Historical biogeography of Caribbean plants: introduction ...geographic history of the Caribbean biota have ignited much interest, resulting in many studies addressing ... tributions

INTRODUCTIONFor years studies on the classification evolution and

geographic history of the Caribbean biota have ignitedmuch interest resulting in many studies addressingissues in a variety of contexts Most of the works on bio-geography of the region are predominantly on animalgroups and there is a marked deficiency in similar stud-ies for plant groups For instance in the proceedings of asymposium (Morin 1982) emphasizing the phytogeogra-phy of Central America in which issues about theCaribbean islands were addressed two of the most rele-vant contributions (Humphries 1982 Savage 1982) dis-cussed only vertebrate groups One of the authors(Humphries 1982) acknowledged that the number ofplant studies then available were very scarce andbecause of the lack of phylogenies were unsuitable toapply in his cladistic methods

A synthetic compilation on West Indian Bio-geography (Woods 1989) includes 32 works on paleon-tology anthropology ichthyology entomology ornithol-

ogy herpetology and conservation biology In these pro-ceedings the editor emphasized that ldquohaving reviewedmuch of the available data on West Indian biogeographyI have been struck by how few studies on plants andinvertebrates there are in the overall body of data onWest Indian biogeography compared with the informa-tion available on other groupsrdquo (Woods 1989) Thoseldquoother groupsrdquo mentioned by Woods are primarily verte-brates which at that time formed the predominant bodyof information for interpreting the biological history ofthe West Indies In that symposium only one of the 50contributors (Adams 1989) addressed ideas on the geo-graphic origin and evolution of a plant group(Juniperus) A second contribution (Woods amp Sergile2001) includes only one biogeographical study (Judd2001) of a Caribbean plant group (Lyonia sect Lyonia)This disparity between knowledge on the evolution andbiogeography of plants relative to animals is a primeindication that our ideas on the history of the Caribbeanbiota have been erected almost exclusively from faunaldata

299

Santiago-Valentin amp Olmstead bull Historical biogeography of Caribbean plants53 (2) bull May 2004 299ndash319

Historical biogeography of Caribbean plants introduction to current know-ledge and possibilities from a phylogenetic perspective

Eugenio Santiago-Valentin12 amp Richard G Olmstead1

1 Department of Botany University of Washington Seattle Washington 98195 USA olmsteaduwashingtonedu (author for correspondence)

2 Present address Jardiacuten Botaacutenico de Puerto Rico Universidad de Puerto Rico Apartado Postal 364984 SanJuan 00936 Puerto Rico goetzeayahoocom

Knowledge of the evolution and biogeography of the Caribbean biota comes primarily from faunal studies inspite of the tremendous richness of the Caribbean flora The limited data to assess Caribbean phytogeographycomes from fossils floristic distributions and phylogenetic studies The geology in the Caribbean is extreme-ly complex but it is widely accepted that part of the Greater Antilles are old and precede the emergence of theLesser Antilles and the Panamanian isthmus Cuba and Hispaniola are each composed of a series of independ-ent land blocks and Jamaica was mainly or entirely submerged during part of its history Paleopalynologicaldata at present the only reliable source for reconstructing past plant communities suggest warm-temperate totropical conditions Contemporary floristic data portray affinities with both Laurasian and Gondwanan derivedplant groups Shortcomings of traditional methodological approaches is a major criticism to studies assessinghistorical biogeography Phylogenetics is an appropriate strategy to apply because it (1) elucidates the sys-tematic relationships of taxa (2) permits assessment of morphological and ecological evolution and (3) illus-trates the direction and sequence in which the distribution of a taxa originated Study of Caribbean taxa withhigh levels of endemicity allows tracing the correspondence between taxa and geographic areas and thus per-mits application of a phylogeny for assessments of geographic relatedness under both dispersal and vicariantscenarios Caribbean plant groups in several angiosperm families have now been studied phylogeneticallySome biogeographic patterns are starting to emerge from these studies but more studies are needed for gener-alizations to be drawn

KEYWORDS Antilles biogeography Caribbean phylogeny West Indies

Historically zoological studies have dominated thescene of Caribbean evolution and biogeography This isstill the case today where new contributions range overa diversity of vertebrate groups including birds (egErard 1991 Bermingham 1994) mammals (eg Breuilamp Masson 1991) and especially reptiles (eg Savage1982 Hedges amp al 1991 Roughgarden 1995 Hass1996 Seidel 1996 Jackman amp al 1997) and amphib-ians (eg Bogart amp Hedges 1995 Hedges 1999)Hedges (1996) reviewed the current knowledge on thehistorical biogeography of West Indian vertebratesZoogeographic studies of the region have diversified toinclude invertebrates (eg Liebherr 1988 Schubart ampal 1998) Although these numerous zoogeographic con-tributions have made the Caribbean one of the most fer-tile regions for the study of biogeography a comprehen-sive understanding of the evolution of the biota of theregion can only be achieved by inclusion of other groupsincluding plants

Interest in applying phylogenetics to study the his-torical biogeography of Caribbean plants has increasedin recent years as evidenced by published works (egJudd 1981 2001 Zona 1990 Lavin amp Luckow 1993Skean 1993 Salzman amp Judd 1995 McDowell ampBremer 1998) and a recent symposium (Fritsch amp Mc-Dowell 2003) In order to stimulate further interestamong students and newcomers we present in this workan outline on the present understanding of Caribbeanphytogeographymdashwith emphasis on the Greater AntillesWe provide general summaries on fossils floristic dataand with emphasis on case studies of Caribbean plantgroups where a phylogenetic approach has been incorpo-rated We also discuss considerations for applying thisapproach to test hypotheses on evolutionary processesand biogeography in other plant groups of the region

Several names are used to define different but over-lapping geographic regions mentioned in this reviewHence we define their use in this work for clarity Theterms ldquoGreater Caribbeanrdquo or ldquoCaribbean Basinrdquo refercollectively to the islands and the neighboring continen-tal regions along the Gulf of Mexico Central Americaand Northern South America The main islands of theCaribbean belong to three different archipelagos TheGreater Antilles the Lesser Antilles and the Bahamas(Fig 1) The Greater Antilles are composed of the fourlargest islands in the region Cuba Hispaniola Jamaicaand Puerto Rico The Lesser Antilles is separated fromthe Greater Antilles by the Anegada Passage and com-prise the arc of smaller islands extending from Sombreroin the north to Grenada Collectively the three archipel-agos are referred to as the Caribbean Islands or the WestIndies This paper covers issues relevant to the entireCaribbean with emphasis on the island systems

PLANT DIVERSITY IN THECARIBBEAN

The Caribbean is the oldest Neotropical region stud-ied by western science The first observations on plantdiversity are documented in Columbusrsquo diary and otherchroniclers of the 15th and 16th centuries with plantdescriptions predating Linneausrsquo Species Plantarum of1753 The long plant collecting history has resulted innumerous new taxa as well as published Floras includ-ing among the most recent ones for Cuba (Leoacuten 1946Leoacuten amp Alain 1951ndash1957 Alain 1963 Liogier 1974)Hispaniola (Liogier 1982ndash1996) Jamaica (Adams1972) Puerto Rico and the Virgin Islands (Acevedo-Rodriacuteguez 1996 Liogier amp Martorell 1982) theCayman Islands (Proctor 1984) the Bahamas (Correll ampCorrell 1982) and the Lesser Antilles (Howard1974ndash1989) Ongoing botanical explorations keep refin-ing present knowledge with the publication of florulasand checklists the report of new records and thedescription of new species

Plant diversity is so remarkable in the Caribbeanislands that the region is considered a distinctive phyto-geographic unit within the Neotropics (Gentry 1982)The vascular plant flora of the region consists of approx-imately 12000 species including the southern tip ofFlorida (Myers amp al 2000 Table 1) in about 200 fami-lies Over fifty percent of the vascular species are endem-ic to the region These endemics represent about 2 ofall vascular plants on Earth placing the Caribbean as oneof the leading hotspots in terms of species-levelendemism (Myers amp al 2000) Endemicity is also pres-ent at the generic level in the Greater Antilles Bothgeneric and species-level endemism are particularlyprominent in the two larger islands of Cuba andHispaniola (Gentry 1982) Cuba the largest island in theCaribbean has the richest flora and highest proportion ofendemism (Table 2) About half of its approximately

Santiago-Valentin amp Olmstead bull Historical biogeography of Caribbean plants 53 (2) bull May 2004 299ndash319

300

Fig 1 Map of the Caribbean region

6000 species of flowering plants and approximately 70genera are unique to the island Hispaniola the secondlargest island contains over 4500 flowering plantsalmost a third of which are endemic (T Zanoni perscomm) and about 30 endemic genera Diversity in thesmaller Jamaica and Puerto Rico is less and endemism isprimarily at the species level The Jamaican floweringplants total over 2500 species with about 30 endemic-ity whereas not more than 10 of Puerto Ricorsquos approx-imately 2000 native flowering plants are endemic

The relatively high plant diversity observed in theCaribbean is explained in part by proximity to the conti-nental Neotropics to the south (South America) west(Mesoamerica) and north (Florida) In addition the widerange of altitudes the contrasting rainfall and tempera-ture regimes and the diverse geology have resulted instriking habitat mosaics even within the same island per-mitting the occurrence of plants species with very differ-ent ecological requirements and tolerances ManyAntillean taxa are endemic to serpentine rocks Suchsubstrates in the Greater Antilles particularly the ones inCuba and Puerto Rico (Cedentildeo-Maldonado amp Breckon1996) are regarded as among the most important of trop-ical America (Brooks 1987) In Cuba 135 of theendemic species (Berazaiacuten-Iturralde 1976 1981) and343 of the endemic genera are confined to serpentine

(Brooks 1987) Antiquity of portions of the GreaterAntilles (of approximately 70 to 100 my Donovan ampJackson 1994) could also be a factor to explainendemism in the area

Comparisons between the Caribbean andother areas mdash Numerically the approximately12000 vascular plants present in the Caribbean are com-parable to other important centers of plant diversity suchas Madagascar and New Caledonia (Table 1) TheCaribbean has roughly the same number of speciesknown today for Madagascar and has more than threetimes the number of species of New CaledoniaFurthermore generic and species diversity in the regionis also higher than in other oceanic archipelagos (Table3) although the latter have considerably less land areawhen compared to the Caribbean Species endemism inthe region is higher than in Central America ~50 ver-sus ~20 respectively (Gentry 1982 Myers amp al2000) It is also higher than in other islands (Table 1)such as the Philippines (39) Borneo (30) Sumatra(12) Java (5) and Japan (4) The total number ofendemics in the Caribbean islands is several timesgreater than in the Hawaiian Islands although the latterhold a higher proportion of endemics per unit area (Table3) Similarly it comprises ten times the endemic speciesthan in the Canary Islands more than twenty times what


301

Table 1 Diversity of vascular floras in selected large islands and archipelagos Estimates are based on WWF amp IUCN1994 1995 1997 and Myers amp al 2000

Land area Max Endemic Endemic Species Region (km2) elevation (m) Families Genera genera Species species endemismMadagascar 594150 2876 160ndash181 gt1289 260 12000 9704 809New Caledonia 18600 1628 191 863 110 3332 2551 765Caribbean Is 263500 3087 186 2500 c 100 12000 7000 583

(incl S-Florida)Borneo 738864 4101 ndash ndash 61 ~25000 ~7000 ~28Philippines 300800 2954 ndash ndash 23 8931 3500 391Japan 376520 3776 229 ndash 17 5565 gt222 ~4Sumatra 427610 3805 ndash ndash 13 gt10000 gt1200 ~12

Table 2 Phanerogamic diversity and endemism in the Caribbean Islands Estimates are based on WWF and IUCN (19941995 1997) Myers amp al (2000) and P Acevedo (pers comm)

Land area Number Number of endemic Number ofArchipelago (km2) of species species () endemic genera

Greater AntillesCuba 110848 06015 03193 (53) c70Hispaniola 0 76190 ~4685 ~1400 (30) c30Jamaica 0 10830 02746 000852 (31) 007Puerto Rico 00 9308 02128 000215 (10) 001

Lesser Antilles 006466 ~2300 00~290 (3) 000Bahamas 010010 01068 000117 (11) 000 = Does not include the offshore islands north of South America

is reported for the Galaacutepagos and over fifty times that ofthe Juaacuten Fernaacutendez Islands At the generic level theCaribbean comprises about the same number of endemicgenera found in Central America and roughly the sameamount in New Caledonia (Table 1) Endemic genera inthe region surpass the total found in Borneo (61) thePhilippines (23) Japan (17) Sumatra (13) and Java (5)Furthermore Caribbean-endemic genera triple theamount found in Hawaii and are nine to ten times theamount for the Galaacutepagos the Juan Fernaacutendez and Fiji(Table 3)

GEOLOGIC HISTORY OF THECARIBBEAN

Although there are still diverse interpretations onkey aspects of Caribbean geologic history we presentbasic geologic information that will allow readers unfa-miliar with this area to formulate an outline on the sub-ject A review including the historical development ofgeohistorical models for the Caribbean is out of thescope of the present work For this purpose we recom-mend Hedges (1996) Iturralde-Vinent and MacPhee(1999) and Graham (2003) The geologic history of theCaribbean is extremely complex and basic issues suchas which areas were above sea level during the history ofthe islands (Hedges 1996) remain unresolved

The Greater Antilles and the younger Lesser Antillesare part of the Caribbean Plate which originated from aportion of the East Pacific plate during the Jurassic-Cretaceous (Pindell 1994) The North and SouthAmerican separation from Laurasia and Gondwanarespectively involved a westward movement towards thePacific which resulted in the intrusion of part of the EastPacific Plate in the area between both continents Thisintrusion eventually fractured and formed the smallerCaribbean Plate Some authors (eg Malfait ampDinkelman 1972 Sykes amp al 1982 Pindell amp Barrett1990) consider that portions of what would form theGreater Antilles termed Proto-Greater Antilles arose

during the Cretaceous as an arc of volcanic islands creat-ed from subduction of oceanic lithosphere locatedbetween North and South American continental plates(Ladd 1976 Coney 1982) Early in the Tertiary com-pression of the westward-moving North and SouthAmerican plates resulted in a relative eastward move-ment of the Caribbean plate and thus drift of the Proto-Greater Antilles (Rosen 1976 1985) During that timesome portions of the Greater Antilles reached their pres-ent-day positions (Coney 1982 Pindell amp Dewey 1982Sykes amp al 1982) as the result of the collision of thenortheastern boundary of the Caribbean plate with theBahamas platform which is fixed to the North AmericanPlate This collision caused subduction and volcanism inthe Proto-Greater Antillean area although orogeny fromfolding faulting and local uplift continued (Lewis ampDraper 1990) Subsequently a major fault was formed tothe south of Cuba along with the small spreading center(the Cayman Trough) that could have influenced theeastward drift of Southwestern Hispaniola and JamaicaToday the Caribbean Plate is still moving eastwards withrespect to North and South America with volcanic activ-ity along the eastern and western margins of the plates inthe Lesser Antilles and Central America respectively

Historical relationships and composition of the landmasses of the Greater Antilles is still very incompletebut it is widely accepted that Cuba and Hispaniola areeach formed by composite regions comprising no lessthan three land blocks (Pindell amp Dewey 1982 Sykes ampal 1982) each of which could have been unrelated toone another The western and eastern portions of Cubafor instance exhibit marked geological differences(Draper amp Barros 1994) the geology of the western por-tion is considered unique in the Antilles (Draper ampBarros 1994) probably due to its relationship with theNorth American Plate (Graham amp al 2000) On theother hand Eastern Cuba North-Central Hispaniola andPuerto Rico were probably connected as a single mag-matic arc during the Paleocene-Eocene (Draper ampBarros 1994) and until the Oligocene (Iturralde-Vinent1994) or Early Miocene (Sykes amp al 1982 Perfit amp


302

Table 3 Diversity of vascular floras in selected island systems Estimates are based on WWF and IUCN (1994 19951997) Marticorena amp al (1998) and Myers amp al (2000) excluding introduced taxa

Region Families Genera Endemic genera Species Endemic species

Caribbean Islands 186 2500 c 100 12000 7000Fiji 210 1013 12 1628 812Madeira-Salvagens 134 ndash 1 1226 123Hawaii 87 216 32 1200 1000Canary ndash ndash 20 1200 500Galaacutepagos 90 292 7 541 224Azores ndash ndash 1 300 81Juan Fernaacutendez 57 107 12 211 127

Williams 1989 Pindell amp Barrett 1990) Coalescence ofeastern and westerncentral portions of Cuba may haveoccurred in the Late Tertiary or Early Quaternary (Draperamp Barros 1994 Iturralde-Vinent amp MacPhee 1999)

Hispaniola is composed of four different land blocksaccording to Sykes amp al (1982) During the Eocene thenorth and central portions of Hispaniola fused and hadclose relationships with eastern Cuba a connection thatremained until Early to Middle Miocene (Graham2003) This long land relationship is invoked by Graham(2003) as a plausible reason to explain the high numberof Cuban-Hispaniola endemic plant genera Both south-western Hispaniola and Jamaica were considerably iso-lated from the rest of the Proto-Greater Antilles until theopening of the Cayman trench and posterior seafloorspreading (Perfit amp Heenzen 1978) moved them in anortheast direction until southern Hispaniola collidedwith the rest of that island probably early in the Miocene(Pindell amp Barrett 1990 Huebeck amp Mann 1991) orPliocene (Sykes amp al 1982) During drift to its present-day position most or all of Jamaica became submergedbetween Late Eocene to part of the Miocene followed byuplift in the Late Miocene (Draper amp Lewis 1990Robinson 1994) Evidence of this mid-Tertiary inunda-tion is the extensive karst topography and exposed lime-stone on much of the island The most recent addition toHispaniola comes via accretion of crustal terrane fromthe Bahaman Bank

Puerto Rico and the Virgin Islands experienced con-siderable eustatic changes although uplifts have appar-ently persisted in the Puerto Rican Bank since lateCretaceous (Lewis amp Draper 1990 Larue 1994) orMiddle-Late Eocene The island reached its current posi-tion during Late Eocene to Early Oligocene and separat-ed from Hispaniola in Oligocene or Miocene (Graham2003) Later during the Quaternary sea level changesdue to glacial-interglacial oscillations resulted in consid-erable modification of the outline and land extension inthe Puerto Rican Bank These changes are responsiblefor the separation of Puerto Rico from the Virgin Islandsand Anegada

The Lesser Antilles are considered as a classic islandarc of volcanic islands emerging from above a subduc-tion zone (Wadge 1994) They originated independentlyfrom the Greater Antilles due to the subduction of theAtlantic plate under the Eastern Caribbean Plate Thevolcanic activity that formed the Lesser Antilles startedat the end of volcanism in the Greater Antilles and per-sists today

In an attempt to bring together geological knowledgeand geographic scenarios of biological importance arecent contribution by Iturralde-Vinent and MacPhee(1999) presents a detailed paleogeographical and pale-oceanic reconstruction for the Caribbean Basin They

consider that the Greater Antillean land blocks that exist-ed previous to the Eocene very likely did not remain con-tinuously as subaerial land masses due to geologic activ-ity and to catastrophic events resulting from the KT bol-lide impact (Hildebrand amp Boynton 1990) Thus suchareas must have limited or no relevance for the biogeo-graphic assessment of the current Greater Antillean biotaBased on geological constitution geographical positionand physical paleogeography Iturralde-Vinent ampMacPhee (1999) infer that the current Greater Antillesmdashor at least portions of themmdashare no older than MiddleEocene Later during the Eocene-Oligocene transitionthe northern Greater Antilles and Northwestern SouthAmerica were connected by a landspan that involved theAves Ridge (ldquoGAARlandiardquo the Greater Antilles + theAves Ridge) The Eocene-Oligocene transition was aperiod of general uplift in this landspan followed byhigh sea levels (and presumably less connectionsbetween island areas) during Late Oligocene DuringEarly to Middle Miocene tectonic activity promoted fur-ther subdividions of Greater Antillean land blocks anevent of potential biogeographic relevance

FOSSILS AND PALEOVEGETATIONAvailability of fossil evidence mdash Plant fossil

data from the Caribbean are scanty several factors makethe reconstruction of paleoenvironments a very difficulttask (Graham 1989a) In a review of the status ofTertiary paleobotanical studies in Northern LatinAmerica (comprised of Mexico Central America andthe Antilles) Graham (1993) reported 59 studies specif-ic to paleobotany and paleopalynology in the Antillesbut only 16 of them include lists of identified plant fos-sils Of these several works are marred by impreciseidentification of the megafossils and are inconsistentwith other more recent evidence derived from fossilmammalian and marine invertebrate fauna 18O isotopestudies and plate tectonics (Graham 1988) Graham(1993) recommended their exclusion from any paleoen-vironmental interpretation until the material is restudiedor until new megafossils are available Palynologicaldata are at present the most reliable basis for recon-structing Tertiary communities in the Caribbean andNorthern Latin America (Graham 1988 1993 Grahamamp al 2000)

The Antillean paleocommunities mdash In gener-al Graham argues that most of the currently-knownTertiary plant fossils of the Antilles are warm-temperateto tropical with affinities to North America (includingMexico) (Graham 1990a 1990b Graham amp al 2000)The Mid-Eocene Chapleton deposit of Jamaica (Graham1993) and the approximately contemporary Sarama-


303

guacaacuten flora of east-central Cuba (Graham amp al 2000)are the oldest plant microfossil sites described for theregion (Graham 1993) The age of the Jamaican depositis supported by independent statigraphic and paleonto-logical evidence (Robinson 1988) The nature of the sed-iment deposition the presence of dinoflagellates mixedwith the pollen and spores and the presence of the pollenof Pelliciera and cf Acrosticum suggests that the depo-sitional environment at Chapleton was a warm-temperate(tropical) coastal one (Graham 1993) The sedimentsand fossil remains in the Saramaguacaacuten flora indicatethat it was a freshwater swamp or marsh community withwarm-temperate to tropical elements Graham amp al(2000) recognized 46 palynomorphs including 16 withNorth American affinity 11 endemics (unique to thatpalynoflora) and only one of South American originThe rest were widespread taxa Comparisons with othercontemporary fossil floras from the Antilles (egChapelton) and Central America are not feasible due tothe small size of those floras However this flora andones from NE Mexico and Panama are more similar thanany of them are to any contemporary South Americanfossil floras This is consistent with the hypothesis thatWest and Central Cuba are of North American continen-tal plate origin (Graham amp al 2000)

The Mid-Oligocene San Sebastiaacuten deposits in PuertoRico (45 million years old) comprise a diversity of habi-tats including coastal brackish-water swamps uplandtropical and subtropical forests and an arboreal cool-temperate forest (Graham amp Jarzen 1969) The majority(75) of the identified microflora consisted of generathat presently grow in Puerto Rico or the Antilles sug-gesting that the climate in the region was not dramatical-ly different from that of today and that evolution of someof these taxa might have occurred ldquoin siturdquo Howeverseveral northern cool-temperate genera that are notextant in Puerto Rico were found Engelhardia FagusLiquidambar and Nyssa Based on similarity of the fos-sils and the present tropical vegetation and assumingthat the Oligocene climate was comparable to that oftoday it can be inferred that a greater physiographicrelief must have existed in Puerto Rico to provide suit-able conditions for the temperate genera and the tropicalcommunities at the same time (Graham amp Jarzen 1969)In addition the authors interpret pathways of migrationin the region They found that all 14 genera from the SanSebastiaacuten flora that are not in the Antilles today arepresently distributed throughout Mexico CentralAmerica and South America an indication thatalthough San Sebastiaacuten contains elements present todayin the temperate Eastern United States it has strongerfloristic affinities with Mexico Central America andNorthern South America

Work on the Late Miocene-Middle Pliocene

Artibonite pollen from Haiti (Graham 1990a b) is thefirst Tertiary paleopalynological study of HispaniolaThe most common microfossils at Artibonite are fernspores of at least three families but also included Pinusand palms among others Two genera represented in thedeposits Alfaroa and Oreomunnea do not occur today inHaiti but are elements of the cloud forests of Mexico andCentral America Graham (1990a) estimates that highestelevations during the Late Miocene-Middle Pliocene inArtibonite could have been about 1400 m (the highestpoint today Morne de la Selle is 2900 m) Furthermorepollen of Asteraceae Chenopodiaceae and Amarantha-ceae suggest local openings in the vegetation but there isno evidence of dry forests The latter as well as savan-nas are thought to have developed later in the Plioceneand during the Quaternary (Leyden 1984)

Insights from Tertiary Paleocommunities inCentral America and Southern Mexico mdash TheGreater Antilles is part of a paleophysiographic provincethat also includes southern Mexico and Central America(Graham 1993) Consequently and given the lack of fos-sil sites in the Antilles our interpretation of the vegeta-tional history of the islands during most of the Tertiarycould be facilitated by analyzing available evidence fromthe Gulf-Caribbean deposits of Mexico and CentralAmerica The vegetational history of these deposits iscongruent with other independent studies of paleotem-perature patterns and paleofauna (Graham 1988)

In general regional cooling since the Miocene alongwith Quaternary climate fluctuations have been the mainfactors shaping the plant communities in southernMexico which has always had a continuous land con-nection to the North Prior to the Miocene temperaturedrop warm tropical conditions predominated in southernMexico as suggested by the elements present in thedeposits (Graham 1999a) On the other hand regionalclimatic cooling has apparently played a lesser role indefining the principal plant communities of CentralAmerica (Graham 1996) The vegetation of proto-Central America remained predominantly tropicalthroughout the Eocene-Miocene until the Pliocene andvery similar to that of todayrsquos low-lying regions(Graham 1989b 1990b 1996) None of the earlierCentral American paleofloras suggest elevations thatsupport high altitude and paacuteramo vegetation and thusthere was not a continuous habitat that would haveallowed interchange of temperate elements betweenNorth and South America (Graham 1990b 2003Burnham amp Graham 1999) Also the surrounding oceanwaters of the low-relief insular proto-Central Americacould have buffered the effects of global cooling(Graham 1989a) The increased diversity of the habitatsand elevations of present-day Central America resultedprimarily from recent volcanism and uplift associated


304

with plate movements and Quaternary climate fluctua-tions (Graham 1996 2003 Burnham amp Graham 1999)One of the most remarkable events in the CentralAmerican Pliocene was the connection of North andSouth America about 3 to 35 million years ago(Graham 1996) It is believed that dry forest originatedas the result of this land connection and the recentlyformed higher mountains ranges factors that deflectedthe northeast winds and produced marked differences inthe rainfall regime seen today between the wetterAtlantic slope and the dryer Pacific slope (Graham1989c 1990b Graham amp Dilcher 1995) Following for-mation of the isthmus of Panama plant migration pro-ceeded in both directions affecting the floras of northernSouth America and Central America Fossil studies(reviewed in Burnham amp Graham 1999 Graham 1999b2003) indicate that North American elements primarilyinvaded high elevation temperate zones in SouthAmerica whereas as much as 75 of the rainforestcanopy trees in southern Mexico may be of SouthAmerican origin (Wendt 1993)

PHYTOGEOGRAPHIC PATTERNSSUGGESTED BY FLORISTIC DATA

Early notions of floristic affinities of Caribbeanplants can be attributed to the authors of the first detailedfloras for the region whose comments revolved primari-ly around the island each one was studying (eg Urban1923 Eggers 1879 Alain 1958) Initial observationsled to distinguish distributions in the Caribbean flora asbeing primarily Greater Antillean or Lesser AntilleanThe Greater Antillean flora was considered to be olderand to have North South and especially CentralAmerican affinities (Asprey amp Robbins 1953 Alain1958) Within the Greater Antilles affinities betweenHispaniola and Cuba were considered stronger thanthose of Hispaniola with Jamaica (Woodson 1940)Affinities were also observed between Puerto Rico andHispaniola Most of the relationships in the region wereexplained by land bridges between Honduras and theGreater Antilles (Schuchert 1935 Asprey amp Robbins1953) and by Cuban land connections with Yucatan andFlorida (Alain 1958) Dispersal by sea and wind cur-rents and by birds and bats was suggested to explainmore recent migrations (Alain 1958) or those for whichland bridges did not fit interpretation of current distribu-tions (Beard 1949 Asprey amp Robbins 1953) Theunique distributions and the unexplained absence of cer-tain taxa in Jamaica led to the development of a com-bined argument involving the existence of an old landbridge with Central America and more recent dispersalevents after the re-emergence of the island (Asprey amp

Robbins 1953) In contrast dispersal and not landbridges was the undisputed interpretation for the originof the vegetation of the Lesser Antilles The advent ofplate tectonics (Hess 1962 Tuzo-Wilson 1965 Morgan1968 Cox 1973 Bird amp Isacks 1980) was a major rev-olution that lead to the rejection of land bridges(Williams 1989) and to the establishment of a differentvicariant scenario

In a study describing the vegetation of the AntillesHoward (1973) provides a summary of generic affinitiesbetween that region and continental America by tabulat-ing distribution data in several combinations of islandsand continental regions Based on the total number ofgenera sharing a similar range he defined five main pat-terns or ldquounitsrdquo The first the Pan-Caribbean unit arethose genera occurring across the Greater and the LesserAntilles Mexico Central America and Northern SouthAmerica The second the Western Continental unitcomprises genera in Central America Northern SouthAmerica and the Greater Antilles That is genera thatare absent from Lesser Antilles This second unit is fur-ther subdivided by whether the genera are exclusive tothe Greater Antilles and Central America or to theGreater Antilles and South America The second unit alsoseparates genera whose distribution exclude Jamaica orPuerto Rico The third or Southern Continental unitincludes genera distributed in Central America NorthernSouth America and the Lesser Antilles but missing fromthe Greater Antilles The fourth or Antillean unit groupsgenera restricted to or better developed in the GreaterAntilles The fifth or Greater Antillean unit is formed ofgenera endemic to one or more islands in the GreaterAntilles which could be considered a subset of unit fourthe Antillean unit Geographic affinities in this work aredescriptive in nature and although some of the units arereminiscent of Leoacuten Croizatrsquos idea of mapping inde-pendent ranges that coincide to define ldquotracksrdquo (Croizat1952 1958) no interpretation of such units is given(Howard 1973) Furthermore the work also addressesthe presence of long distance disjuncts in the Antillesbased on the works by Good (1964) but besides statingthat this region shares genera with Africa Madagascarand Asia there is no inference to explain how these dis-juncts arose

Phytogeographical patterns of the Caribbean wererevisited as apart of a larger scale assessment ofNeotropical floristic diversity using distributional dataon available regional floras published monographsherbarium specimens and fossils (Gentry 1982) Gentryalso considered the following geological and climaticforces as relevant to his interpretations the emergence ofthe Proto-Greater Antilles between North and SouthAmerica (Late Cretaceous-Early Tertiary) the morerecent Central American isthmian connection (Pliocene)


305

the uplift of the Northern Andes (Pliocene-Pleistocene)and the Quaternary climatic fluctuations (Pleistocene)(Gentry 1982)

The Neotropical flora presents two majordichotomies The first distinguishes Laurasian-derivedfrom Gondwanan-derived taxa (Gentry 1982) In mostof the Neotropics Laurasian-derived taxa if present areprimarily montane higher altitude plant groups of herbsand sometimes canopy trees mixed within theGondwanan elements Some examples in the Caribbeaninclude Magnoliaceae Pinaceae AquifoliaceaeCyrillaceae Rosaceae and Ranunculaceae They are notvery species-rich a fact interpreted by Gentry as evi-dence of their recent (Late Tertiary-Quaternary) arrivalOn the other hand Gentry points out that in theCaribbean islands some old Laurasian-derived groupshave thrived and diversified well in lowland dry areasincluding members of Aristolochiaceae VitaceaeRhamnaceae Boraginaceae and Buxaceae In generaltaxa of Laurasian derivation are better represented in theCaribbean than in other parts of the Neotropics and thusthe Laurasian-Gondwanan dichotomy is more pro-nounced in the islands than in other Neotropical regions(Gentry 1982) Some of the northern taxa of the dry low-lands may be considered reminiscent of the endemictropical Laurasian families and could have been some ofthe stocks that during the Late Cretaceous-Early Tertiaryreached the Proto-Greater Antilles an idea inferred byother authors (eg Graham 1990b Lavin 1993)

Gentryrsquos second dichotomy is the subdivision of theGondwanan group into Neotropical Amazonian-centeredand extra-Amazonian (ldquoAndean-Centeredrdquo) subgroupsThe Amazonian-centered subgroups include most of theNeotropical lowland canopy trees and lianas and gener-ally have few species per genus On the other hand taxapredominantly epiphytic or shrubby have their distribu-tion centers in low and mid-elevations of the AndesMany exhibit explosive radiation including Bromelia-ceae Gesneriaceae Piperaceae Orchidaceae Melastom-ataceae Acanthaceae Myrsinaceae Rubiaceae andSolanaceae (Gentry 1982) Both Gondwanan groups arecommon forest components of the Caribbean islands

STUDIES APPLYING PHYLOGENET-ICS TO BIOGEOGRAPHICAL IN-TERPRETATIONS OF CARIBBEANPLANTS

In recent years a growing number of plant systema-tists and biogeographers have applied phylogeneticinformation to test hypotheses of evolutionary relation-ship and historical biogeography Analytical approachesto biogeography are varied depending upon the ques-

tions involved (Page amp Lydeard 1994 Morrone ampCrisci 1995) We believe however that phylogeneticanalysis provides the most objective approach to assessthe historical biogeography of a given taxon A phyloge-ny provides a direction and a sequence to the reconstruc-tion of geographic distributions whether dispersal vic-ariance or a mix is inferred

Few studies on Caribbean plant groups have incor-porated phylogenetic reconstructions Consequentlyeven answers to basic questions on evolutionary rela-tionships systematic placement morphological or eco-logical evolution are lacking for most taxa in the regionHere we present outlines of studies on Caribbean plantsin which a phylogenetic approach most using moleculardata has been incorporated into biogeographic assess-ments Various methods of tree construction and biogeo-graphic inference were used It is not our intent to reana-lyze these data but rather to look for common patterns

Lyonia (Ericaceae) mdash One of the pioneeringstudies applying phylogenetics to the biogeography ofCaribbean plants was that for the genus Lyonia sectionLyonia (Judd 1981 1995 2001) The genus with mem-bers in Eastern Asia and Eastern North America com-prises a monophyletic group (Judd 2001) of approxi-mately 25 species in the Greater Antilles Many of theGreater Antillean Lyonia are narrow endemics that occurin a variety of environments including low-elevationthickets savannas and moist montane and cloud forestsWithin-island isolation is in many cases due to edaphicconditions as well as by altitudinal differences (Judd2001) The major centers of diversity of Lyonia in theAntilles are the Cordillera Central and the Massiff de laSelle-Sierra de Bahoruco both in Hispaniola and themountains of Eastern Cuba (Judd 1981)

Phylogenetic reconstructions using morphologicalcharacters were analyzed by with a Wagner-Groundplan-Divergence Analysis (Judd 1981) and later reassessedusing heuristic and branch-and-bound algorithms (Judd1995 2001) A strict consensus tree based on this latterreassessment (Fig 2) illustrates that all Antillean mem-bers of Lyonia sect Lyonia (except the Jamaican Ljamaicensis see below) belong in two major clades TheldquoHispaniolan claderdquo is formed by taxa ocurring inHispaniola Puerto Rico and St Thomas (VirginIslands) The second clade or ldquoCuban claderdquo includes allthe Cuban taxa as well as Lyonia octandra fromJamaica Within the Cuban clade all but one of the taxafrom Cuba group together In this phylogenetic recon-struction the lineage of the Jamaican taxon L jamaicen-sis comes out unresolved and outside of the Hispaniolanand Cuban clade

An area cladogram resulting from a BrooksParsimony Analysis for the Antillean Lyonia (Judd2001) was in agreement with geographic relationships


306

suggested by the species phylogeny This was not sur-prising because species of Lyonia growing on eachisland (especially in the case of Cuba and Hispaniola)came out in the phylogeny as closely related to speciesfrom the same island In the area cladogram taxa fromthe Cordillera Central and Massif de la Selle(Hispaniola) formed a clade with a taxon distributed inPuerto Rico and the Virgin Islands and taxa of the north-ern Oriente region (Cuba) form a clade with other Cubantaxa and one Jamaican species The derived taxa in bothislands exhibit adaptations to extreme habitats (Judd2001)

Results of this study were also evaluated in light ofRosenrsquos (1976 1985) vicariant model to explain the ori-gin of the biota of the Greater Antilles and the geologichistory of the Antilles (Judd 2001) A close relationshipis seen between Eastern Cuba North-Central Hispaniolaand Puerto Rico as well as between Western Cuba andSouthwestern Hispaniola Results of this study are inagreement with the close relationship between EasternCuba North-Central Hispaniola and Puerto Rico Thusthis suggests the possibility of a vicariant scenario insome lineages of Lyonia However Western Cuba comesout as closely related to Eastern Cuba and Southwestern

Hispaniola as closer to Central Hispaniola An interpre-tation of this disagreement with Rosenrsquos vicariant modelis that Lyonia was originally restricted to Eastern Cubaand North-Central Hispaniola and that these areas pro-vided the source of more recent lineages in Western Cubaand Southwestern Hispaniola respectively (Judd 2001)This could have occurred as the result of within-islanddispersal events One argument to provide support for theantiquity of Eastern Cuba and Southwestern Hispaniolais that Lyonia is most diverse in those areas TheJamaican species are also considered to be the result ofdispersal That most species of Lyonia occur in limitedranges and species in the same island are closely relatedindicates that successful long-distance dispersal (at leastbetween islands) is not very common The eastward (ieCuba to Virgin Islands) decrease in Lyonia diversity (theldquoWestern continental distribution patternrdquo described byHoward in 1973) and the absence of the genus in theLesser Antilles suggests that taxa arrived in the islandsfrom MexicoEastern North America (Judd 2001)

Sabal (Arecaceae Coriphoideae) mdash A mono-graph of Sabal includes a phylogenetic analysis based onmorphology leaf anatomy and flavonoid phytochem-istry (Zona 1990) This work recognizes 15 species dis-


Fig 2 Strict consensus tree of selected species of Lyonia sect Lyonia based on morphological characters (adaptedfrom Judd 1995 2001)

307

tributed throughout the southern United States (fourspecies three endemic) Mexico and northern CentralAmerica (six species five endemic) Cuba (three speciesnone endemic) Jamaica (one species none endemic)Hispaniola (two species one endemic) Puerto Rico (onespecies none endemic) and Bermuda (one speciesendemic) One species (Sabal mauritiiformis) shows anotably patchy distribution occurring in GuatemalaBelize southeastern Costa Rica eastern Panama andscattered along northern South America

The cladistic analysis of Zona (1990) identified fivemajor lineages (Fig 3) Using the Mexican palmsWashingtonia filifera and Brahea dulcis as outgroupsthe resulting tree shows the most basal clades withspecies endemic to (or mainly with) a US distributionThe western Mexican species form a clade that is sisterto the southern Mexican species and the Antillean-endemic species which in turn form clades that are sis-ter to each other One species each from the SE US andMexican clades also occurs in Cuba The presence of tan-nins within the bundle sheath surrounding the vascular

bundles is a synapomorphic trait for the Antillean-endemic clade (Zona 1990)

Based on the patterns of phylogenetic relationshipsand distributional data Zona hypothesized origin ofSabal in North America with the Antillean-endemicspecies being derived from Mexican lineages rather thanfrom those in the Southern United States The presenceof a species of Sabal in Bermuda and the apparent originsof Greater Antillean taxa from three areas outside theAntilles together with evidence of zoochory (Zona ampHenderson 1989) and hydrochory (in at least somespecies in the genus) points towards over-water disper-sal as a plausible mechanism to explain the distributionsof this palm genus in the Caribbean (Zona 1990) Thephylogeny of Asmussen amp al (2000) indicates that aclade of five genera (Chelyocarpus CoccothrinaxCrysophila Thrinax and Trithrinax) are sister to Sabalwith Washingtonia and Brahea more distantly relatedThe inferred ancestry of this sister clade is SouthAmerican It is uncertain whether this would alter theinference of a North American origin for extant Sabal


Fig 3 Phylogeny of Sabal and related genera (adapted from Zona 1990 and Asmussen amp al 2000)

Coccothrinax

Thrinax

Chelyocarpus

Crysophila

Trithrinax

S minor

S miamiensis

S etonia

S palmetto

S bermudana

S uresana

S rosei

S pumos

S yapa

S mauritiiformis

S guatemalensis

S mexicana

S maritima

S domingensis

S causiarum

Antilles

S America

C America

S America

S amp SE United States

Florida

SE US Bahamas W amp C Cuba

Bermuda

W Mexico

Mexico

S Mexico Guatemala

S Mexico to northern S America

S Mexico W Cuba

Cuba Jamaica

N Hispaniola

SW amp E Hispaniola Puerto Rico

308

species but it is unlikely to alter the inference of aMexican origin for Antillean species

Mecranium (Melastomataceae) mdash A mono-graphic study on the Antillean-endemic genusMecranium included a very detailed assessment of mor-phology as well as anatomical phenological and cyto-logical data (Skean 1993) Mecranium is most species-rich in Hispaniola with 16 species 14 of which areendemic The southwestern portion of Hispaniola (theldquosouth islandrdquo of Skean) contains most of the diversity inthis genus Of the 13 species occurring there 10 areendemic to that portion of the island and seven of thoseare unique to the Massif de la Hotte of Haiti (Skean1993) Northern Hispaniola contains only five speciestwo of which are endemic to the region The otherspecies in the genus are in Cuba (five species threeendemic) Jamaica (three endemic species) and PuertoRico (one endemic species)

Using cladistic analysis Skean divided the genusinto two sections sect Sagreoides with two Hispaniolanendemic species and sect Mecranium with the remain-ing species (Fig 4) The presence of a calyptra is asynapomorphy for the large section Mecranium The lat-ter clade presents a basal trichotomy One of the lineag-es consists of a single Cuban species A second lineageconsists of a trichotomy that includes two southwesternHispaniola and one Jamaican species The third cladecontains Cuban and Hispaniolan species plus a polyto-my with taxa from the four islands (the Mecranium mul-tiflorum complex Skean 1993) Despite these threepolytomies the phylogeny demonstrates that none of thespecies occupying islands form monophyletic groupsexcept the trivial case of the sole species in Puerto Rico

Skean proposes that the current distribution ofMecranium has resulted from tectonic events dispersalby birds and climatic changes The striking level ofendemism in southwestern Hispaniola suggests that iso-lation there must have been of much greater extent thanthe 20 km-wide Cul de Sac-Enriquillo plain that todayseparates this region from Northern Hispaniola This isconsistent with the geological hypothesis of the inde-pendent insular nature of southwestern HispaniolaBecause Hispaniola is the center of diversity forMecranium (Skean 1993) the author suggested that theancestral stock that gave raise to the genus might havefirst occurred and radiated in insular southwesternHispaniola followed by dispersal to northern portions ofHispaniola Puerto Rico Cuba and finally Jamaica Thishypothesis is consistent with the phylogeny but thesequential order of some of these island colonizationscannot be assessed with the present cladogram due tolack of resolution especially in the M multiflorum com-plex which comprises 11 species representing all fourislands (Skean 1993) The fact that most members of

Mecranium grow in higher elevations mainly above 800m (Skean 1993) and that many species have veryrestricted rangesmdashin some cases confined to a singlemountain rangemdashled Skean to infer that Pleistocenecooling might have played a role in shaping the currentdistribution of the genus The sister group to Mecraniumis not clear (W Judd pers comm) thus no continentalorigin of the ancestral stock that gave rise to Mecraniumcan be inferred

Poitea (Fabaceae) mdash Lavin (1993) assessed thebiogeography and systematics of the Antillean-endemicgenus Poitea a member of the North-American-centeredtribe Robinieae using phylogenetic reconstruction basedon morphological and chloroplast DNA restriction sitedata This was expanded to the entire Robinieae usingcpDNA and ITS sequences (Lavin amp al 2003) Poiteawith 12 species is confined to Cuba (two endemicspecies) Hispaniola (seven species six endemic) PuertoRico (three species two endemic) and the LesserAntillean island of Dominica (one endemic species) Thegenus does not occur in Jamaica Current circumscriptionof the genus includes the formerly recognized generaSabinea Sauvallella Corynella Notodon andBembicidium (Polhill amp Souza 1981 Lavin 1993)Hebestigma with two species endemic to Cuba was alsoincluded in the study of Lavin amp al (2003)

The cladistic analysis reflects two main lineages orsubgroups within Poitea (1) the Poitea galegoidesalliance comprised of species with reddish tubular corol-las and imparipinnate leaves and (2) the Poitea floridaalliance with papilionaceous flowers and paripinnateleaves (Lavin 1993 Fig 5) The Poitea galegoidesalliance is distributed in Western Cuba and southwesternto north-central Hispaniola whereas the Poitea floridaalliance is found in eastern Cuba north-centralHispaniola the Puerto Rican bank and Dominica Thebiogeographic assessment of Poitea by Lavin (1993) isbased on the ideas of geohistory of the Greater Antillesoutlined by Rosen (1976 1985) Pindell amp Barrett(1990) and Perfit amp Williams (1989) Lavin initiallysuggested that if a vicariant model is applied to interpretthe current distribution of Poitea the group must haveoriginated in the Early Tertiary (Lavin 1993) Close rela-tionships between members of the Poitea galegoidesalliance from western Cuba and Hispaniola (primarily inthe southwestern peninsula of Hispaniola) is consistentwith diversification during the Early Eocene when west-ern Cuba and southwestern Hispaniola were closer toeach other and to northern Central America Later in theTertiary during the accretion of southwestern Hispaniolato the rest of the island members of this alliance expand-ed its range to central Hispaniola For members of thePoitea florida alliance Lavin found that the EasternCuban and the north-central Hispaniolan species are sis-


309

ter to each other and this clade is sister to three speciescentered in Puerto Rico This pattern of relationship isconsistent with the hypothesis of a close land connectionbetween eastern Cuba Central Hispaniola and PuertoRico during the Early Tertiary (Rosen 1985 Perfit ampWilliams 1989 Pindell amp Barrett 1990) Howeverbased on the molecular clock in Lavin amp al (2003) thedates for this diversification appear to be much morerecent than the vicariant model would predict They esti-mated minimum divergence times of 16 mya for Poiteaand its sister Gliricidia and ca 93 mya for extant Poiteaspecies The vicariant model therefore seems highlyunlikely

Gliricidia the sister to Poitea is distributed inMexico and northern South America Sister to thePoiteaGliricidia clade is a group of nine genera com-prising the remainder of tribe Robinieae The first branchin this clade comprises Lennea (Central America) andHebestigma the next branch is Olneya (SW NorthAmerica) The remaining six genera are widely distrib-uted in North Central and South America A simple par-

simony optimization of ancestral distributions suggests aprobable origin for Robinieae in Central America orMexico This further suggests two colonizations of theAntilles (Hebestigma and Poitea) most likely in Cubawith subsequent eastward colonization in Poitea (but notHebestigma) including two lineages reaching Hispaniolaand diversifying there one of these spawning twomigrants to Puerto Rico and another dispersing toDominica

Bactris (Arecaceae) mdash Salzman amp Judd (1995)investigated the ingroup relationships of the Antilleanspecies of Bactris which were previously found to be amonophyletic group embedded in a larger clade contain-ing northern South American species (Sanders 1991)Bactris one of the largest Neotropical palm genera (over60 species according to Henderson amp al 1995) has onlythree species endemic to the Greater Antilles one each ineastern Cuba Hispaniola and Jamaica

The cladistic analysis of morphological and anatom-ical characters by Salzman and Judd confirmed mono-phyly of the Antillean Bactris placing the Hispaniolan Bplumeriana as sister to the Jaimaican B jamaicensis andboth as sister to the Cuban species B cubensis (Fig 6)Salzman and Judd invoke both vicariance and dispersalexplanations to reconstruct the historical development ofthe Antillean clade basing their ideas on the obtainedphylogeny current distributions of species within theislands and the present models of geological evolution inthe Caribbean In their interpretation the stock to whichBactris belongs occurred in western Gondwana (Uhl ampDransfield 1987) and could have reached the Proto-Greater Antilles during late Cretaceous-Early TertiaryUnder this scenario the eastward drift of the Proto-Greater Antilles represented the vicariant event responsi-ble for the split between the Antillean and the northernSouth American lineages (Salzman amp Judd 1995) Theoriginal Antillean stock could have been on the EasternCuba-Central Hispaniolan block which eventually splitSubsequently the Hispaniolan populations could havereached Southwestern Hispaniola and then the youngerJamaica via water dispersal

Exostema (Rubiaceae) mdash Phylogenetic recon-struction and biogeographic interpretations on the basisof ITS and rbcL sequence data (Fig 7) were carried outfor Exostema a neotropical genus of 25 species of treesand shrubs with a high level of endemicity in the GreaterAntilles (McDowell amp Bremer 1998 McDowell amp al2003) The initial biogeographic conclusions based onmorphology and ITS sequences but with limited out-group sampling (McDowell amp Bremer 1998) were over-turned with the addition of rbcL sequences and moreextensive outgroup sampling (McDowell amp al 2003)Exostema comprises four predominantly Antillean clades(one clade includes a derived Central American species)


310

Fig 4 Phylogeny of Mecranium (adapted from Skean1993)

M acuminatum

M amygdalinum

N Hispaniola

M crassinerve

M haitiense

M purpurascens

M racemosum

M multiflorumcomplex (11 spp)

M birimosum

M microdyctium

M puberulum

M haemanthum

M tuberculatum

N amp SW Hispaniola

SW Hispaniola

Jamaica

W Cuba

SW Hispaniola (5 spp)N Hispaniola (1 sp)N Hispaniola C amp E Cuba (1 sp)Cuba (1 sp)W Cuba (1 sp)

Jamaica (2 spp)Puerto Rico (1 sp)

SW Hispaniola

N Hispaniola

W amp E Cuba

SW Hispaniola amp E Cuba

sect Sagreoides

sect M

ecra

nium

and one South American clade which also includes thegenus Coutarea None of the Antillean clades representtaxa of a single island each includes at least one speciesendemic to Cuba and to Hispaniola Nested withinExostema is a clade comprised of Chiococca (AntillesFlorida and tropical America) and Erithalis (AntillesBahamas and Florida see below) The three sequentialoutgroups to the clade of Exostema (Cubanola Isidoreaand Catesbaea in order of increasing distance) and itsthree derivative genera are all Antillean in distributionThus this entire clade diversified in the Caribbean basinand has at least three descendant lineages in Central andSouth America (Fig 7)

Erithalis (Rubiaceae) mdash A study on the phylo-genetics systematics and biogeography of the genusErithalis (Rubiaceae) was carried out by Negroacuten-Ortiz ampWatson (2002 2003) using chloroplast DNA sequences(trnF-trnL) as well as nuclear sequences (ITS and ETS)Erithalis is a small group of about eight to ten species oftrees and shrubs distributed in the West Indies MexicoHonduras Venezuela and Brazil Some species have aPan-Caribbean distribution whereas others are narrow

endemics In this analysis taxa with broad distributionalranges were sampled from different geographic areas

Parsimony analysis of the combined dataset supportsmonophyly of Erithalis The obtained phylogeny con-sists of two major clades (Fig 8) One clade groups sev-eral lineages of E fruticosa (from Cuba Jamaica PuertoRico and Florida) as sister to the Jamaican E harisiiThe other clade groups five other species with theJamaican E quadrangularis and one lineage of E odor-ifera (from the Bahamas) coming out as early lineagesplits the rest of the lineages form two subclades One ofthe subclades forms a polytomy consisting of lineagesfrom the Bahamas Cuba and Hispaniola Accessionsidentified as E fruticosa from Florida and the Bahamaswere also part of this clade but were suspected to be ofhybrid origin and not included in subsequent studies(Negron-Ortiz amp Watson 2002 2003) The other sub-clade comprises several lineages of E odorifera fromPuerto Rico and the Lesser Antilles (Dominica and StVincent) Erithalis is sister to Chiococca (AntillesCentral and South America) and together are nestedwithin the paraphyletic genus Exostema (see above) thus


311

Fig 5 Phylogeny of Poitea and related genera of Robinieae (adapted from Lavin amp al 2003)

P emarginata

P multiflora

P glyciphylla

P galegoides

P campanilla

P punicea

P florida

P carinalis

P paucifolia

P dubia

P gracilis

Gliricidia

Lennea

Hebestigma

Olneya

Poissonia

Robinia

Coursetia

Genistidium

Sphinctospermum

Peteria

W Cuba

SW C Hispaniola

SW Hispaniola

Hispaniola

C Hispaniola

Puerto Rico

Dominica

N C Hispaniola Puerto Rico

C Hispaniola

Cuba

5 spp Mexico C amp S America

3 spp C America

1 sp Cuba

1 sp SW N America

4 spp S America

4 spp N America


1 sp Mexico

1 sp SW N America

4 spp SW N America

suggesting an in situ origin of Erithalis in the AntillesIn this study branch lengths are short and most

clades have weak bootstrap support Extremely lowsequence divergence indicates recent diversificationTaken together with the widespread occurrence of sever-al species (E fruticosa E odorifera E salmeoides) sug-gests dispersal as the primary historical factor in presentdistributions

Goetzeoideae (Solanaceae) mdash A study bySantiago amp Olmstead (2003) investigated the evolution-ary relationships of the Antillean genera GoetzeaEspadaea Henoonia Coeloneurum and the SouthAmerican genera Duckeodendron and MetternichiaGoetzea consists of two species one each in Hispaniolaand Puerto Rico Coeloneurum is a monotypic genus ofHispaniola Espadaea and Henoonia are each monotypicgenera endemic to Cuba No member of this plant groupis present in Jamaica Duckeodendron and Metternichiaare monotypic genera endemic to Brazil The systematicplacement of all of these taxa except Metternichia havebeen a long-standing question The Antillean generawere suggested (Hunziker 1979) to form the only plantfamily (Goetzeaceae) endemic to the Greater Antillesand Duckeodendron was believed to represent the onlyplant family endemic to the Amazon basin (Kuhlmann1930 1947)

Phylogenetic reconstruction using nuclear andchloroplast DNA sequence data (Fay amp al 1998Santiago amp Olmstead 2003) strongly supported mono-phyly of these genera and placed them in Solanaceae(Fig 9) Within the clade the Antillean genera are mostclosely related to each other and the Brazilian genera arethe basal lineages These taxa represent an early diverg-ing lineage within Solanaceae (Fay amp al 1998) provid-ing evidence for the establishment of a new subfamily inSolanaceae Goetzeoideae (Olmstead amp al 1999) Basedon the phylogeny and current geological knowledge aplausible explanation of the history of the group in the

Antilles involves colonization from South America toHispaniola or Cuba The Puerto Rican lineage then arosefrom Hispaniola Phylogenetic reconstruction inGoetzeoideae indicates evolution towards xeric environ-ments (in Coeloneurum and Henoonia) and a shift in thepollination syndrome from moth pollination (in theSouth American Metternichia and Duckeodendron) tobird pollination (in the Antillean taxa)

Additional studies mdash Several additional studieshave addressed phylogeny and biogeography ofAntillean plant groups Pictetia (Fabaceae) is a genus ofeight species with one endemic to Puerto Rico oneendemic to Hispaniola one occurring on both Cuba andHispaniola and the other five endemic to Cuba A phy-logeny based on ITS and morphological data found thegenus to be monophyletic and sister to a clade occurringpredominantly in Africa and Madagascar and the inclu-sive clade is sister to Diphysa which is restricted toCentral America (Beyra M amp Lavin 1999) This distri-bution of related groups and the relatively high sequencedivergence among species in Pictetia suggests a longhistory in the Antilles with probable origin from what isnow Central America Present distributions are consis-tent with an early diversification on a land mass consist-ing of the present-day Puerto Rico north-centralHispaniola and eastern Cuba with a basal polytomy inthe phylogeny between the three islands Subsequentradiation has been restricted to Cuba and an apparent dis-persal event in P sulcata between Cuba and Hispaniola

Ginoria including Haitia (Lythraceae) is a clade of16 species 14 of which are restricted to either Hispaniolaor Cuba one in Puerto Rico and the Virgin Islands andone in Mexico A phylogenetic analysis based on mor-phology found that the Hispaniolan plus Cuban speciesform a clade with the Mexican and east Caribbeanspecies unresolved at the base (Graham 2002 2003)Neither of the groups of species occurring on Cuba orHispaniola form monophyletic groups Sister to Ginoria


312

Fig 6 Phylogeny of Bactris (adapted from Salzman amp Judd 1995)

is a small clade of two species inhabiting coastal envi-ronments in northern South America and the island ofMauritius in the Indian Ocean Outgroups to this inclu-sive clade are Afro-Asian in distribution thus the geo-graphic ancestry of this group is uncertain

Four species of Styrax sect Valvatae (Styracaceae)occur in the Antilles one endemic to Puerto Rico oneendemic to Hispaniola one on both Cuba andHispaniola and one in the Lesser Antilles and northernSouth America A phylogenetic study of sect Valvataefound that three separate origins in the Antilles arerequired to explain present distributions with two origi-nating in South America and one from Mexico (Fritsch2003) Another case study similar to Styrax in whichthere were multiple introductions to the Antilles

involves Cuphea (Lythraceae) A phylogenetic studybased on morphology and ITS sequence data found atleast five and more likely as many as eight (includingone human introduction) separate origins of Antilleanlineages (Graham 2003) Colonization from easternSouth American ancestors occurred at least three timesand from Central American ancestors at least twice(including the recent introduction) the remaining intro-ductions are of uncertain origin because the clades towhich they belong contain species in each area and sam-pling is not complete There has been limited subsequentspeciation in the Antilles with only seven species beingendemic

Major biogeographic patterns from the previousphylogenies include the following


313

Fig 7 Phylogeny of Exostema (adapted from McDowell amp Bremer 1998 McDowell amp al 2003)

E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca

Erithalis vacciniifolia

Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola

Cuba Hispaniola Puerto Rico

S America

7 spp Mexico to S America

Peru

Hispaniola

Cuba Hispaniola

Antilles Florida Mexico C America

6 spp Florida tropical America

10 spp Antilles Bahamas Florida

Hispaniola

Cuba

2 spp Cuba

20 spp Antilles

6 spp N America to tropical America

1 sp Antilles

Catesbaea 20 spp Antilles Florida

E lancifolium

E purpureum Cuba

Cuba

E brachycarpum Jamaica

E stenophyllum Cuba

1 North American and South American ori-gins of Caribbean plant groups mdash Phylogenies forLyonia Poitea Hebestigma Pictetia and Sabal indicatecloser relationship to stocks from North America main-ly Mexican lineages This contrasts with Bactris andGoetzeoideae which are derived from South AmericaStyrax and Cuphea each have one or more lineages in theAntilles derived from each continent

2 Groups of in situ origin mdash Exostema andErithalis both belong to the same lineage of Rubiaceaethat apparently occupied the Antilles before the origin ofeither genus Each has members (Exostema) or a closelyrelated group (Erithalis) with continental distributionsleading to erroneous earlier hypotheses

3 Groups of unknown origin mdash Mecraniumand Ginoria (including Haitia) both have geographicallydistant andor diverse continental relatives and thus areof unknown continental origin

4 Plant groups recolonizing continentalland masses mdash This is illustrated best in Exostemawhere three colonizations of Central and South America

are inferred Erithalis exemplifies another pattern withtwo lineages that have colonized Florida from Antilleanancestors

5 In Cuba and Hispaniola species within anisland do not form monophyletic groups mdashSpecies on one island often have their closest relatives onanother island (Exostema Goetzeoideae Mecraniumand Poitea) The presence of a diverse assemblage ofunrelated lineages has been suggested as evidence for thecomposite nature of these islands (Lavin 1993) but italso could indicate relatively frequent inter-island colo-nization events A contrasting pattern is found in Lyoniawhere all the Cuban species are in the same lineage andanother lineage comprises all the Hispaniolan species(plus Puerto Rico and the Virgin Islands see below)Pictetia is similar with a clade of Cuban species one ofwhich also occurs on Hispaniola In Ginoria (includingHaitia) the primary radiation was on Cuba with two orthree colonizations of Hispaniola Evaluation of diasporedispersal will be of importance to understand these dif-ferences (Judd 2001)


314

Fig 8 Phylogeny of Erithalis (adapted from Negroacuten-Ortiz amp Watson 2003)

Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii

Exostema longiflorum

Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera

6 Relationship of Jamaica taxa with Cubaand Hispaniola mdash Lyonia octandra (Jamaica) isclosely related to the Cuban lineages of the genusRelationships of Jamaican Lyonia jamaicensis is stillunclear A lineage in Sabal is shared between Jamaicaand Cuba as is an unresolved lineage of four species ofExostema with two species each on Cuba and HispaniolaAffinities between Jamaica and southwestern Hispaniolaare found in a clade in Mecranium A similar relationshiphas been suggested for Bactris (Salzman amp Judd 1995)but the phylogeny is equivocal

7 Relationship of Puerto Rican taxa withHispaniola mdash Phylogenies of four independent plantgroups support the close relationship of Puerto Rico andHispaniola Sabal Lyonia Poitea and Goetzea

8 Greater Antillean-centered groups withrepresentatives in the Virgin Islands or theLesser Antilles which are closely related to andmost likely derived from Puerto Rico mdash This isobserved in Erithalis (a relationship of Puerto Rico withSt Vincent and Dominica) Lyonia (Puerto Rico and StThomas) and Poitea (Puerto Rico and Dominica)

9 Relationship between eastern Cubanorth-central Hispaniola and Puerto Rico mdashLineages that to different extents suggest a relationship

between these land areas include Lyonia Poitea andGoetzeoideae These taxa are absent or not very well rep-resented in Jamaica Judd (2001) infers that members ofLyonia may be the result of vicariant events betweenCuba north-central Hispaniola and Puerto Rico but thepattern is obscured by subsequent dispersal events ofrecent lineages

10 Relationships between western Cubaand southwestern Hispaniola mdash This pattern isfound only in the Poitea galegoides alliance

ACKNOWLEDGEMENTSThanks to J Ammirati and C Hamilton for reading an early

draft of this manuscript Pedro Acevedo for providing estimates ofplant diversity in the Lesser Antilles and two anonymous review-ers for helpful comments on literature we missed Support wasprovided by the University of Washington Botany DepartmentUniversity of Colorado Department of EPOB University of PuertoRico and the American Society of Plant Taxonomists as well asNSF grant DEB-9509804 to RGO


315

Fig 9 Phylogeny of Goetzeoideae (from Santiago-Valentiacuten amp Olmstead 2003)

LITERATURE CITEDAcevedo-Rodriacuteguez P 1996 Flora of St John US Virgin

Islands New York Botanical Garden New YorkAdams C D 1972 Flowering Plants of Jamaica Robert

MacLehose amp Company Ltd GlasgowAdams R P 1989 Biogeography and evolution of the

junipers of the West Indies Pp 167ndash190 in Woods C A(ed) Biogeography of the West Indies Past Present andFuture Sandhill Crane Press Gainesville

Alain H 1958 La Flora de Cuba sus principales caracteriacutesti-cas y su origen probable Rev Soc Cubana Bot 15 36ndash5984ndash96

Alain H 1963 Flora de Cuba vol 5 Univ de Puerto RicoRio Piedras

Asmussen C B Baker W J amp Dransfield J 2000Phylogeny of the palm family (Arecaceae) based on rps16intron and trnL-trnF plastid DNA sequences Pp 525ndash535in Wilson K L amp Morrison D A (eds) MonocotsSystematics and Evolution CSIRO Melbourne

Asprey G F amp Robbins R G 1953 The vegetation ofJamaica Ecol Monogr 23 359ndash412

Beard J S 1949 The Natural Vegetation of the Windwardand Leeward Islands Oxford Univ Press Oxford

Berazaiacuten-Iturralde R 1976 Estudio preliminar de la floraserpentiniacutecola de Cuba Ciencias Ser Bot 12 11ndash26

Berazaiacuten-Iturralde R 1981 Reporte preliminar de plantasserpentiniacutecolas acumuladoras e hiperacumuladoras dealgunos elementos Rev Soc Cubana Bot 2 48ndash59

Bermingham E 1994 Historical biogeography of thebananaquit (Coereba flaveola) in the Caribbean region amitochondrial DNA assessment Evolution 48 1041ndash61

Beyra M A amp Lavin M 1999 Monograph of Pictetia(LeguminosaemdashPapilionoideae) and review of theAeschynomeneae Syst Bot Monogr 56 1ndash93

Bird J M amp Isacks B 1980 Plate Tectonics AmerGeophysical Union Washington DC

Bogart J P amp Hedges S B 1995 Rapid chromosome evo-lution in Jamaican frogs of the genus Eleutherodactylus(Leptodactylidae) J Zool 235 9ndash31

Breuil M amp Masson D 1991 Some remarks on LesserAntillean bat biogeography Compt Rend SommaireSeacuteances Soc Biogeacuteogr 67 25ndash39

Brooks R R 1987 Serpentine and its Vegetation DioscoridesPress Portland

Burnham R J amp Graham A 1999 The history of neotrop-ical vegetation new developments and status AnnMissouri Bot Gard 86 546ndash589

Cedentildeo-Maldonado J amp Breckon G J 1996 Serpentineendemism in the flora of Puerto Rico Carib J Sci 32348ndash356

Coney P J 1982 Plate tectonic constraints on the biogeogra-phy of Middle America and the Caribbean region AnnMissouri Bot Gard 69 432ndash443

Correll D S amp Correll H B 1982 Flora of the BahamaArchipelago (including the Turk and Caicos Islands) JCramer Vaduz

Cox A 1973 Plate Tectonics and Geomagnetic Reversals WH Freeman San Francisco

Croizat L 1952 Manual of Phytogeography Or an Accountof Plant Dispersal throughout the World N V Drukkerij

Hooiberg EPE The HagueCroizat L 1958 Panbiogeography Published by the author

CaracasDonovan S K amp Jackson T A 1994 Caribbean Geology

an Introduction Univ West Indies KingstonDraper G amp J A Barros 1994 Cuba Pp 65ndash86 in

Donovan S K amp Jackson T A (eds) CaribbeanGeology an Introduction Univ West Indies Kingston

Draper G amp Lewis J F 1990 Geology and tectonic evolu-tion of the northern Caribbean margin Pp 1ndash14 inDengo G amp Case J E (eds) The Geology of NorthAmerica Vol H The Caribbean Region Geol SocAmerica Boulder

Eggers B H F A 1879 The Flora of St Croix and the VirginIslands Government Printing Office Washington DC

Erard C 1991 Landbirds of the Lesser Antilles ComptRend Sommaire Seacuteances Soc Biogeacuteogr 67 3ndash23

Fay M F Olmstead R G Richardson J E Santiago EPrance G T amp Chase M W 1998 Molecular data sup-port the inclusion of Duckeodendron cestroides in theSolanaceae Kew Bull 53 203ndash212

Fritsch P W 2003 Multiple geographic origins of AntilleanStyrax Syst Bot 28 421ndash430

Fritsch P W amp McDowell T D 2003 Biogeography andphylogeny of Caribbean plantsmdashintroduction Syst Bot28 376ndash377

Gentry A H 1982 Neotropical floristic diversity phytogeo-graphical connections between Central and SouthAmerica Pleistocene climatic fluctuations or an accidentof the Andean orogeny Ann Missouri Bot Gard 69557ndash593

Good R 1964 The Geography of the Flowering PlantsWiley New York

Graham A 1988 Some aspects of the Tertiary history in theGulfCaribbean region Transactions of the 11thCaribbean Geological Conference Barbados 3 1ndash18

Graham A 1989a Late Tertiary paleoaltitudes and vegeta-tional zonation in Mexico and Central America Acta BotNeerl 38 417ndash424

Graham A 1989b Paleofloristic and paleoclimatic changes inthe Tertiary of northern Latin America Rev PaleobotanyPalynology 60 283ndash293

Graham A 1989c Studies in neotropical paleobotany VIIThe lower Miocene communities of Panamandashthe La BocaFormation Ann Missouri Bot Gard 76 50ndash66

Graham A 1990a Late Tertiary microfossil flora from therepublic of Haiti Amer J Bot 77 911ndash926

Graham A 1990b New angiosperm records from theCaribbean Tertiary Amer J Bot 77 897ndash910

Graham A 1993 Contribution toward a Tertiary palyno-stratigraphy for Jamaica the status of Tertiary paleobotan-ical studies in northern Latin America and preliminaryanalysis of the Guys Hill Member (Chapelton Formationmiddle Eocene) of Jamaica Mem Geol Soc America182 443ndash461

Graham A 1996 Paleophysiographic and paleoenvironmen-tal histories in northern Latin Americandashpossible impact onmodes of speciation Paleobotanist 45 143ndash147

Graham A 1999a Studies in neotropical paleobotany XIIIAn Oligo-Miocene palynoflora from Simojovel (ChiapasMexico) Amer J Bot 86 17ndash31

Graham A 1999b The Tertiary history of the northern tem-


316

perate element in the northern Latin America biota AmerJ Bot 86 32ndash38

Graham A 2003 Geohistory models and Cenozoic paleoen-vironments of the Caribbean region Syst Bot 28378ndash386

Graham A Cozadd D Areces-Mallea A amp FrederiksenN O 2000 Studies in Neotropical paleobotany XIV Apalynoflora from the Middle Eocene SaramaguacaacutenFormation of Cuba Amer J Bot 87 1526ndash1539

Graham A amp Dilcher D 1995 The Cenozoic record of trop-ical dry forest in northern Latin America and the southernUnited States Pp 124ndash145 in Bullock S H Mooney HA amp Medina E (eds) Seasonally Dry Tropical ForestsCambridge Univ Press Cambridge

Graham A amp Jarzen D M 1969 Studies in neotropicalpaleobotany I The Oligocene communities of PuertoRico Ann Missouri Bot Gard 56 308ndash357

Graham S A 2002 Phylogenetic relationships and biogeog-raphy of the endemic Caribbean genera Ginoria Haitiaand Crenea (Lythraceae) Carib J Sci 38 195ndash204

Graham S A 2003 Biogeographic patterns of AntilleanLythraceae Syst Bot 28 410ndash420

Hass C A 1996 Relationships among West Indian geckos ofthe genus Sphaerodactylus a preliminary analysis of mito-chondrial 16S ribosomal RNA sequences Pp 175ndash194 inPowell R amp Henderson R W (eds) Contributions toWest Indian Herpetology A Tribute to Albert SchwartzSoc Study Amphibians and Reptiles Ithaca New York

Hedges S B 1996 Historical biogeography of West Indianvertebrates Annu Rev Ecol Syst 27 163ndash196

Hedges S B 1999 Distributional patterns of amphibians inthe West Indies Pp 211ndash254 in Duellman W E (ed)Regional Patterns of Amphibian Distribution A GlobalPerspective John Hopkins University Press

Hedges S B Bezy R L amp Maxson L R 1991Phylogenetic relationships and biogeography of xantusiidlizards inferred from mitochondrial DNA sequencesMolec Biol Evol 8 767ndash780

Henderson A Galeano G amp Bernal R 1995 Field Guideto the Palms of the Americas Princeton Univ PressPrinceton New Jersey

Hess H Ed 1962 History of Ocean Basins PterologicalStudies A Volume in Honor of A F BuddingtonGeological Society of America Boulder

Hildebrand A R amp Boynton V W 1990 ProximalCretaceous-Tertiary boundary impact deposits in theCaribbean Science 248 843ndash847

Howard R A 1973 The Vegetation of the Antilles Pp 1ndash38in Graham A (ed) Vegetation and Vegetational Historyof Northern Latin America Elsevier ScientificAmsterdam

Howard R A 1974ndash1989 Flora of the Lesser AntillesArnold Arboretum of Harvard Univ Jamaica PlainMassachusetts

Huebeck C amp Mann C 1991 Structural geology andCenozoic tectonic history of the southeastern terminationof the Cordillera Central Dominican Republic Geol SocAmerica Special Paper 262 315ndash336

Humphries C J 1982 Vicariance biogeography inMesoamerica Ann Missouri Bot Gard 69 444ndash463

Hunziker A T 1979 South American Solanaceae a synopticsurvey Pp 49ndash86 in Hawkes J G Lester R N amp

Skelding A D (eds) The Biology and Taxonomy of theSolanaceae Academic Press London

Iturralde-Vinent M A 1994 Cuban geology a new platetectonic synthesis J Petrol Geol 17 39ndash70

Iturralde-Vinent M A amp MacPhee R D E 1999Paleogeography of the Caribbean region implications forCenozoic biogeography Bull Amer Mus Nat Hist 2381ndash95

Jackman T Losos J B Larson A amp de Queiroz K 1997Phylogenetic studies of convergent adaptative radiationsin Caribbean Anolis lizards Pp 535ndash557 in Givnish T Jamp Sytsma K J (eds) Molecular Evolution and AdaptiveRadiation Cambridge Univ Press New York

Judd W S 1981 A monograph of Lyonia (Ericaceae) JArnold Arb 62 63ndash128

Judd W S 1995 13 Lyonia Nuttall Pp 222ndash294 in LuteynJ L (ed) Ericaceae Part II mdash The Superior-OvariedGenera Flora Neotropica Monograph 66 New YorkBotanical Garden Bronx New York

Judd W S 2001 Phylogeny and Biogeography of Lyoniasect Lyonia (Ericaceae) Pp 63ndash75 in Woods C A ampSergile F E (eds) Biogeography of the West IndiesPatterns and Perspectives ed 2 CRC Press Boca Raton

Kuhlmann J G 1930 Contribuiccedilao para o conhecimento dealgumas novas especies de regiatildeo amazonica e uma do Riode Janeiro bem como algumas notas sobre especies jaacute con-hecidas Arch Jard Bot Rio de Janeiro 5 209ndash211

Kuhlmann J G 1947 Duckeodendraceae Kuhlmann (novafamilia) Arq Ser Florest 3 7ndash8

Ladd J W 1976 Relative motion of South America withrespect to North America and Caribbean tectonics BullGeol Soc America 87 969ndash976

Larue D K 1994 Puerto Rico and the Virgin Islands Pp151ndash166 in Donovan S K amp Jackson T A (eds)Caribbean Geology an Introduction Univ West IndiesKingston

Lavin M 1993 Biogeography and systematics of Poitea(Leguminosae) Syst Bot Monogr 37 1ndash87

Lavin M amp Luckow M 1993 Origins and relationships oftropical North America in the context of the Boreotropicshypothesis Amer J Bot 80 1ndash14

Lavin M Wojciechowski M F Gasson P Hughes C ampWheeler E 2003 Phylogeny of Robinioid legumes(Fabaceae) revisited Coursetia and Gliricidia recircum-scribed and a biogeographical appraisal of the Caribbeanendemics Syst Bot 28 387ndash409

Leoacuten H 1946 Flora de Cuba vol 1 Cultural S A LaHabana

Leoacuten H amp Alain H 1951ndash1957 Flora de Cuba vols 2ndash4Cultural S A La Habana

Lewis J F amp Draper G 1990 Geology and tectonic evolu-tion of the northern Caribbean margin Pp 77ndash140 inDengo G amp Case J E (eds) The Geology of NorthAmerica Vol H The Caribbean Region Geol Soc ofAmerica Boulder

Leyden B W 1984 Guatemalan forest synthesis afterPleistocene aridity Proc Natl Acad Sci USA 814856ndash4859

Liebherr J K 1988 The Caribbean fertile ground for zoo-geography Pp 121ndash152 in Liebherr J K (ed)Zoogeography of Caribbean Insects Cornell Univ PressIthaca


317

Liogier A H 1974 Flora de CubamdashSuplemento InstitutoCubano del Libro La Habana

Liogier A H 1982ndash1996 La Flora de la Espantildeola vols 1ndash8Univ Central del Este San Pedro de Macoris

Liogier A H amp Martorell L F 1982 Flora of Puerto Ricoand Adjacent Islands Univ de Puerto Rico Rio PiedrasPuerto Rico

Malfait B T amp Dinkelman M G 1972 Circum-Caribbeantectonic and igneous activity and the evolution of theCaribbean plate Bull Geol Soc Amer 83 251ndash272

Marticorena M Stuessy T F amp Baeza C M 1998Cataacutelogo de la flora vascular de las Islas Robinson Crusoeo Juaacuten Fernaacutendez Chile Gayana Bot 55 187ndash211

McDowell T amp Bremer B 1998 Phylogeny diversity anddistribution in Exostema (Rubiaceae) implications ofmorphological and molecular analyses Pl Syst Evol 212215ndash246

McDowell T Volovsek M amp Manos P 2003Biogeography of Exostema (Rubiaceae) in the Caribbeanregion in light of molecular phylogenetic analyses SystBot 28 431ndash441

Morgan W J 1968 Rises trenches great faults and crustalblocks J Geophys Res 73 1959ndash1982

Morin N R 1982 Biological studies in Central America Thetwenty-eighth annual Systematics Symposium AnnMissouri Bot Gard 69 431

Morrone J J amp Crisci J V 1995 Historical biogeographyintroduction to methods Annu Rev Ecol Syst 26373ndash401

Myers N Mittermeier R A Mittermeier C G daFonseca G A B amp Kent J 2000 Biodiversity hotspotsfor conservation priorities Nature 403 853ndash858

Negroacuten-Ortiz V amp Watson L E 2002 Molecular phyloge-ny and biogeography of Erithalis (Rubiaceae) an endem-ic of the Caribbean Basin Pl Syst Evol 234 71ndash83

Negroacuten-Ortiz V amp Watson L E 2003 Hypotheses for thecolonization of the Caribbean Basin by two genera of theRubiaceae Erithalis and Ernodea Syst Bot 28 442ndash451

Olmstead R G Sweere J A Spangler R E Bohs L ampPalmer J D 1999 Phylogeny and provisional classifica-tion of the Solanaceae based on chloroplast DNA Pp111ndash137 in Nee M amp Symon D (eds) Solanaceae IVAdvances in Taxonomy and Utilization Royal BotanicGardens Kew

Page R G M amp Lydeard C 1994 Towards a cladistic bio-geography of the Caribbean Cladistics 10 21ndash41

Perfit M R amp Heenzen B C 1978 The geology and evolu-tion of the Cayman Trench Bull Geol Soc America 891155ndash1174

Perfit M R amp Williams E E 1989 Geological constraintsand biological retrodictions in the evolution of theCaribbean Sea and its islands Pp 47ndash102 in Woods C A(ed) Biogeography of the West Indies Past Present andFuture Sandhill Crane Press Gainesville

Pindell J 1994 Evolution of the Gulf of Mexico and theCaribbean Pp 13ndash40 in Donovan S K amp Jackson T A(eds) Caribbean Geology an Introduction Univ WestIndies Kingston

Pindell J amp Barrett S F 1990 Geological evolution of theCaribbean a plate tectonic perspective Pp 405ndash432 inDengo G amp Case J E (eds) The Geology of NorthAmerica Vol H The Caribbean Region Geol Soc

America BoulderPindell J amp Dewey J F 1982 Permo-Triassic reconstruction

of western Pangea and the evolution of the Gulf ofMexico-Caribbean region Tectonics 1 179ndash211

Polhill R M amp Souza M 1981 Robinieae Pp 283ndash288 inPolhill R M amp Raven P H (eds) Advances in LegumeSystematics Part 1 Royal Botanic Gardens Kew

Proctor G R 1984 Flora of the Cayman Islands HerMajestyrsquos Stationary Office London

Robinson E 1988 Late Cretaceous and early Tertiary sedi-mentary rocks of the Central Inlier Jamaica J Geol SocJamaica 24 49ndash67

Robinson E 1994 Jamaica Pp 111ndash127 in Donovan S ampJackson T A (eds) Caribbean Geology an IntroductionUniv West Indies Kings

Rosen D E 1976 A vicariance model of Caribbean biogeog-raphy Syst Zool 24 431ndash464

Rosen D E 1985 Geological hierarchies and biogeographiccongruence in the Caribbean Ann Missouri Bot Gard72 636ndash659

Roughgarden J 1995 Anolis Lizards of the CaribbeanOxford New York

Salzman V T amp Judd W S 1995 A revision of the GreaterAntillean species of Bactris (Bactridinae Arecaceae)Brittonia 47 345ndash371

Sanders R W 1991 Cladistics of Bactris (Palmae) survey ofcharacters and refutation of Burretrsquos classificationSelbyana 12 105ndash133

Santiago-Valentiacuten E amp Olmstead R G 2003 Phylogenet-ics of the Antillean Goetzeoideae (Solanaceae) and theirrelationships within the Solanaceae based on chloroplastand ITS DNA sequence data Syst Bot 28 452ndash460

Savage J 1982 The enigma of the Central American herpeto-fauna dispersals or vicariance Ann Missouri Bot Gard69 464ndash547

Schubart C D Diesel R amp Hedges S B 1998 Rapid evo-lution to terrestrial life in Jamaican crabs Nature 393363ndash365

Schuchert C 1935 Historical Geology of the Antillean-Caribbean Region or the Lands Bordering the Gulf ofMexico and the Caribbean Sea Wiley amp Sons New York

Seidel M E 1996 Current status of biogeography of the WestIndian turtles in the genus Trachemys (Emydidae) Pp169ndash174 in Powell R amp Henderson R W (eds)Contributions to West Indian Herpetology A Tribute toAlbert Schwartz Soc Study Amphibians and ReptilesIthaca New York

Skean J D 1993 Monograph of Mecranium (Melastomata-ceae-Miconieae) Syst Bot Monogr 39 1ndash116

Sykes L R McCann W R amp Kafka A L 1982 Motionof Caribbean plate during the last 7 million years andimplications for earlier Cenozoic movements J GeophysRes 87 10656ndash10676

Tuzo-Wilson J 1965 A new class of faults and their bearingon continental drift Nature 207 343ndash347

Uhl N W amp Dransfield J 1987 Genera PalmarumInternational Palm Society L H Bailey HortoriumIthaca

Urban I 1923 Zur Pflanzengeographie von HispaniolaFratres Borntraeger Lipsiae

Wadge G 1994 The Lesser Antilles Pp 167ndash178 inDonovan S K amp Jackson T A (eds) Caribbean


318

Geology an Introduction Univ West Indies JamaicaKingston

Wendt T 1993 Composition floristic affinities and originsof the canopy tree flora of the Mexican Atlantic slope rainforests Pp 595ndash680 in Ramamoorthy T P Bye R LotA amp Fa J (eds) Biological Diversity of Mexico Originsand Distribution Oxford Univ Press New York

Williams E E 1989 Old problems and new opportunities inWest Indian Biogeography Pp 1ndash46 in Woods C A(ed) Biogeography of the West Indies Past Present andFuture Sandhill Crane Press Gainesville

Woods C A (ed) 1989 Biogeography of the West IndiesPast Present and Future Sandhill Crane PressGainesville

Woods C A amp Sergile F E (eds) 2001 Biogeography ofthe West Indies Patterns and Perspectives ed 2 CRCPress Boca Raton

Woodson R E Jr 1940 The Apocynaceous flora of theYucatan Peninsula Carnegie Inst Washington Publ 52262ndash102

WWF amp IUCN 1994 1995 1997 Centres of Plant DiversityA Strategy for their Conservation IUCN publications unitCambridge UK

Zona S 1990 A monograph of Sabal (ArecaceaeCoryphoideae) Aliso 12 583ndash666

Zona S amp Henderson A 1989 A review of animal-mediat-ed seed dispersal in palms Selbyana 11 6ndash21


319

Historically zoological studies have dominated thescene of Caribbean evolution and biogeography This isstill the case today where new contributions range overa diversity of vertebrate groups including birds (egErard 1991 Bermingham 1994) mammals (eg Breuilamp Masson 1991) and especially reptiles (eg Savage1982 Hedges amp al 1991 Roughgarden 1995 Hass1996 Seidel 1996 Jackman amp al 1997) and amphib-ians (eg Bogart amp Hedges 1995 Hedges 1999)Hedges (1996) reviewed the current knowledge on thehistorical biogeography of West Indian vertebratesZoogeographic studies of the region have diversified toinclude invertebrates (eg Liebherr 1988 Schubart ampal 1998) Although these numerous zoogeographic con-tributions have made the Caribbean one of the most fer-tile regions for the study of biogeography a comprehen-sive understanding of the evolution of the biota of theregion can only be achieved by inclusion of other groupsincluding plants

Interest in applying phylogenetics to study the his-torical biogeography of Caribbean plants has increasedin recent years as evidenced by published works (egJudd 1981 2001 Zona 1990 Lavin amp Luckow 1993Skean 1993 Salzman amp Judd 1995 McDowell ampBremer 1998) and a recent symposium (Fritsch amp Mc-Dowell 2003) In order to stimulate further interestamong students and newcomers we present in this workan outline on the present understanding of Caribbeanphytogeographymdashwith emphasis on the Greater AntillesWe provide general summaries on fossils floristic dataand with emphasis on case studies of Caribbean plantgroups where a phylogenetic approach has been incorpo-rated We also discuss considerations for applying thisapproach to test hypotheses on evolutionary processesand biogeography in other plant groups of the region

Several names are used to define different but over-lapping geographic regions mentioned in this reviewHence we define their use in this work for clarity Theterms ldquoGreater Caribbeanrdquo or ldquoCaribbean Basinrdquo refercollectively to the islands and the neighboring continen-tal regions along the Gulf of Mexico Central Americaand Northern South America The main islands of theCaribbean belong to three different archipelagos TheGreater Antilles the Lesser Antilles and the Bahamas(Fig 1) The Greater Antilles are composed of the fourlargest islands in the region Cuba Hispaniola Jamaicaand Puerto Rico The Lesser Antilles is separated fromthe Greater Antilles by the Anegada Passage and com-prise the arc of smaller islands extending from Sombreroin the north to Grenada Collectively the three archipel-agos are referred to as the Caribbean Islands or the WestIndies This paper covers issues relevant to the entireCaribbean with emphasis on the island systems

PLANT DIVERSITY IN THECARIBBEAN

The Caribbean is the oldest Neotropical region stud-ied by western science The first observations on plantdiversity are documented in Columbusrsquo diary and otherchroniclers of the 15th and 16th centuries with plantdescriptions predating Linneausrsquo Species Plantarum of1753 The long plant collecting history has resulted innumerous new taxa as well as published Floras includ-ing among the most recent ones for Cuba (Leoacuten 1946Leoacuten amp Alain 1951ndash1957 Alain 1963 Liogier 1974)Hispaniola (Liogier 1982ndash1996) Jamaica (Adams1972) Puerto Rico and the Virgin Islands (Acevedo-Rodriacuteguez 1996 Liogier amp Martorell 1982) theCayman Islands (Proctor 1984) the Bahamas (Correll ampCorrell 1982) and the Lesser Antilles (Howard1974ndash1989) Ongoing botanical explorations keep refin-ing present knowledge with the publication of florulasand checklists the report of new records and thedescription of new species

Plant diversity is so remarkable in the Caribbeanislands that the region is considered a distinctive phyto-geographic unit within the Neotropics (Gentry 1982)The vascular plant flora of the region consists of approx-imately 12000 species including the southern tip ofFlorida (Myers amp al 2000 Table 1) in about 200 fami-lies Over fifty percent of the vascular species are endem-ic to the region These endemics represent about 2 ofall vascular plants on Earth placing the Caribbean as oneof the leading hotspots in terms of species-levelendemism (Myers amp al 2000) Endemicity is also pres-ent at the generic level in the Greater Antilles Bothgeneric and species-level endemism are particularlyprominent in the two larger islands of Cuba andHispaniola (Gentry 1982) Cuba the largest island in theCaribbean has the richest flora and highest proportion ofendemism (Table 2) About half of its approximately


300

Fig 1 Map of the Caribbean region






301















302














303








304








305












306







307







Coccothrinax

Thrinax

Chelyocarpus

Crysophila

Trithrinax

S minor

S miamiensis

S etonia

S palmetto

S bermudana

S uresana

S rosei

S pumos

S yapa

S mauritiiformis

S guatemalensis

S mexicana

S maritima

S domingensis

S causiarum

Antilles

S America

C America

S America


Florida


Bermuda

W Mexico

Mexico

S Mexico Guatemala


S Mexico W Cuba

Cuba Jamaica

N Hispaniola


308









309








310


M acuminatum

M amygdalinum

N Hispaniola

M crassinerve

M haitiense

M purpurascens

M racemosum


M birimosum

M microdyctium

M puberulum

M haemanthum

M tuberculatum

N amp SW Hispaniola

SW Hispaniola

Jamaica

W Cuba



SW Hispaniola

N Hispaniola

W amp E Cuba


sect Sagreoides

sect M

ecra

nium






311


P emarginata

P multiflora

P glyciphylla

P galegoides

P campanilla

P punicea

P florida

P carinalis

P paucifolia

P dubia

P gracilis

Gliricidia

Lennea

Hebestigma

Olneya

Poissonia

Robinia

Coursetia

Genistidium

Sphinctospermum

Peteria

W Cuba

SW C Hispaniola

SW Hispaniola

Hispaniola

C Hispaniola

Puerto Rico

Dominica


C Hispaniola

Cuba


3 spp C America

1 sp Cuba

1 sp SW N America

4 spp S America

4 spp N America


1 sp Mexico

1 sp SW N America

4 spp SW N America









312







313


E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca


Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola


S America


Peru

Hispaniola

Cuba Hispaniola




Hispaniola

Cuba

2 spp Cuba

20 spp Antilles


1 sp Antilles


E lancifolium

E purpureum Cuba

Cuba


E stenophyllum Cuba








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319






301















302














303








304








305












306







307







Coccothrinax

Thrinax

Chelyocarpus

Crysophila

Trithrinax

S minor

S miamiensis

S etonia

S palmetto

S bermudana

S uresana

S rosei

S pumos

S yapa

S mauritiiformis

S guatemalensis

S mexicana

S maritima

S domingensis

S causiarum

Antilles

S America

C America

S America


Florida


Bermuda

W Mexico

Mexico

S Mexico Guatemala


S Mexico W Cuba

Cuba Jamaica

N Hispaniola


308









309








310


M acuminatum

M amygdalinum

N Hispaniola

M crassinerve

M haitiense

M purpurascens

M racemosum


M birimosum

M microdyctium

M puberulum

M haemanthum

M tuberculatum

N amp SW Hispaniola

SW Hispaniola

Jamaica

W Cuba



SW Hispaniola

N Hispaniola

W amp E Cuba


sect Sagreoides

sect M

ecra

nium






311


P emarginata

P multiflora

P glyciphylla

P galegoides

P campanilla

P punicea

P florida

P carinalis

P paucifolia

P dubia

P gracilis

Gliricidia

Lennea

Hebestigma

Olneya

Poissonia

Robinia

Coursetia

Genistidium

Sphinctospermum

Peteria

W Cuba

SW C Hispaniola

SW Hispaniola

Hispaniola

C Hispaniola

Puerto Rico

Dominica


C Hispaniola

Cuba


3 spp C America

1 sp Cuba

1 sp SW N America

4 spp S America

4 spp N America


1 sp Mexico

1 sp SW N America

4 spp SW N America









312







313


E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca


Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola


S America


Peru

Hispaniola

Cuba Hispaniola




Hispaniola

Cuba

2 spp Cuba

20 spp Antilles


1 sp Antilles


E lancifolium

E purpureum Cuba

Cuba


E stenophyllum Cuba








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319








302














303








304








305












306







307







Coccothrinax

Thrinax

Chelyocarpus

Crysophila

Trithrinax

S minor

S miamiensis

S etonia

S palmetto

S bermudana

S uresana

S rosei

S pumos

S yapa

S mauritiiformis

S guatemalensis

S mexicana

S maritima

S domingensis

S causiarum

Antilles

S America

C America

S America


Florida


Bermuda

W Mexico

Mexico

S Mexico Guatemala


S Mexico W Cuba

Cuba Jamaica

N Hispaniola


308









309








310


M acuminatum

M amygdalinum

N Hispaniola

M crassinerve

M haitiense

M purpurascens

M racemosum


M birimosum

M microdyctium

M puberulum

M haemanthum

M tuberculatum

N amp SW Hispaniola

SW Hispaniola

Jamaica

W Cuba



SW Hispaniola

N Hispaniola

W amp E Cuba


sect Sagreoides

sect M

ecra

nium






311


P emarginata

P multiflora

P glyciphylla

P galegoides

P campanilla

P punicea

P florida

P carinalis

P paucifolia

P dubia

P gracilis

Gliricidia

Lennea

Hebestigma

Olneya

Poissonia

Robinia

Coursetia

Genistidium

Sphinctospermum

Peteria

W Cuba

SW C Hispaniola

SW Hispaniola

Hispaniola

C Hispaniola

Puerto Rico

Dominica


C Hispaniola

Cuba


3 spp C America

1 sp Cuba

1 sp SW N America

4 spp S America

4 spp N America


1 sp Mexico

1 sp SW N America

4 spp SW N America









312







313


E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca


Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola


S America


Peru

Hispaniola

Cuba Hispaniola




Hispaniola

Cuba

2 spp Cuba

20 spp Antilles


1 sp Antilles


E lancifolium

E purpureum Cuba

Cuba


E stenophyllum Cuba








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319











303








304








305












306







307







Coccothrinax

Thrinax

Chelyocarpus

Crysophila

Trithrinax

S minor

S miamiensis

S etonia

S palmetto

S bermudana

S uresana

S rosei

S pumos

S yapa

S mauritiiformis

S guatemalensis

S mexicana

S maritima

S domingensis

S causiarum

Antilles

S America

C America

S America


Florida


Bermuda

W Mexico

Mexico

S Mexico Guatemala


S Mexico W Cuba

Cuba Jamaica

N Hispaniola


308









309








310


M acuminatum

M amygdalinum

N Hispaniola

M crassinerve

M haitiense

M purpurascens

M racemosum


M birimosum

M microdyctium

M puberulum

M haemanthum

M tuberculatum

N amp SW Hispaniola

SW Hispaniola

Jamaica

W Cuba



SW Hispaniola

N Hispaniola

W amp E Cuba


sect Sagreoides

sect M

ecra

nium






311


P emarginata

P multiflora

P glyciphylla

P galegoides

P campanilla

P punicea

P florida

P carinalis

P paucifolia

P dubia

P gracilis

Gliricidia

Lennea

Hebestigma

Olneya

Poissonia

Robinia

Coursetia

Genistidium

Sphinctospermum

Peteria

W Cuba

SW C Hispaniola

SW Hispaniola

Hispaniola

C Hispaniola

Puerto Rico

Dominica


C Hispaniola

Cuba


3 spp C America

1 sp Cuba

1 sp SW N America

4 spp S America

4 spp N America


1 sp Mexico

1 sp SW N America

4 spp SW N America









312







313


E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca


Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola


S America


Peru

Hispaniola

Cuba Hispaniola




Hispaniola

Cuba

2 spp Cuba

20 spp Antilles


1 sp Antilles


E lancifolium

E purpureum Cuba

Cuba


E stenophyllum Cuba








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319








304








305












306







307







Coccothrinax

Thrinax

Chelyocarpus

Crysophila

Trithrinax

S minor

S miamiensis

S etonia

S palmetto

S bermudana

S uresana

S rosei

S pumos

S yapa

S mauritiiformis

S guatemalensis

S mexicana

S maritima

S domingensis

S causiarum

Antilles

S America

C America

S America


Florida


Bermuda

W Mexico

Mexico

S Mexico Guatemala


S Mexico W Cuba

Cuba Jamaica

N Hispaniola


308









309








310


M acuminatum

M amygdalinum

N Hispaniola

M crassinerve

M haitiense

M purpurascens

M racemosum


M birimosum

M microdyctium

M puberulum

M haemanthum

M tuberculatum

N amp SW Hispaniola

SW Hispaniola

Jamaica

W Cuba



SW Hispaniola

N Hispaniola

W amp E Cuba


sect Sagreoides

sect M

ecra

nium






311


P emarginata

P multiflora

P glyciphylla

P galegoides

P campanilla

P punicea

P florida

P carinalis

P paucifolia

P dubia

P gracilis

Gliricidia

Lennea

Hebestigma

Olneya

Poissonia

Robinia

Coursetia

Genistidium

Sphinctospermum

Peteria

W Cuba

SW C Hispaniola

SW Hispaniola

Hispaniola

C Hispaniola

Puerto Rico

Dominica


C Hispaniola

Cuba


3 spp C America

1 sp Cuba

1 sp SW N America

4 spp S America

4 spp N America


1 sp Mexico

1 sp SW N America

4 spp SW N America









312







313


E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca


Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola


S America


Peru

Hispaniola

Cuba Hispaniola




Hispaniola

Cuba

2 spp Cuba

20 spp Antilles


1 sp Antilles


E lancifolium

E purpureum Cuba

Cuba


E stenophyllum Cuba








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319








305












306







307







Coccothrinax

Thrinax

Chelyocarpus

Crysophila

Trithrinax

S minor

S miamiensis

S etonia

S palmetto

S bermudana

S uresana

S rosei

S pumos

S yapa

S mauritiiformis

S guatemalensis

S mexicana

S maritima

S domingensis

S causiarum

Antilles

S America

C America

S America


Florida


Bermuda

W Mexico

Mexico

S Mexico Guatemala


S Mexico W Cuba

Cuba Jamaica

N Hispaniola


308









309








310


M acuminatum

M amygdalinum

N Hispaniola

M crassinerve

M haitiense

M purpurascens

M racemosum


M birimosum

M microdyctium

M puberulum

M haemanthum

M tuberculatum

N amp SW Hispaniola

SW Hispaniola

Jamaica

W Cuba



SW Hispaniola

N Hispaniola

W amp E Cuba


sect Sagreoides

sect M

ecra

nium






311


P emarginata

P multiflora

P glyciphylla

P galegoides

P campanilla

P punicea

P florida

P carinalis

P paucifolia

P dubia

P gracilis

Gliricidia

Lennea

Hebestigma

Olneya

Poissonia

Robinia

Coursetia

Genistidium

Sphinctospermum

Peteria

W Cuba

SW C Hispaniola

SW Hispaniola

Hispaniola

C Hispaniola

Puerto Rico

Dominica


C Hispaniola

Cuba


3 spp C America

1 sp Cuba

1 sp SW N America

4 spp S America

4 spp N America


1 sp Mexico

1 sp SW N America

4 spp SW N America









312







313


E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca


Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola


S America


Peru

Hispaniola

Cuba Hispaniola




Hispaniola

Cuba

2 spp Cuba

20 spp Antilles


1 sp Antilles


E lancifolium

E purpureum Cuba

Cuba


E stenophyllum Cuba








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319












306







307







Coccothrinax

Thrinax

Chelyocarpus

Crysophila

Trithrinax

S minor

S miamiensis

S etonia

S palmetto

S bermudana

S uresana

S rosei

S pumos

S yapa

S mauritiiformis

S guatemalensis

S mexicana

S maritima

S domingensis

S causiarum

Antilles

S America

C America

S America


Florida


Bermuda

W Mexico

Mexico

S Mexico Guatemala


S Mexico W Cuba

Cuba Jamaica

N Hispaniola


308









309








310


M acuminatum

M amygdalinum

N Hispaniola

M crassinerve

M haitiense

M purpurascens

M racemosum


M birimosum

M microdyctium

M puberulum

M haemanthum

M tuberculatum

N amp SW Hispaniola

SW Hispaniola

Jamaica

W Cuba



SW Hispaniola

N Hispaniola

W amp E Cuba


sect Sagreoides

sect M

ecra

nium






311


P emarginata

P multiflora

P glyciphylla

P galegoides

P campanilla

P punicea

P florida

P carinalis

P paucifolia

P dubia

P gracilis

Gliricidia

Lennea

Hebestigma

Olneya

Poissonia

Robinia

Coursetia

Genistidium

Sphinctospermum

Peteria

W Cuba

SW C Hispaniola

SW Hispaniola

Hispaniola

C Hispaniola

Puerto Rico

Dominica


C Hispaniola

Cuba


3 spp C America

1 sp Cuba

1 sp SW N America

4 spp S America

4 spp N America


1 sp Mexico

1 sp SW N America

4 spp SW N America









312







313


E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca


Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola


S America


Peru

Hispaniola

Cuba Hispaniola




Hispaniola

Cuba

2 spp Cuba

20 spp Antilles


1 sp Antilles


E lancifolium

E purpureum Cuba

Cuba


E stenophyllum Cuba








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319







307







Coccothrinax

Thrinax

Chelyocarpus

Crysophila

Trithrinax

S minor

S miamiensis

S etonia

S palmetto

S bermudana

S uresana

S rosei

S pumos

S yapa

S mauritiiformis

S guatemalensis

S mexicana

S maritima

S domingensis

S causiarum

Antilles

S America

C America

S America


Florida


Bermuda

W Mexico

Mexico

S Mexico Guatemala


S Mexico W Cuba

Cuba Jamaica

N Hispaniola


308









309








310


M acuminatum

M amygdalinum

N Hispaniola

M crassinerve

M haitiense

M purpurascens

M racemosum


M birimosum

M microdyctium

M puberulum

M haemanthum

M tuberculatum

N amp SW Hispaniola

SW Hispaniola

Jamaica

W Cuba



SW Hispaniola

N Hispaniola

W amp E Cuba


sect Sagreoides

sect M

ecra

nium






311


P emarginata

P multiflora

P glyciphylla

P galegoides

P campanilla

P punicea

P florida

P carinalis

P paucifolia

P dubia

P gracilis

Gliricidia

Lennea

Hebestigma

Olneya

Poissonia

Robinia

Coursetia

Genistidium

Sphinctospermum

Peteria

W Cuba

SW C Hispaniola

SW Hispaniola

Hispaniola

C Hispaniola

Puerto Rico

Dominica


C Hispaniola

Cuba


3 spp C America

1 sp Cuba

1 sp SW N America

4 spp S America

4 spp N America


1 sp Mexico

1 sp SW N America

4 spp SW N America









312







313


E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca


Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola


S America


Peru

Hispaniola

Cuba Hispaniola




Hispaniola

Cuba

2 spp Cuba

20 spp Antilles


1 sp Antilles


E lancifolium

E purpureum Cuba

Cuba


E stenophyllum Cuba








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319







Coccothrinax

Thrinax

Chelyocarpus

Crysophila

Trithrinax

S minor

S miamiensis

S etonia

S palmetto

S bermudana

S uresana

S rosei

S pumos

S yapa

S mauritiiformis

S guatemalensis

S mexicana

S maritima

S domingensis

S causiarum

Antilles

S America

C America

S America


Florida


Bermuda

W Mexico

Mexico

S Mexico Guatemala


S Mexico W Cuba

Cuba Jamaica

N Hispaniola


308









309








310


M acuminatum

M amygdalinum

N Hispaniola

M crassinerve

M haitiense

M purpurascens

M racemosum


M birimosum

M microdyctium

M puberulum

M haemanthum

M tuberculatum

N amp SW Hispaniola

SW Hispaniola

Jamaica

W Cuba



SW Hispaniola

N Hispaniola

W amp E Cuba


sect Sagreoides

sect M

ecra

nium






311


P emarginata

P multiflora

P glyciphylla

P galegoides

P campanilla

P punicea

P florida

P carinalis

P paucifolia

P dubia

P gracilis

Gliricidia

Lennea

Hebestigma

Olneya

Poissonia

Robinia

Coursetia

Genistidium

Sphinctospermum

Peteria

W Cuba

SW C Hispaniola

SW Hispaniola

Hispaniola

C Hispaniola

Puerto Rico

Dominica


C Hispaniola

Cuba


3 spp C America

1 sp Cuba

1 sp SW N America

4 spp S America

4 spp N America


1 sp Mexico

1 sp SW N America

4 spp SW N America









312







313


E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca


Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola


S America


Peru

Hispaniola

Cuba Hispaniola




Hispaniola

Cuba

2 spp Cuba

20 spp Antilles


1 sp Antilles


E lancifolium

E purpureum Cuba

Cuba


E stenophyllum Cuba








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319









309








310


M acuminatum

M amygdalinum

N Hispaniola

M crassinerve

M haitiense

M purpurascens

M racemosum


M birimosum

M microdyctium

M puberulum

M haemanthum

M tuberculatum

N amp SW Hispaniola

SW Hispaniola

Jamaica

W Cuba



SW Hispaniola

N Hispaniola

W amp E Cuba


sect Sagreoides

sect M

ecra

nium






311


P emarginata

P multiflora

P glyciphylla

P galegoides

P campanilla

P punicea

P florida

P carinalis

P paucifolia

P dubia

P gracilis

Gliricidia

Lennea

Hebestigma

Olneya

Poissonia

Robinia

Coursetia

Genistidium

Sphinctospermum

Peteria

W Cuba

SW C Hispaniola

SW Hispaniola

Hispaniola

C Hispaniola

Puerto Rico

Dominica


C Hispaniola

Cuba


3 spp C America

1 sp Cuba

1 sp SW N America

4 spp S America

4 spp N America


1 sp Mexico

1 sp SW N America

4 spp SW N America









312







313


E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca


Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola


S America


Peru

Hispaniola

Cuba Hispaniola




Hispaniola

Cuba

2 spp Cuba

20 spp Antilles


1 sp Antilles


E lancifolium

E purpureum Cuba

Cuba


E stenophyllum Cuba








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319








310


M acuminatum

M amygdalinum

N Hispaniola

M crassinerve

M haitiense

M purpurascens

M racemosum


M birimosum

M microdyctium

M puberulum

M haemanthum

M tuberculatum

N amp SW Hispaniola

SW Hispaniola

Jamaica

W Cuba



SW Hispaniola

N Hispaniola

W amp E Cuba


sect Sagreoides

sect M

ecra

nium






311


P emarginata

P multiflora

P glyciphylla

P galegoides

P campanilla

P punicea

P florida

P carinalis

P paucifolia

P dubia

P gracilis

Gliricidia

Lennea

Hebestigma

Olneya

Poissonia

Robinia

Coursetia

Genistidium

Sphinctospermum

Peteria

W Cuba

SW C Hispaniola

SW Hispaniola

Hispaniola

C Hispaniola

Puerto Rico

Dominica


C Hispaniola

Cuba


3 spp C America

1 sp Cuba

1 sp SW N America

4 spp S America

4 spp N America


1 sp Mexico

1 sp SW N America

4 spp SW N America









312







313


E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca


Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola


S America


Peru

Hispaniola

Cuba Hispaniola




Hispaniola

Cuba

2 spp Cuba

20 spp Antilles


1 sp Antilles


E lancifolium

E purpureum Cuba

Cuba


E stenophyllum Cuba








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319






311


P emarginata

P multiflora

P glyciphylla

P galegoides

P campanilla

P punicea

P florida

P carinalis

P paucifolia

P dubia

P gracilis

Gliricidia

Lennea

Hebestigma

Olneya

Poissonia

Robinia

Coursetia

Genistidium

Sphinctospermum

Peteria

W Cuba

SW C Hispaniola

SW Hispaniola

Hispaniola

C Hispaniola

Puerto Rico

Dominica


C Hispaniola

Cuba


3 spp C America

1 sp Cuba

1 sp SW N America

4 spp S America

4 spp N America


1 sp Mexico

1 sp SW N America

4 spp SW N America









312







313


E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca


Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola


S America


Peru

Hispaniola

Cuba Hispaniola




Hispaniola

Cuba

2 spp Cuba

20 spp Antilles


1 sp Antilles


E lancifolium

E purpureum Cuba

Cuba


E stenophyllum Cuba








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319









312







313


E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca


Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola


S America


Peru

Hispaniola

Cuba Hispaniola




Hispaniola

Cuba

2 spp Cuba

20 spp Antilles


1 sp Antilles


E lancifolium

E purpureum Cuba

Cuba


E stenophyllum Cuba








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319






313


E myrtifolium

E ixoroides

E parviflorum

E mexicanum

E lineatum

E longiflorum

E ellipticum

E maynense

Coutarea

E corymbosum

E nitens

E spinosum

E caribaeum

Chiococca


Erithalis fruticosa

E acuminata

E salicifolia

Cubanola

Isadorea

Cephalanthus

Strumpfia

Cuba

Hispaniola

Mexico C America

Hispaniola

Cuba Hispaniola


S America


Peru

Hispaniola

Cuba Hispaniola




Hispaniola

Cuba

2 spp Cuba

20 spp Antilles


1 sp Antilles


E lancifolium

E purpureum Cuba

Cuba


E stenophyllum Cuba








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319








314


Cuba Hispaniola

Bahamas

Bahamas

Hispaniola

Dominica

Puerto Rico

St Vincent

St Vincent

Bahamas

Jamaica

Florida

Puerto Rico

Jamaica

Cuba

Jamaica

TropicalAmerica

E quadrangularis

E vacciniifolia

E diffusa

E salmeoides

E salmeoides

E odorifera

E harrisii


Chiococca alba

E fruticosa

E fruticosa

E fruticosa

E fruticosa

E odorifera

E odorifera

E odorifera

E odorifera

E odorifera










315
















































316








































317











































318











319










315
















































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317











































318











319















































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Documents

Historical biogeography of Caribbean plants: introduction ...geographic history of the Caribbean biota have ignited much interest, resulting in many studies addressing ... tributions