes 26 endemic genera (Table XII). Jamaica has four endemic genera (Table XIII) and approximately 20% of the flora is of endemic species. Puerto Rico (Table ~IV) has one endemic genus and about 4% of the species are endemic (Britton and Wilson, 1923/ 1924). In contrast, there are no endemic genera in the Bahama islands, but Britton and Millspaugh (1920) suggest that 13 % of the species are endemic. These species have not received study by monographers in recent years, and the figure is certainly high. There are no endemic genera in the Lesser Antilles and a realistic estimate of 12% represents the endemic species. In addition to the local, single island endemics, there are a number of genera restricted to one or more islands. Lists are given of examples of genera found only on Cuba and H ispaniola (Table XV), Cuba and Jamaica (Table XVI), Cuba and Puerto Rico (Table XVII), Cuba and the Bahamas (Table XVIII), Hispaniola and Jamaica

TH E VEGETATION OF THE ANT I LLES

Fig.5. Some distributional patterns within the G reate r Antilles. Estimates of specific endemism a rc given as a percentage, followed by the number o f e ndemic genera of Oowering plants. Generic distribut ion patterns whic h include more tha n one island arc indicated as follows: - . - . -Âˇ . Cuba, Jamaica, Hispanio la, Puerto Rico; ... .. Cuba, Hispaniola and J amaica, but excluding Puerto Rico; - - - - Cuba and Hispaniola, excluding J amaica and Puerto Rico; Cuba, Hispanio la and Puerto R ico, excluding Jamaica; --- - Hispaniola and Puerto Rico, excluding Cuba ~and Jamaica; - ... - ... Cuba and Jamaica. excl uding Hispaniola and Puerto Rico.

(Table XIX), Hispaniola and Puerto Rico (Table XX) and Hispaniola and the Bahamas (Table XX I). There are patterns of distribution of genera which involve th ree of the islands of the Greater Antilles or, in other words, in which a genus has not been reported-from one of the major islands. There are genera which are not present on Cuba, but do occur on Hispaniola, Jamaica and Puerto Rico (Table XX Il) and genera which are absent from Hispaniola, but occur on Cuba, Ja maica and Puerto Rico (Table XX I B). The listings are larger for genera which a ppear to be lacking on either Jamaica (Table XXIV) or Puerto Rico (Table XXV). TABLE Ill EXAMI'LES O F GENERA O F PAN-CARIUBEAN D IST RIBUTION

Amyris (Rutaceae) Anoda (Malvaceae) Apteria (Burmanniaceae) Ardisia (Myrsinaceae) Beilschmiedia (Lauraceae) B;1melia (Sapotaceae) Burmannia (Burmanniaceae) Cedrela (Meliaceae) Celtis (Ulmaceae) Chlorophora (Moraceae) Clidemia (Melastomatacea e)

HMella (Rosaceae) Homalium (Fiacourtiaceae) Jacquinia (Theophrastaceae ) Leiplwmos (Gentianaceae) Linociera (Oteaceae) Manellia (Rubiaceae) Morcgravia (Marcgraviaceae) Miconia (Melastomataceae) Myrica (Myricaceae) Ocoteo (Lauraceae) Ossaea (Melastomataceae)

T HE VEGETATI ON OF THE ANTILLES

R. A. HOWARD

TABLE V

TABL E Ill (conriuuctl)

- - -- --·--·------Co!J11mtcll (Gcsncria~cac) Collo.,rcgiu (Mclastomat<lccac) Comuria (Ycrbcna~cac) Cyrilla (Cyrillaccac) Daplmupsis (Thymcl:tcaccac) Eryrltroxylwn (Erythroxylaccac) Ewmcma (Rubiaccac) Grit.f}enriedia (Mclastomataccac) Guarea (Meliaccac) Gtutzwna (Eiacocarpaccac) Guellarda (Rubiaccac) Hemandia (HcrnallCiiaccac) Hillia (Rubiaccac) Hippocrarea (Hippocratcaccac)

EXAM I'll:._') OF G f. N I:I~A W II ICil DO NOT tti:ACI I PU_f-UTO IUCO AS AN E,\STEH. N I.:X'n~NSION OF A. Wt:STERN

Ourarca (Ochnaccac) Oxoudm (Annonaccac) Picramnia (Silmrubaccac) Pudocarpu.1· (Podocarpaccac) Rondeletia (Rubiaccac) SaMa (Labiatae) Sopium (Euphorbiaccac) Scltlegelia (13ignoniaccac) Scltoepfia (Oiacaccac) · Si11wrouba (Simaroubaccac) Strucltiwn (Compositac) Tiboucltina (Mclastomataccac) Tricltilia (Mcliaccac) WeiunWJmia (Cunoniaccac)

C'ONTIN I: NTI\L H AN(iJ:

MatlfJia (!cacinaccac) CA. SA Mascagnia (Malpighiaccac) CA, SA Neurolaena (Compositae) CA , SA Oocarpwr (Onagraccac) CA, SA Plryllosry/on (Ulmaceae) CA, SA Rammw/us (Ranunculaccac) CA, SA Samolus (Primulaccac) US, B, CA, SA Srexnaspemw (Phytolaccaccac) CA Tapura (Dichapctalaccac) CA, SA Va/lcsia (Apocynaccac) US, B, CA, SA Zuclania (F iacourtiaccac) B, CA

All•ttrttduo (Simaroubaccac) US, B, CA, SA Areuaria (Caryophyllaccac) US, CA, SA Budt!/eia (Log;•niaccac) CA , SA Esenbeckia (Rutaceac) CA, SA Eusroma (Gcntianaccac) CA, SA Forchlwmmeria (Capparaccac) CA, SA Gonya (Garryaccac) US, CA Kosrelerzkya (Malvaccac) US, CA, SA Laeria (FiacourtiacC<lC) CA, SA Liabum (Compositac) CA, SA Lwwnia (Fiacou rtiaccac) CA, SA

TABLE IV WESTERN CONTI NENTAL UNIT-'EXAM I'Lf.S OF GENERA MISSING FROM THE LESSI! R ANTILL ES

- - -- - - -- - - - · -·

lld.wmtltera (Mclastomataceae) GA, CA, SA Alcltomca (Euphorbiaccac) GA, CA, SA Augdnnia (Scrophulariaccac) GA, CA, SA Atriplex (Chcn opodiaccac) GA, CA, SA Bmnellia (Brunelliaceac) GA. CA, SA Buclmera (Scrophulariaccac) US, B, GA, CA, SA Callimrpa (Yerbcnaccac) US, GA, CA, SA Cleyera (Thcaceac) GA, CA Dct~drorJcmoll

( Loranthaccac) GA, CA, SA Det1drophrltora (Loranthaceac) GA. CA, SA Drosera (Droscraceae) NA, GA, CA, SA &·hires (Apocynaceae) US, GA, CA, SA Ehretia (Borag inaccae) GA, CA, Asia Esenbeckia (Rutaceae) C, H, J , CA, SA Eusloma (Gcntianaceae) US, B, C, H, J, C A Forchlwmmeria (Capparaceae) C, H, J, CA, SA Forsreronia (Apocynaccae) GA , CA, SA Gerardia (Scrophula riaceae) US, GA G01·rya (Garryaceae) C, H , J, CA, US Gyminda (Celastraceae) US, GA, CA He/icleres (Sterculiaceae) GA, CA, SA Hieronima (Euphorbiaceac) GA, CA, SA Hyperbaena (Me(lispermaccac) G A, CA, SA lsocarpha (Compositac) US, GA, CA, SA Jug/01rs (Juglandaceae) C, H, PR, CA, SA, US Laetia (Fiacourtiaceac) C, H, J, CA, SA Laplacea (Theaceae) GA, CA, SA Liabum (Compositae) C, H,J, CA, SA Licoria (Lauraceae) US, GA, CA, SA Lwrania (Fiacourtiaceae) C, H, J, CA, SA Lyonia (Ericaceae) GA, CA, US, Asia Machaonia (Rubiaceae) GA, CA, SA Macrocarpaea (Gentianaceae) GA, SA, CA

TABLE Vf

Magnolia ( Magnoliaccac) C, H, PR, CA, SA Mappia (lcacinaccac) GA, CA, SA Mai'sdcnia (Asclepiaclaccac) GA, CA, SA Mawyba (Sapindaccac) G A, CA, SA Meriania (Mclastomataccac) GA, CA, SA Nama (Hydro phyllaccacl US, GA, CA, SA Neea (Nyctaginaceac) GA, CA, SA Newolaena(Composi tac) C, H. J , CA, SA .Oocarpon (Onagraceac) C, H, J , CA, SA Ossaea (Melastomataccae) GA, CA, SA Oxyperaltan (Asclcpiadaceac) GA, CA, SA Phaeo.1plwerion (Commelinaceae) C, H , PR, CA,

EXMII'LES O F GENERA OF T i lE GREATER ANTI LLES W HI CII REACII THE LEEWARD (NORT"IIEH N) ISL ANDS OF HI E L ESSER ANTILLES

Borrichia (Compositac) US, B, GA, Guadeloupe Forcsriera (Oicaccac) US, B, GA, CA. Antigua Gltinia (Vcrbcnaccac) C , H, PR, CA. SA, Antigua Hybam/ms(Violaccac) C, H, PR, CA, SA, Guadeloupe Hype/arc (Sapindaceac) US, B, GA, Anguilla Metopium (Anacardiaceac) US, GA, Anguilla Mouriri (Mclastomataceae) GA. CA. SA. G uadcloupc NymtJiwides (Mcnyanthaccae) GA, CA, SA, Guadeloupe Rubus (Rosaceae) US, GA, CA, SA,St. Kitts Srrrmrpjia (R ubiaceac) US, GA, Guadeloupe

-Phyllosrylon (Ulmaceae) C, H, CA, SA Portlandia (Rubiaceac) GA, CA .Pseudo/media (Moraceae) GA, CA , SA Rachicallis (Rubiaceae) B, GA, CA ROirrmcrt/us (Ranunculaceae) C, H , J, CA, SA Ravenia (Rutaceae) GA, CA, SA Sag if/aria (Aiismataccae) US, GA, CA , SA Salicomia (Chenopodiaceae) B, GA, SA, CA Sa/mea (Compositae) GA, CA, SA Samolus (Primulaceae) C, H , J, CA, SA Sarcoslema (Asclepiadaceac) US, B, GA, CA, SA Schaefleria (Celastraceae) US, GA, CA, SA Scybalium (Balan ophoraceae) GA, SA Sregnosperma (Phytolaccaceae), C , H , J, CA Stel/aria (Caryophyllaceae) B, GA, CA, SA Solandra (Solanaceae) GA, CA, SA Thalia (Marantaceae) GA, CA, SA Valeriana (Yalerianaceac) NA, GA, CA, SA Ximenia (Oiacaceae) US, GA, CA, SA Zamia (Cycadaceae) GA, CA, SA

T ABLE Vfl

EXAMPLES OF WEST INDIAN GENERA WH ICH A P PEAR TO MISS SOUTH AMERICA

Bor.-iclria (Co mpositae) US, B, GA, LA Carpadiprera (Tiliaceae) C, H, LA, CA

Lyonia (Ericaceae) US, GA, CA Masrichodendron (Sapotaceae) US, GA, LA, CA

Dip/ro/is (Sapotaceac) US, GA, LA, CA

Phialam/rus(Rubiaceae) GA , LA, CA

Elrrelia (Boraginaccae) GA, CA Eritlral/is (Rubiaceac) GA, LA, CA

Samyda (Fiacourtiaceae) GA, LA, CA Stegnosperma (Phytolaccaccae) C, H, J , CA Terrazygia (Melastomataceae) GA , LA Tropidia (O rch idaceae) US, GA, LA, CA Ureclrites (Apocynaceae) GA, L A, CA

Ernodea(Rubiaceae) US, GA, LA Exot/rea (Sapindaceae) US, GA, LA, CA Gyminda (Celastraccae) US, GA, CA Krugiodendron (Rhamnaceae) US, GA, LA, CA

R. A. HOWARD

T HE VEGETATION OF THE ANTILLES

TAllLE VIII

TABLE X

SOUTIWRN CONTINENTAL l!NIT- EXMII'LES 0 1' v ENf.RA OCCUIHU I':G IN TilE LESSF.R ANTI LLES HUT Wll l{'ll DO NOT OCCUI{ IIEYOND I'UERHl RICO IN Ti lE v REATER ANTII.LI;5

I.XAMI'LI~S

ANTILI..I~AN

/Jomia (Myoporaccac) ll, GA, LA, SA Casasia (Rubiaccac) US. ll, C, H, J, LA, SA Cymlu>carpa (llurnmn niaccac) C, H, J, SA Gesneria (Gcsncrinccilc) GA, LA, SA HuriWIIWIIIIia (Ericaccac) GA, LA, SA Nepsera (Mclastoma taccac) GA. LA, SA Rhytidopltyllum (Gcsncriaccac) GA, LA Schradera (Rubiaccac) GA. LA, SA Tetrazn:ia (Mclastomataccac) GA, LA , SA

A11umoa (Euphorbiaccac) SA, Guade loupe, Dominica A11iba ( Lauraccac) LA, CA, SA, Puerto Rico Browallit1 (Solanaceae) LA, SA Ca/u/isiant/ms(Gcntiana=c) LA, SA

Cenlropogon (Campa.nulaL·cac) LA, CA, SA C/wme/ia (Rubiaccac) LA, CA, SA Cltrysuch/amys (Guttifcrac) LA, CA, SA Codonantlte (Gcsncriaccac) LA, SA, CA Coutonbea (Gcntianaccac) LA, SA, CA Dacryodcs (llurscraccac) LA, SA, CA, Puerto Rico Drymonia (Gcsncriaceae) LA, SA, CA Dussia (Lcg uminosac) LA, SA, CA Enicostema (Gcntianaccac) LA Gonolobus (Asclcpiadaccac) LA, SA, CA, Puerto Rico Isclmm·il'holl (Marantaccac) LA, SA, CA, Puerto Rico Licania (Chrysobalanaccac) LA , SA, CA Malanea (Rubiaceac) LA, SA Nauti/o caly.r (Gcsncriaccac) LA, SA, CA Nomntea (Marcgraviaccac) LA, SA, CA Petrea (Vcrbcnaceae) LA, SA, CA Pres1o11ia (Apocynaceac) LA, SA, CA Plerc/epis(Mclastomataccae) LA, SA, CA Riclteritl (Euphorbiaceae) LA, SA Rola11dm (Compositae) LA, SA, CA, Puerto Rico Ruyschia (Ma rcgraviaceae) LA, SA, CA Sipanma (Monimiaceae) LA, SA, CA Srylogyl/e (Myrsinaceac) LA, Puerto Rico To••omita (Guttifcrac) LA, SA, CA Willmackia (Bromcliaceae) LA, SA, CA, Puerto Rico

(ii::"\F.RJ\ W I IICJI ARI-: LAC KING IN CI:I'\'T!tAL AMERICA

TABLE XI G ENERA

Ancistrimt/w;- (Acanthaccac) Behaimia (Lcguminosac) !Jembi<'idium (Lcguminosac) Caribea (Nyctaginaccac) Cerlllopyxis (Rubiaccac) Cubat'I'OI<III (Euphorbiaccac) Dm·ytropi.~

(Acan thaccac)

Doerpfcldia (Rhamnaceac) Eosanthe (Rubiaccac) Espadaca (Solanaceae) Euc/wrium (Sapindaccac) Euleria (AnaCHrdiaceac) Goerzie//a (Amaranthaccae) Hamackia (Compositac) Hebe.~ti~tma (Lcguminosac)

Hen/eaphytu111 (Malpighiaccac) Heptwlllws (Compositac) Herpyza (Lcguminosac) Koe/meo/a (Compositae) Lescai//ea (Composilae)

~NDEM IC

TO CUll/\

Linude11dmn (Thymclaeaceae) Megalopanax (Araliaceae) Microcycm· (Cycadaccac) Nodoct/J}Jea (Rubiaccac) Pllidiasia (Acanthaccac) Plty//acamlms (Rubiaccac) Phyllomelia (Ru biaccac) Piuosia (Caryophyllaceac) Rlwdogerou (Compositac) Sapphoa (Acanthaccac) Sclllnidtollia (Rubiaccac) Slwfera (Compositae) Sieme11sia (Rubiaccac) Solonia (Myrsinaccac) Spirotemma (Bignoniaccae) Tetralix (Tiliaccac) Tetraperoue (Compositac) Web;-teria (Cypcraceae) Woehleria (Amaranthaceae) Zollalllhus (Gentianaceae) TABLEX!l GENERA ENDI: MI C TO HISPANIOLA

TABLE IX EXAM P LES OF ANTILLEAN GENERA MISSING IN CENTRAL AMERICA AND SOUTH AMERICA

Ca/ycogonium (Melastomataceae) Catesbaea (Rubiaceae) Hype/ate (Sapindaceae) Lasiantlws (Rubiaceae) Reynosia (Rhamnaceae) Strumpfia (Rubiaceae) Wa//enia (Myrsinaceac)

Arcoa (Legum inosae) Coe/olleurum (Solanaceae) Cryplorhiza (Myrtaceae) Ekmmtiocltaris (Melastomataceae) Fuertesia (Loasaceae) Haitia (Lylhraceae) Herodtia (Compositae) Hybosperma (Rhamnaceae) Leprogommt (Polygonaceae) Mallfe/dia (Compositae) Mommseuia (Melastomataceac) Neobuchio (Bombacaceac) Pedinopetalum (Umbelliferac)

Peue/opeia (Cucurbilaceae) Peratautbe (Rubiaceae) Piperomhera (Piperaceae) Poitaea (Leguminosae) Pterocissus (Vitaceae) Rhodopis (Leguminosae) Sarcopilea (Urticaceae) Se//eo/a (Caryophyllaceae) Stevensia (Ru biaceae) Ulbrichia (Malvaceae) Vegaea (Myrsinaceae) Wuuschmannia (Bignoniaceae) Zombia (Palmae)

R. A. HOWARD

T HE VEGETATION OF H IE ANTILLES

T AllLE XIII GENER A I' NI>IO MIC TO JAMA ICA

Dcndroeoasinia (Euphorbiaccac) Jamima (Asclcpiadacc:.c) Peltosligma (Rutaceac) Telrasiphon (Cclastraceac)

TABLE XIV

TA BLE XV I foXAMI'I.I'S OF GI'NiillA FOU ND IN C UBA AND JA MAI C'A, BUT W ill('! I AP P EAR TO llli MISSING IN IIISl'ANIOLA ANI) I'UI'IlTO RICO

Acro.\ytwmlw.l· (Rubiaceae) endemic Auerudemlrou (Rhamnaccac) B Bemardia (Euphorbiaccac) LA Cenlaurium (Gcntianaceac) US, B. SA Cal)'plrolloma (Palmac) e ndemic Clrrys01hcmi.1· (Gesncriaccac) LA, CA, SA Cionocisyos (Cucurbitaccac) endemic Cle1hra (Ciclh raccac) US. CA, SA Dmilla (Dil lcniaccac) endemic Freziem (Thcaccac) CA, SA

GENERA ENDEMIC TO I'UERTO RICO

Grimmeotlendw11 (Euphorbiaccac) ll Hydrulea (Hyd rophyllaccac) US, LA, CA, SA Qnii11a (Quiinaceac) SA, CA Rha11midim11 (Rhamnaceae) CA, SA Simuia (Cucurbilaccac) SA S/mlhelia (Rutaceae) B, SA Strempe/iopsis (Apocynaceae) endemic Teimcerct (Dillcniaceae) CA, SA Vibumum (Caprifoliaccae) US, CA, SA Xylopia (Annonaccae) CA, SA Zueltmia (Fiacourtiaceae) B

Cybitmllwpsis (Myrsinaceac) TABLE XVII EXAMI'LE OF Gf:NERA FOU NO IN CUDA AND I' U ERTO RICO

Cau~:ria (PalnHie)

endemic

TABLE XV TABLE XV III

EXAMI'LES OF G I:NERA \VHI Cii OCCUR ON LY ON CUOA AND HISI)ANIOLA IN GRI!ATI:R ANTILLES

EXA MPLES Or GF.~F.ItA FO UND IN CUllA AN D TilE OAt-l AMAS

Ampeloc-cras (Ulmaceae) endemic Ale/cia (Leguminosae) CA, SA Bellcmia (Gesneriaceae) endemic Bisgoeppet"lio (Gentianaceae) endemic Bonottia (Euphorbiaceae) ll Cm·podiptcra (Tiliaceae) LA, CA, SA Cltascollteca (Euphorbiaceae) end emic Cubantitus (Euphorbiaceae) endemic Curalella (Dilleniaceae) CA, SA Ekmanianllte (Bignon iaceae) endemic Eriosema (Leguminosae) CA, SA Goc/matia(Compositae) US, B, CA, SA Hucr/ea (Staphy leaceae) SA Illicium (!lliciaceae) CA, US lsidorea (Rubiaceae) endemic Jacaranda (Bignoniaceae) B, CA, SA Junipems (Pinaceae) US, B, CA Kanvittskia (Rhamnaceae) US, CA, SA Lattlanopsis (Compositae) endemic Limnocltaris (Butomaceae) CA, SA Macrocarpaea (Gentianaceae) endemic Marila (Guttiferae) L A, CA, SA Mayaca (Mayacaceae) CA, SA Mesechites (Apocynaceae) CA, SA Neomazea (Rubiaceae) endemic Neoregnellia (Sterculiaceae) endemic

Pacltyanllttt.s (Mclastomataccae) SA Paconritta (Compositae) CA, SA Paepa/mulms (Eriocaulaceac) US, CA, SA Pararlteria (Gramineae) SA Pcm (Euphorbiaceac) B, CA, SA Phiuaea (Gesncriaceac) SA Phyllo.l'l)'lon (Ulmaceae) CA, SA Picardea (Rubiaceae) endemic Piuillosa (Compositae) endemic Pimts (Pinaceae) US, B, CA Pleilwdenia (Rutaceae) endemic Prolitmt (Proteaceae) LA , CA, SA Pseudocarpidium (Verbenaceae) B Sabatia (Gentianaceae) US, CA Saugelia (Gramineae) endemic Sc/w/resia (Genlianaceae) CA, SA Scutachue (Gramineae) e ndemic Sip!tocampylus (Campanulaceac) SA Strychnos (Logan iaceac) CA, SA Tapura (Dichapetalaceae) LA, CA, SA Thouiuidium (Sapindaceae) CA Tibouchiua (Melastomataceae) LA, CA, SA Verheu/lia (Piperaceae) e ndemic Victorino (Euphorbiaceae) endemic Wigandia (Hydrophyllaceae) CA, SA

Neobmcea (Apocynaccae) endcm ic Neolhymopsis (Compositae) endemic

T ABLE XIX EXAMI'LioS O F GEN ERA FOUND IN H ISPAN IOLA AND JAMAI CA, OUT NOT llEI'OilTED IN CU BA OR PUERTO RICO

Cmtaeva (Capparaceae) CA, SA Oreopauax (Araliaceae) LA, CA, SA Symplto11ia (Guttiferae) LA, CA, SA

TABLE XX EXAMPLES OF GENERA FOUND IN HJSI'ANIO LA AND PUERTO RICO, BUT NOT REI'ORTEO FROM CUBA O R J AMA ICA

Alchomeopsis (Euphorbiaceae) SA Coetzia (Solanaceae) endemic Krameria (Krameriaceae) LA, CA, SA Morrisonia (Capparaceae) LA, CA, SA Ottoschmidtia (Rubiaceae) endemic Piptocoma (Compositae) e ndemic

Pleode11drou (Canel!aceae) endemic Quararibea (Bombacaceae) LA, CA, SA Ro/linia (Annonaceae) LA Staltlia (Leguminosae) e ndemic Terragaslris (Burseraceae) CA, SA

R. A. HOWARD

T I-l E VEGET AT ION O F T HE ANT IL LES

TABLE XX I

T ABL E XXV

FXAMI 'L~ OF GENU< A I"OUNO I N II ISI'ANIOLA ANilll ll! IIAIIAMAS. BUT NOT ON C Ull.\, JAMAICA U~ I'U[I{TO

f:.XAMI'LL"i OF (ii:NHtA \V I tiCH AI'PJ:Ail TO IJE AIJSI:Nl" FH.OM I'UI!KTO K.ICO, UUTOCCUR ON CUBA, II ISPANIOLA

ltiCO

ANI)

Tcmmthus {Compositac) endemic

Aciducrowu {Euphorbiaccac)cndcmic Ail'fu·udoll (Simaroubaccac) US, ll, CA. SA ;lreuaria (Caryophyllaccac) US, CA, SA /Jesleria {Gcsncriaccac) LA, CA · Broughtunia (Orchidaceac) endemic Brya ( Lc~:uminosac) endemic Budd/cia (Loganiaccac) CA, SA Cameraria (Apocynaccac) endemic Co.wsia (Ru biaccac) US, ll, LA, SA · Cotalp11 (Bigno niaceac) US, Asia Cimwmodcudron (Cancllaceac) SA Cyatlwla {Amara nthacc;tc) LA, CA, SA Cymbocarpit (Burma nniaccae) SA Eseubeckia (R utaccac) CA, SA Eustoma (Gentianaccac) CA, SA Forchhlllllllterio {Capparaccac) CA, SA G11rry11 (GMryaccac) US, CA Grimmeodendrou (Eup horbiaccac) D Gyrotaenill (Urticaccac) endemic

TA BLE XX !l I'XA ~ I I'LES Of G ENERA W III C H AI'I'I!AR TO liE ABSENT FROM CUBA, BUT OCCUII ON IIISI'A NIOLA, J AMAICA

AND I'UERTO RICO

Alloplectus {dcsncriaccac) LA, CA, SA Ormo.l"ia (Lcguminosac) LA, CA, SA Pouteria {Sapotaccac) LA, CA, SA Sabicea (Rubiaccac) CA, SA

Jnncns{J uncaccac) US, LA, CA, SA Ko;·u:letzkya (Malvaceac) US, CA, SA, Africa

TABLE XXIII

JA~IAlCA

J.aetia (Fiacourtiaccac) CA, SA Lagella (Thymclae<lccac) endemic Lasiocroton {Euphorbiaccac) B Letteocroton (J::uphorbiaccac)-cnctctn ic Liabum {Compositac) CA, SA Lmumia (Fiacourtiaccac) CA, SA Mascaguia {Malpighiaccac) CA. SA Neuro/aeua (Compositac) CA, SA Omplwlea (Euphorbiaccac) LA, CA , SA Oocarpon (Onagraccac) CA, SA Phyl/ostylo11 (Ulmaceae) CA. SA Pkrodeudrou (Picrodcndraccac) endemic Rmllmmlw· (R<In unculaccac) CA, SA

Samo/11;· {Primulaccac) US, B, CA, SA S teguospemw (Phytolaccaccac) CA Tttpura (Dichapctalaccac) CA, SA Triopteris (M alpighiacc:\C) B Vallesia (Apocynaccac) US, B, CA, SA Vacciui11111 (Ericaccac) US, LA, CA, SA Zuelania (Fiacourtiaccac) B, CA

EXA MI' LES OF GENERA W HIC H Al'l'EAR TO BE ABSENT FROM HI Sl'A NIOLA, BUT OCCUK ON CUUA, JAMA ICA AN D PU ERTO RICO

Holtenbergia {Bromeliaccac) LA, SA Marsypiautlw.1· {labiatae) LA, CA, SA

Disjunct distribution patterns in the Antilles

J'dicmpho/is{Sapotaceae) LA, CA

TABLE XX IV EXAMPLES OF GENERA WHIC H AP P EAR TO BE ABS ENT FKOM JAMAICA, llUT OCCUR ON CUBA, HISI'AN IOLA AND PUERTO RICO

Da/ec/rampia {Euphorbiaceae) CA, SA Didymopauax {Araliaceae) L A, CA, SA Ditta {Euphorbiaceae) endemic Doliocarpus {Dilleniaceae) LA, CA, SA Guattaria (Annonaceae) LA, CA, SA Hybauthus (Violaceae) LA, CA, SA Ixora (Rubiaceae) LA, CA, SA Jug /aus (Juglandaceae) US, CA, SA Magnolia (Magnoliaceae) US, C A, SA Me/iosma (Sa biaceae) LA, CA, SA Ottosc/rulzia (Icacinaceae) endemic Parathesis (Myrsinaccae) CA

Plraeosphaeriou {Commelinaceae) CA, SA Pilocarpus (Rutaceae) LA, CA, SA P iuus(Pinaceae) US, B, CA Prockia (Fiacourtiaceae) LA, CA, SA Proustia (Compositae) SA Rlrexia (Melastomataceae) US Scolosantlws (Rubiaceae) endemic Scrop!tularia (Scrophulariaceae) US Styrax (Styracaceae) US, L A, SA Terebraria (Ru biaceae) B, LA Tetragastris (Burseraceae) CA, SA Torralbasia (Celastraceae) endemic

l· l

T here a re many exa mples of d isjunct distribution patterns of genera between one or mo re islands of the Greater Antilles with isla nds of the Lesser Antilles, with Central A merica, o r with South America. These can be genera wi th several species in the Antilles a nd a large n umber in a distant area. Table XXVI consists of genera indigenous to C uba alone of the Greater Antilles. Only three o ft he examples cited occur also in the Lesser Antilles, fo r the more com mon range extension is to Central America and/or Sou th America. Lists for Hispaniola (Table XXVII) and Jamaica (Ta ble XXV lll) show a smaller number of genera comparably represented only on each island of the Greater Antilles. O nly two genera are cited for Puerto R ico (Table XXIX) as indigenous to that island but occurring elsewhere in Latin America. Examples of genera occurring on two isla nds of the Greater Antilles and with a disjunct distribution elsewhere are infrequent. Tapura (Dichapetalaceae) occurs on Cuba and Hispaniola, in the Lesser Antilles and South America, but is unreported from Central America. Alchorneopsis (Euphorbiaceae) is reported from· Hispaniola and Puerto Rico a nd from South America. Symphonia (Guttiferae) occurs on Hispaniola and Jamaica and is fo und in the southern Lesser Antilles as well as in Central America and South America. Cymbocarpa (Burmanniaceae) is found in C uba, Hispaniola and Jamaica and occurs elsewhere in Latin America in South America.

R. A. HOWARD

TH E VEG ETATION OF TH E ANTILLES

TABLE XXVI

TABLE XXVIII

I:XAM I'Lf.S OF (if.NEilA INDIGENOUS TO CUUA ALONE OF TilE (illFATII\ ANTILLES

l :XAM I'LES OF G[Nt'ltA INDIGENOUS TO JAM AICA Ol>.LY OF Till: Glt[ All;R ANTILLES

--- - ·- ---.

A coelorraplte( Palm<~c)

US, CA Actinostemt'll (Euphorbiaccac) LA, SA Amaitma (Rubiaccac) CA. SA Bcfm'ia (Eric<~ccac) US, CA. SA /Jeloria (Tili<~ccac) CA Bcrbcri.1· (Bcrbcridaccac) CA /Jonuetitt (Thcacc<IC) SA Brao•tti.-ia (Acant haccac) CA, SA Ca /ycophylluut (Rubiaccac) CA, SA Calyptroclu'f)/1.\' (Composilac) US, CA Cephalaurlm.\· (Rubiaccac) US, CA Clwerolepi.l' (Mclastomataccac) SA Cienfuego.ria (Ma lvaccac) US, CA, SA Cneorum (Cncoraccac) Canary lsi., Mcditcrr. Cou.mwea (Rubiaccac) CA, SA Dec/ieuxia (Rubiaccac) CA, SA Deheriania (Thcophrastaccac) CA Desmopsis(Annonaccae) CA Dmccna (Liliaceae) CA Elaeagia (Rubiaceac) CA, SA EMra (Compositac) CA, SA Eriocaulon (Eriocaulaccac) US, CA, SA Eriope (Labiatae) SA Fmxinus (Oicaccac) US, CA Gen/isea (Lcntibulariaccac) SA Gossypio.rpermum (Fiacourtiaccac) SA Harpalyce (Lcguminosac) CA, SA Helielfa (Rubiaccac) CA Helo>ir (Balanophoraccac) LA, CA, SA lsertia (Rubiaceae) CA, SA Kalmia (Ericaccae) US

Laclwamltes (Haemodoraceac) US Larluwctmlon (Eriocaulacc;oe) US Leberiua (Compositac) CA Lee/tea (Cistaccae) US Lepidesmia (Compositae) SA Lueltea (Tiliaceae) CA, SA Mamuthmm ( Po dostem~tceae) CA, SA Margarantlm.r (Solanaceae) Me/mwullws (Solanaceac) CA, SA Milleria (Compositac) CA, SA Neoma<fad)'ll (Uignoniacene) CA Oenot!tera (Onagraccae) US, SA Pam thesis (Myrsinaceae) CA Picras11w (Simaroubaccac) LA, CA, SA Pieris (Ericaccae) US Poln>retmml (Loganicene) US, CA, SA PriSJimeria ( Hippocratcaceac) CA, SA Proserpinaca (1-la loragaccae) US, SA Purdiaea (Cyrillaceae) CA, SA Quercus (Fagaceae) US, CA Rims (Anacardiaccae) US Salacia (Hippocrateaceae) CA, SA Schweuckia (Solanaceae) CA, SA Stipulicida (Caryophyllaceae) US Syugo11a1111ws (Eriocaulaceae) US, CA, SA Talauma (Magnoliaceac) LA, CA, SA Tocoyeua (Rubiaceac) SA Touiua (Eriocaulaceae) CA, SA Tric!tmpira (Compositac) CA, SA Tristiclta (Podostcmaceae) CA, SA Unciuia (Cyperaccae) CA, SA

Kegeliella (Orchidaccae) CA. SA Loci.1·tcma ( Lacistcmataccac} CA, SA MtnTomemum (Rubiaccac) CA, SA Mmulco•illa (AilOcynaceac).CA, SA Mosquiroxylou (Anacardiaccac} CA Notoptem (Composirae) CA Prioria (Leguminosac) CA Pseudocentrum (Orch idaceac) LA, CA. SA P.>ittacantlws (Loranthaccae) LA, CA, SA Pteric!tis (Orchidaccae) CA, SA Tetrorchidium (Euphorbiaccae) LA, CA, SA

TABLE XXIX f.XAMI'LES OF GENERA INDIGENOUS TO I'UERTO RICO ONLY Of THE

ANTILLE~

Gonoca/yx (E ri~accac) SA Piptocarpa (Compositae) CA, SA

Distributian patterns within the islands Cuba and Hispanio la are the two largest islands of t he Greater Antilles. They possess the largest flora, the greatest d iversity of topography, great isolation of ecological niches, the highest percentage of endemism and offer the most frequent examples of disj unci ranges. The uniqueness of the flora in various sections of Cuba (Fig.6) has been emphasized in the publications of Marie-Victorin and Leon (1942, 1944, 1956) and of Sei friz (1943). The unusual floras of Pinar del Rio and the Isle o f Pines, the_ serpentine lands of Las Villas (Santa Clara) provi nce and the mountains of the · Oriente province have received considerable attention in the past. Moscoso (1943) delimited thirteen units of geographic distribution in considering

TABLE XXVII

Lepechiua (Labiatae) CA, SA Linaria (Scrophulariaceae) NA, CA, SA Loasa (Loasaceae) CA, SA Narvaliua (Compositae) SA Pereskia (Cactaceae) CA, SA Sclerothrix (Loasaccae) CA, SA Spananthe (Umbclliferae) CA, SA Sphyrospermum (Edcaceae) CA, SA Spiracautha (Compositae) CA, SA Viola (Violaceae) NA, LA, CA, SA

----------- --------

AdclobotrJ•s (Mclastomataccac) CA, SA Ariiii'Osll!lllltW (Mclastoma taceac) CA, SA Blakea (Mclastomataccac) LA, CA, SA Calea (Compositae) CA, SA Ceplwefi.,· (R ubiaccac) LA, CA, SA Clmeuocep!ta/us(Compositae) CA, SA Chariantlws (Melastomntaccae) LA Coccoloba (paniculatc inOorcsccnt forms) SA Geissomeria (Gcsncriaccac) SA Gria.1· (Lccythidaceac) CA, SA Gynoxys (Compositae) SA Ho./finannia (Rubiaccac) CA, SA

EXAMPLES OF GENERA FOUND IN HISPANIOLA ALONE OF TH E GREATER ANTILLES

Arceutltobium (Loranthaceae) NA, CA, SA Aspidosperma (Apocynaceae) SA Castilleja (Scrophulariaceae) NA, CA Cftimaphila (Pyrolaceae) NA, CA Clavija (Theophrastaceae) CA, SA Coreopsis (Compositae) NA, CA, SA Dactylaeua (Capparaccae) SA Disciphania (Menispermaceae) CA, SA Dussia (Leguminosae) LA, CA, SA Gaultheria (Ericaceae) NA, LA, SA Halimium (Cistaceac) LA, Afr., Eur.

Fig.6. The provinces of Cuba. PR = P inardel Rio; IP = Isle of Pines; Ii = Havana; M = Matanzas; LV= Las Villas; CAM = Camaguey; OR= Oriente.

R. A. IIOWARD

Ti lE VEGETATION OFTHE ANTILLES

T/\ilLE XXX

the ot her western provi nces, but only two in eac h of Camagucy and Oriente provinces. The. sandy plains and swamps of Cuba prQ.vide the habi tat for continental North American genera such as Kalmia, Pieris, Quercus and Fraximts. The ge nera of Cuba which also occur in Central America arc predomina ntly in the Pinar del Rio province with less num bers successively in Oriente and the Isle of Pines and generally di minish in number!. in an easterly progression. Those genera of Cuba wh ich are also found in South America follow the same pattem as tha t for Central American genera in Cuba. Two well-known and striking cases of disj unct distribution of great distance are the genera Cneorum and Dracena. The genus Cneomm (Cncoraceae) is found in the West Indies only in the Sierra Maestra range on the south side of the Oriente prov ince in Cuba. lt is represented elsewhere in t he Canary Islands and the Mediterra nean region. The species Dracena cubensis (Agavaceae).is found in the Sierra de Moa range on the north coast of the Oriente province in Cuba with additional species of the genus occurring in Central America and in Africa. The island of Hispaniola cannot be d ivided into easily recognized and named provinces comparable to those of Cuba. Province or area names and boundaries have been changed frequently in the Dominican Republic following various pol itical reorganizations. The accompanyin3 map ( Fie.7) delimits nine geographic zones

T I U: LOCA'J10N ANO (ii:OG itAI,III C KAN(il: 01' SELE(.TI:O GENERA IN Til l.: CUUAN FLOKA

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- ---·---.. Pinar del Rio Isle of Pines Havana Matanzas Las Vi llas Camagucy Oriente

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the vegetation of Hispaniola. Severa l of these represent isolated mountain ranges o!Tering ltSLia l plant formations as the southern peninsula of.Haiti extending into the Barahona province, the Cordillera Central and the Samana peninsula. Asprey and Robbins (1953) call attention to areas of specialil!ed vegetation in Jamaica as does the work of Gleaso n and Cook ( 1926) for Puerto Rico. It is possible to ta bulate the endemic genera as well as those genera indigenous to Cuba and Hispaniola cited in Tables XI , X! f, XXVI and XXV!f in their geographic position on lhe islands. The vertical columns in Table XXX represent the geographic distribution of genera within Cuba. Genera which occu r in more than one province are credited to each province. The dominance of endemic genera in the Pinar del Rio and Oriente provinces is striking. The ind igenous genera with extra-territorial relationsh ips outside the Greater Antilles are a lso more abundant in Pinar del Rio and Oriente than in other provinces. Genera which are foun d in both Cuba and Hispaniola of the Greater Antilles show a nearly uniform distribution across the length of Cuba. The Oriente province, that is closest to Haiti, does not dominate the pattern of distribution as might be expected. While the distribution by province is more uniform for genera ocfurring i11 Jamaica and Cuba, there are more example~ cited for Pinar del Rio and for Oriente than for the int~rvening provinces. The pal m gen us Gaussia is the only genus found only in Cuba and Puerto Rico. It is found in the Pinar del Rio province, the westernmost of Cuba and in the limestone western districts of Puerto Rico. The two genera which share a pattern of Cuba and the Bahama~ are found as expected on the north coastal districts in all provinces of Cuba. Four genera are cited which occur in Cuba and the Lesser Antilles. Two of these occur in Cuba and in the Oriente province alone, one in Pinar del Rio alone and one in the Pinar del Rio and Havana provinces. Of genera occurring in Cuba and the United States, sixteen occur in Pinar del Rio, ten on the I sle of Pines, five in each of

-------~--- - - -· 5 i I ,............--- ·- ·- ·- ·- ·- ...._f 1 ·- -

..........

~. - .........

'-...

.:7

·,

. _ _/

. / --·

·,j

---·- ·-" - ·-·-t

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0 Fig.7. Geographic districts of Hispaniola as significant areas of endemism. F or explanation, see text.

combining several of th ose recognized by Moscoso (1943). In Fig.7, I is the southern peninsula of Haiti (Massif de !a Hotte, Massif de la Salle) extending eastward as the Sierra de Bahoruco range; 2 is the Cui de Sac in Haiti and the Enriquillo Basin in the Dominican Republic; 3 is the Cordillera Central or a central chain of mountains; 4 is the northern mountain range of Haiti, (Montagnes du Nord Ouest, Massif du Nord); 5 is the Rio del Yaque valley extending into the Cibao near Santiago; 6 is the Cordillera Septentrional ; 7 is the valley of the Rio Yuna; 8 is the plains of BayaguanaHato Mayor ; 9 is the Samana peninsula.

T H E VEGETATION OF T HE ANTI LLES

T A LILE XXX I

be fou nd on nearly every land mass in t he Ca ribbea n and some are regarded as large genera with many species within t he area. There are some disj unct dist ri b.utio n patterns rep resen ted by genera in tropical A merica and these have been commen ted upon and have often been the basis of speculation involving continental drift, lo ng-range dispcrs<d by sea or by birds or even land b ridges. T hey have not, however, been considered o n the basis of representatives within the Antj lles . · The suggesti ons o f Svenson (1933) and others of d istan t relationships between the floras of the A ntilles and the Galapagos 1slands have n ot been able t o withsta nd the scrutiny of closer in vestigation. Al though these fl oras have genera in common, the d isjunct ranges of species previously reported are now discounted by the correct naming of the specimens or by the recent collectio ns of the species in intermediate areas making the gaps in distribution less significant. T he most recent summary work on pla nt distribution is that of Good (1964): In an appendix he considers genera found ent irely or predominantly in the tropical regions, but exclud ing pan-tropical genera. He gives fo ur lists of genera which occur in the fo llowing pallerns; (1 ) America, Africa and/or M adagascar; (2) America and continenta l Africa ; (3) America and Madagascar only; (4) A merica and Asia. Of the thirty-one genera Good cites having a distribution of America, Africa and/or Madagascar the following twelve genera occur in the West Indies. Following each generic name l have indicated the total number of species cu rrently recognized in the genus (Willis, 1966) followed by t he number o f species recogn ized within the A ntillean flora. Some personal tax onom ic judgemen t is employed in the number of taxa recognized. Bertiera (R ubiaceae) 30/2; Caperonia (Eupho rbiaceae) 60/3; Carpodiptera (Tiliaceae) 80/4; Cassipourea (Rhi zophoraceae) 80/7; Eichhornia (Pontederiaceae) 7/5; Hirtel/a (Chrysobalanaceae) 95/3; Paul/inia (Sapindaceae) 180/8 ; Piriqueta (Tu rneraceae) 20/5; Sabicea (R ubia.ceae) 130/3 ; SaPia (Eu pho rbiaceae) 3 1/ 12; Symphonia (Guttiferae) 21/1; Trichi/ia (Meliaceae) 300/20. Good lists 66 genera as examples o f a distt ibution pattern of America and continental Africa . Twen ty-five of these genera have representatives in the Antilles. A manoa (Euphor biaceae) 7/ 1; Andira 75/1; Brachypteris (Malpighiaceae) 3/2; C!tlorophora (Moraceae) 12/1 ; Chrysobalanus (Chrysobalanaceae) 2/2; Coi!OC0/1Jus (Combretaceae) 2/1; Copaifera (Leguminosae) 25/1; Genlisea ( lentibulariaceae) 15/1; Gum·ea (Meliaceae) 150/7; Heisteria (Olacaceae) 50/ 1; Heteropteris (Malpighiaceae) 100/9; Laguncularia (Combretaceae) 2/ 1; Machaerium, as Drepanocorpus (Leguminosae) 150/ 1; Ma!ouetia (Apocynaceae) 25/ 1 ; Microtea (Phytolaccaceae) 10/2 ; Mayaca (Mayacaceae) 10/ 1; Parkinsonia (Legum inosae) 2/ 1; Quassia (Simaroubaceae) 40/1; Renealmia (Zingiberaceae) 75/ 10; Schultesia (Gen tianaceae) 20/2; Struchium as Sporganophorus (Compositae) 1/ 1; Talinum (Portulacaceae) 50/2 ; Tapura (Dicbapetalaceae) 20/4; Thalia (M arantaceae) 11/ 1 ; Voyria (Gentianaceae) 15/5. Four genera a re cited by G ood showing a distribution of America and Madagascar only. The genus Rheedia (Guttiferae) is now considered a section of Garcinia and

Tl-11.: LOCATION ANO GEOGRAI'IIIC RAh:GE Of SELECTED (ii:N t:I<A IN Tilt; FLORA Of III SI•ANIOLA

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R. A. HOWA RD

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Each of these areas rep resent a reas of speciation. In a n analysis of the selected examples o f genera of Hispaniola and their d istribution in- Latin America, several interesting patterns of plan t distribution are evident (Table XXXI). Endemic genera a re fou nd most often in the limestone mou ntains o f northern and southern Hispani ola (Fig.7,/,4). The indigenous- genera are foun d most fre quently in the sou thern peninsula area, 1, and the Cordi llera Central, 3. Genera with ranges extending into Cuba alone are most abu ndant in the southern peni nsula, 1, and to a less amount in the northern peninsula, 4.. and the dry valley, 2. N o conclusive pattern occurs in the ge nera of Hispani ola fo und also in Jamaica or the Bahamas while genera which occur also in Puerto R ico occur most frequen tly in eastern Hi spaniola. Genera with a geographic range reaching the Uni ted States occur in the sout hern peninsula, 1, and the centra l range of mounta ins, 3. The same pattern exists fo r genera of Hispan iola which also occur in the Lesser Antilles, in Central America o r in Sout h America.

Long-distance relationships and disjunct distribution patterns A number of large families of flowe ring plants are recognized as being primarily t ropical in their d istribution and of these the Acanthaceae, Ebenaceae, Euphorbiaceae, Flacourtiaceae, Ges neriaceae, Melastomataceae, Moraceae, Myrtaceae, Piperaceae, R ubiaceae, Sapotaceae, and Urticaceae are indeed well represented in the Antillean vegetation. Such genera as Casearia (Fiacourtiaceae), Cassia (Leguminosae), Chrysophyllum (Sapotaceae), Croton (Euphorbiaceae), Colubrina (Rhamnaceae), Diospyros (Ebenaceae), Eugenia (Myrtaceae), Ficus (M oraceae), Omphalea (Euphorbi aceae), Pilea (U rticaceae), Peperomia (Piperaceae) and Psychotria (R u biaceae) will

R. A. HOWARD TH E VEGETATIO N O F T H E ANTILLES

therefore a genus primarily of Asian distribution. Two genera have species occurring in the Antilles. Paepafmrthus (Eriocaulaceae) 450/ 10; and Pedifa11tftlrS 14/1. To these examples can be added such genera as Andira (Leguminosac) 75/1; Boucltea (Verbcnaccae) 39/ I ; Diocfea (Leguminosae) 50/3; Hafimium (Cistaceae) 16/2; Jatropha (Euphorbiaccae) 175/9; Kostefetzkya (Malvaccae) 30/6; Pitcaimea (Bromeliaceac) 250/1 1; Pril'{/ (Verbenaceae) 28/3; Rllipsalis (Cactaceae) 60/3; Ximenia (Oiacaccae) 19/4; Tetrorclridiw11 (Euph orbiaceae) 16/ I (Brcnan, 1965). The majority of the gel)cra cited with American-A frican disjunct distribut ion arc New World genera represented in the Antilles by single species and in Africa by single species. Sympho11ia (Guttiferac), Amanoa (Euphorbiaceae), Heisteria (Olacaceae) occur in the Lesser Anti lies and are not represented in the Greater Antilles. Bertiera (R\Ibiaceae) is foun d only in Hispaniola. Ge11/isea (Lentibulariaceae) and Ma/ouetia (Apocynaceae) are found only in Cuba, Tetrorchidium (Euphorbiaceae) is fou nd only in Jamaica. All othe1 genera are represented iri both the Grea ter and the Lesser Antilles. T he genus Symphonia (Guttiferae) is the only genus cited which has more species in Africa and Madagascar than occu r in tlw New World. The importance of Quassia (Simaroubaceae) may be questioned since the plant is both ornamental and medicinal and often cultivated and may have been introduced and persisted following cu lt ivation. The distribution and variation _of Rhipsa/is (Cactaceae) in Africa has been the subject of much discussion concerning its introduction. The problem has had advocates in theories of distribution by birds, man and continental drift, and the study Camp (1948) suggested remains to be done. Equally troublesome and unsolved is the occurrence of Pitcaimea (13romeliaceae), a large tropical Am~r ican genus in West Africa. Good cites forty genera having a disjunct distribution of America and Asia, often extending into Australasia and the Pacific Islands, and the following sixteen genera have representatives in the Antilles. T he numbers given again represent respectively the total number of species in the genus and the number in the An-tilles. Cal/icarpa (Verbenaceae) 140/17; Capsicum (Solanaceae) 50/2 ; Cedrela (Meliaceae) 6/1; Dendropauax, as Gilibertia (Araliaceae) 60/13; Heficteres (Sterculiaceae) 60/3; Ieiman thus (Graminae) 26/5; Laplacea (Theaceae) 30/1 1; Meliosma (Sabiaceae) 100/7; Mifl·eola (Loganiaceae) 6/2; Nefumbo (Nelumbonaceae) 2/ 1; Sapindus (Sapindaceae) 13/1; Schoepfta (Olacaceae) 35/ 7; Sloanea (Eiaeocarpaceae) 120/13; Symplocos (Symplocaceae) 350/21; Ta/auma (Magnoliaceae) 50/4; Turpinia (Staphyleaceae) 40/4; and Xylosma (Flacourtiaceae) 100/ 15. To this list can be added a number of genera which are well developed in Asia, although theit range may be pan-tropical or extend into temperate areas. The occurrence and development of these genera in the Antilles is of comparative interest. Antirlrea (Rubiaceae) 40/24; Chrysophyllum (Sapotaceae) 150/7; Casearia (Flacourtiaceae) 160/21; Catalpa (Bignoniaceae) 11/2; Cephaelis (Rubiaceae) 180/4; Clethra (Ciethraceae) 120/4; Dacryodes (Burscraceae) 50/l; Ehretia (Boraginaceae) 50/1; Hernandia (Hernandiaceae) 20/3; J..asianthus (Rubiaceae) 150/ 1; Linociera (Oleaceae)

100/ 10; Podocarpus (Podocarpaccae) I00/8 ; Protium (Burseraceae) 90/3; Scae•·ola (Goodcniaccae) 100/ 1; Scrophufaria (Scrophu lariaceac) 300/6 ; Styl'frx (Styracaceae) 130/4; SlrJ:clmos (Loganiaceac) 200/1; and Viburnum (Capri foliaceae) 200/4. Although these genera arc represented by one to twen ty-one species within the Antilles, all bu t Lasicmtlrus (Rubiaceae) arc better represented within Central or South America than in the Antillc~. Scae•·o/a (G oodeniaceae) in the New World is represented by a si ngle coastal species weli distrib uted around the Caribbean periphery, but poorly reprc~ented in South Ameri ca, although also found on the coast of Africa. Sapindus (Sapindaceae) has had an economic use, and its distribution may be the result of its usefulness to man. Dacryodes (Bu rsc raccae) is t he sole genus rest ricted to the Lesser Antilles in its distribution within the Caribbean archipelago.

Conclusions

The Caribbean islands form an archipelago of abou t 1,000 islands havi ng a con~ider able range in size, alt itude, soil types and environmental niches. The total fl ora is estimated to be 8,000 species and the flo ra as a whole is in great need of monographic treatments of all groups. Such studies must consider the frequent distribut ion of the genera in Central America and Sou th America as well as the Greater and the Lesser Antilles. The larger islands of the Greater Antilles have the largest number of species and the greatest amount of endemism. On the larger islands, especially Cuba and Hispaniola, there are geographic areas of high speciation and these are largely areas of mesophytic to xerophytic environments. Al though there are many small endemic genera restricted to one or a few islands, on ly two genera of more than twenty species each appear to be limited to the Anti lles. The family Picrode ndraceae with one genus and three species is the sole family restricted to the Antilles. Patterns of disjunct distribution are abundant, involving the Greater Antilles and the Lesser Antilles, the Greater Antilles and Central America, and the Greater Antilles and South America (Fig.3). There are also example~ of relationships of the Anti llean flora to Africa and to Asia. Paleobotanical studies in the Antilles are not numerous. The identification of fossil materials, however, will be a major effort in view of the complex relationships and distributional patterns of the existing vegetation.

References Alain, H., 1962. Flora de Cuba, 5. Editorial Universita ria, Rio Piedras, Puerto Rico, 362 pp. Asprey, G. F. and Robbins, R. G., 1953. The vegetation of J amaica. Ecol. Monogr., 23 :359-412. Bannister, B. A., 1970. Ecological life cycle of Euterpe globosa Gaertn. ln: H. T. Odum (Editor},~A Tropical Rain Forest. U.S. Atomic Energy Comm., Oak Ridge, Tenn., pp. 299-314. Beard, J. S.,l944. Climax vegetation in tropical America. Ecology, 25:127- 158.

R. A. HOWARD

Ocard, J. S., 1949. T he natural vegeta tion of the Windward and Leeward fslands. 0 .1 jo rd For. Mcm., 192 pp. lleanJ, J. S., 1955. The classific:llion of tropical American vegetation-types. Ecolofly, 36 :89- 100. Brcna n, J. P. M., 1965. The geographical relationships or the geriera of Lcguminosae in tropical Africa. Wcbbia, 19: 545-578. Britton, N. L. and Millspaugh, C. F. , 1920. The Bahama Flom. New York Botanical Ga rden, New York, N.Y., 694 pp. Britton, N. Land Wilson, P. , 1923/ 1924; 1925/1930. Scientific Strrl'c)' of Porto Rico and the Virgin Island;·, 5, 6. New York Acad. Sci., New York, N.Y., 5 (1923/ 1924): 626 pp; 6 (1925/1930): 663 pp. . Camp, W. H., 1948. Rhipsalis- and plant distribtltions in the Southern Hem isphere. J. N.Y. Bot. Card., 49:33- 38. Gleason, H. A. and Cook, M. T ., 1926. Plant Ecology of Porto Rico- Scientific Swwy ofPorto Rico and the Virgin Islands, 7. New York Acacl. Sci., New York, N .Y. , 96 pp. Good, R., -1964. Tile Geography ofthe Flowering Plant;·. Wiley, New York, N.Y., 518 pp. Guilding, L., 1825. An Account of the Botanic Garden in the lslaud of St. Vincent. Glasgow, 47 pp. (Now from Arnold Arboretum, Jamaica Plain, Mass.) Hodge, W. H., 1954. Flora of Dominica, B.W.L, I. Lioydia, 17:1- 238. Howard, R. A., 1953. Botanical Gardens in West Ind ies Histo ry. Card. J., 2: 11 7- 120. Howa rd, R. A., 1954. A history of the Botanic Garden of St. Vincent, British West Indies. Geogr. Re1•., 44:38 1-393. · Howa rd, R. A., 1962. Volcanism and vegetation in the Lesser Antilles. J. Amold Arboretum, 43: 279-3 11. Howard, R. A., 1968. The ecology of an elfin forest in Puerto Rico, I. fnt roduction and compositio n studies. J. Amold Arboretum, 49: 381-418. H oward, R. A., 1969. The ecology o f an elfin forest in Puerto Rico, 8. Studies of stem growth and for m of leaf s tructure. J. A m old Arboretum, 50:225-262. Howard, R. A. and Briggs, W., 1953. The vegetation on coastal dogtooth limestone in southern Cuba. J. Amold Arboretum, 34:88- 94. Howard, R. A. and Powell, D. A., 1963. The identifiCation of rubber-producing species in the West Indies. Ecan. Bot., 12:337-349. . Howard, R. A. and Proctor, G. R ., 1957. The vegetation on bauxite soils in J amaica. J. Amold Arboretum, 38: 1-41; 151-169. JacqLLin, N.J., 1797. Plant. Rar. Horti Schiinbrwm, Vietma, I: viii- xii. K irby, I. A. E., 1971. Pre-Columbian lndiam in St. Vincent, West Indies. St. Vincent Archaeol. and Hist. Soc., Kingstown, St. Vincent, 6 pp. • l eon, H., 1946. Flora de Cuba, I. Coutrib. Occas. Col. de Ia Salle, 8: 1- 441. l eon, H. and Alain, H., 1951. Flora de Cuba, 2. Coli/rib. Occas. Col. de Ia Salle, 10: 1-456. Marie-Victorin, F . and Leon, F., 1942, 1944, 1956. ltineraircs botanique dans l'i le de Cuba. Coutrib. !ns f. Bot. Univ. Mont real, 41:496 pp; 50 :410 pp; 68:227 pp. Moscoso, R. M., 1943. Catalogus Florae Domingeusis. Univ. Santo Domingo, Santo Domingo, 732 pp. Proctor, G . R., 1970. Mason River. Jamaica J., 4:29-33. Sauer, J. S., 1967. Geographic reconna issance of seashore vegetation a long the Mexican Gulf coast. Coastal Stud. lnst. La. State Uuiv., Botou Rouge, Tech. Rep., 56:59 pp. Seifr iz, W., 1943. T he plant life of Cuba. Ecol. Mouogr., 13:375-426. Stehle, H., 1945. Fo rest types of the Caribbean Islands. Caribb. For. , 6(suppl.) : 273-408. Stehle, H ., 1946. Les types forestiers des lies Caraibes. Caribb. For. , 7(suppl.) : 337-709. Svenson, H: K., 1933. Vegetation of the coast of Ecuador and Peru a nd its relation to the Galapagos Islands. Am. J. Bot., 33:394-498. Willis, J. C., 1966. A Dictionary of the Floweriug Plauts and Fems ( Revised by H. K. Airy Shaw.) University Press, Cambridge, 12 14 pp.

Chapter 2

A Paleoclimatic Interpretation of the Eocene Floras of Southeastern North America DAVID L. DILCHER Deiwrtmeu/ of Botany, Indiana i!ttil•er;'ity, Blooming/Oil, Ind. (U.S.A.)

Summary The Eocene flo ras of the Mississippi embay ment are well known through the p ublished works of Berry {1916, 1924, 1930, 1941). As part of a rei nvestigation of these Eocene floras in western Kentucky and Tennessee, thei r age, d epositional environment and paleoclimate arc considered. The age of the Lower Eocene Wilcox flora in western Kentucky and Tennessee is revised to M idd le Eocene, Cla iborne Formation, primarily on the basis o f pollen data. Berry interpreted the depositional environment of these deposits to be tropical or subtropical coastal-strand with deposition in near-shore lagoons or backwater areas. The leaf-bearing clay deposits in western Kent ucky and Ten nessee a re here recognized as oxbow lake sed iments, possibly laid down some distance from the actual embayment. The nature of the climate is critically examined by three approaches: (/) identifying ind iv id ual fossi ls to their nearest li vi ng similar forms a nd basing paleoclimatic interpretations o n the climatic range of these modern form s; (2) identifying fossils to similar living forms and analyzing the climate of the commu nities in which these living similar forms are fo und today; and (3) using the form of the fossil vegetation, its fo liar p hysiognomy, and relating this to modern climates where vegetation wit h a similar foliar physiogno my is found. Each of these approaches is analyzed in terms of its usefulness for understanding the paleoclimate of t he Middle Eocene in the upper portion of the Mississippi embayment. The fi rst two approaches are o f some value if their limitations are recognized and they are used with care. Foliar phys iognomy, when both leaf-margin analysis and leaf-size analysis are combined with temperature and moisture data, provides an important index of m odern climates and paleoclimates. The response of modern vegetation to climatic zones is discussed and the value of foliar physiognomy indexes evaluated. The approximate paleoclimate of this flora is concluded to be seasonally dry to slightly moist moisture regime, and an equable warm temperate to cool subtropical temperature regime. T he paleoclimatic conclusions drawn are based on all the available data. However, they are only approximations, since our present knowledge of the evolution of plants and plant communities and of the relatio nship of foliar physiognomy and climate is not refined enough to allow a precise statement of Early Tertiary climates .