biogeography of the antilles based on a parsimony...

Biogeography of the Antilles basedon a parsimony analysis of orchid distributionsJ. Carlos Trejo-Torres and James D. Ackerman Department of Biology, University of Puerto

Rico-Rõ�o Piedras, PO BOX 23360, San Juan, Puerto Rico 00931-3360

Abstract

Aim We obtain biogeographical patterns based on the distributions of shared orchidspecies of the Caribbean. These patterns are used to de®ne biogeographical zones. Wethen analyse the concordance between the distributional patterns with ecological andphysical features of the islands.

Location We use orchid species recorded on 49 islands of the Greater, Lesser, andsouthern Antilles, and the Bahamas. Three continental areas are included: Florida(North America), the Yucatan (Central America), and the Guianas (South America).

Methods We use a parsimonious analysis of species distributions that produces the bestarrangements of shared taxa among areas. The analysis uses 356 shared orchid species ofthe 863 species recorded for studied areas. The methodology has been used to inferhistorical relationships among areas but we interpret the results as static or ecologicalpatterns of biogeographical af®nities.

Results Two kinds of island groupings are revealed. (1) Groups with common geologyand geomorphology: the Bahama Archipelago, the Virgin Islands, the Cayman Islandsand the southern Dutch Antilles. (2) An aggregation of distant islands with aheterogeneous geology but a common physiography: the Greater Antilles/Trinidad/Lesser Antilles/Margarita-Tobago. The Guianas are linked with the Greater Antilles,while the Yucatan and Florida are linked to the Bahamas.

Main conclusions Groupings of islands are congruent with their gross ecologicalfeatures either from similar geomorphology or common physiography. The strongaf®nity among islands considerably distant among each other is explained by the highvagility of dust-seeded orchids. Then, ¯oristic af®nities seem determined by ecologicalcharacteristics of islands rather than by dispersal barriers. We predict that other plantgroups with dust-like diaspores and animals with good vagility should show comparablebiogeographic patterns. Parsimony analysis of distributions (PAD) is an alternativemethodology to multivariate analysis to compare biotas, and a graphic complement toquantitative methods producing numerical values.

Keywords

Antilles, Caribbean, biogeography, Orchidaceae, parsimony analysis of endemicity,distributions, dispersal, islands.

INTRODUCTION

The Antilles, or West Indies, have been frequently studied bybiogeographers. These islands make up one of the largesttropical archipelagos in the world, second only to the islands

between Asia and Australia. The Antilles are also signi®cantbecause they represent one of the two connections, by way ofan island chain, between two major biogeographical realms:the Neartic and the Neotropic. The majority of studies onAntillean biogeography concern the distributional patternsor the cladistic biogeography of animals, especially verte-brates and insects (e.g. Liebherr, 1988; Woods, 1989;Page & Lydeard, 1994; Hedges, 1996). Surprisingly, thereare only a few publications on the phytogeography of this

Correspondence: J. Carlos Trejo-Torres, Department of Biology, University of

Puerto Rico-RõÂo Piedras, PO BOX 23360, San Juan, Puerto Rico 00931-3360.

E-mail: [email protected]

Journal of Biogeography, 28, 775±794

Ó 2001 Blackwell Science Ltd

region (Howard, 1973; Samek, 1988; Adams, 1989; Lavin,1993; Borhidi, 1996), although the ¯oristics of the area havebeen actively studied for decades (Zanoni, 1986; Liogier,1996).

The Caribbean region has been divided phytogeographi-cally in three subregions by Samek (1988). These main divi-sions are: (1) Mexico to Panama, (2) Colombia to Venezuela,and (3) an insular subregion including the Antilles proper.The southern Antilles, from Aruba to Tobago are consideredpart of the Colombian±Venezuelan subregion. Borhidi(1996) joins both continental subregions, from Mexico toVenezuela, into a single one. While these authors considersouth Florida as part of the Antillean subregion, Gentry(1982) included only the Florida Keys in the Antillean region.

Distinguishing biotic regions, or de®ning biologicalboundaries, has been among the major concerns of bioge-ographers. Traditionally, this has been performed withsubjective methods (e.g. Gentry, 1982; Samek, 1988;Borhidi, 1996). However, more objective or analyticalapproaches have been developed for the analysis of distri-butional data of organisms (e.g. Gauch, 1982; McCoy et al.,1986; Patterson & Atmar, 1986; Rosen & Smith, 1988;Rosen, 1988; Vargas, 1991; Real et al., 1992; Scheiner,1992; Worthen, 1996; Puente et al., 1998). Here, we use analternative method based on a parsimonious analysis of taxadistributions (Rosen & Smith, 1988). This method repre-sents a direct way to search for the biogeographical af®nitiesamong areas (Connor, 1988; Vargas, 1991), for the detec-tion of areas of endemism (Morrone, 1994a; Cardoso daSilva & Oren, 199622 ; Bates et al., 1998), and for thedelimitation of biological boundaries (Posadas, 1996;Posadas et al., 1997; Morrone, 1998). The parsimonyanalysis presented here is a tool for searching the mostparsimonious arrangement of shared species among areas, asa means of revealing the biogeographical af®nities in ahierarchical pattern (Rosen & Smith, 1988; Brady, 1994).

The analysis presented here was originally called parsi-mony analysis of endemicity (PAE), and was suggested byRosen (198533 , cited in Rosen & Smith, 1988) and developedby Rosen & Smith (1988). It was also independentlysuggested by Legendre (1986) and Connor (1988). Sincethen it has been employed in the study of extant taxa ofNew Zealand (Craw, 1988), Australia (Cracraft, 1991),Southeast Africa (Morrone, 1994a), the Patagonia(Posadas, 1996), the Andes (Morrone, 1994b; Posadaset al., 1997), the Amazonia (Cardoso da Silva & Oren,1996; Bates et al., 1998), the Austral region (Craw, 1989;Morrone, 1998; Glasby & Alvarez, 1999), Mexico (Lunaet al., 1999), and the entire world (Conran, 1995). Theunits of comparison that have been used are sites, quad-rants or sections of regions, biogeographical areas, ornatural geographical areas (e.g. islands, continents, oceanbasins). We use entire islands or groups of them, as theunits of study. While the method has been mainly used fordiscovering the historical relationships among areas, wegive a static or non-historical interpretation to the patternsobtained (Rosen, 1992; Posadas, 1996; Posadas et al.,1997). Although the method has been called PAE, we use a

more generic name: parsimony analysis of distributions(PAD). The method excludes single-site species (includingsingle-site endemics), and takes into account shared specieswhether endemic or not to the Antilles or adjacent areas.We substitute the term distribution for endemicity to avoidconfusion.

Our goal here is to distinguish biogeographical patterns inthe Antilles based on one plant family, Orchidaceae. Thisgroup of plants, of which there are approximately 700species in the Antilles, is relatively well known taxonomi-cally and geographically because there are relatively recentorchid treatments for the Cayman Islands (Proctor, 1984,1996), Puerto Rico (Ackerman, 1995) and the Bahamas(Sauleda & Adams, 1982; Sauleda, 1992). Furthermore, atreatment for the Greater Antilles is underway (Ackerman,1997; in press). The homogeneous wind-dispersal mechan-ism (except in bird-dispersed Vanilla) of the dust-seedsamong the species of this family, makes orchids a good focalgroup for the study of distribution. Assuming that mostorchids have the same dispersal capacity, their distributionalpatterns may be explained in terms of other ecological,geographical and historical factors.

The questions we address are: (1) what are the phyto-geographical relationships among the Antillean islandsbased on shared orchid species? (2) are these relationshipsaffected when neighbouring continental regions are consid-ered? (3) what is the phytogeographical regionalization ofthe Antilles based on orchids, and (4) how do the distribu-tional patterns of orchids match with geography, geology,physiography and ecology of the region? Apart from thebiogeographical analysis per se, we discuss the use ofparsimony analysis of naturally de®ned areas (i.e. islands)with extremely dissimilar number of species, and the use ofsingle-site species to look at ®ner degrees of biogeographicaldifferentiation.

METHODS

A total of ®fty-two areas (Table 1) were included in thestudy. Data on the distribution of species were taken fromliterature and from unpublished sources (Table 2). Synony-mies and valid species names were standardized mainlyaccording to J.D. Ackerman (unpublished data).

A presence/absence matrix of the 863 orchid speciesreported for the studied areas was constructed in MacClade3.01 (Maddison & Maddison, 1992). Using this program,areas were entered in the place of taxa, while taxa wereentered in the place of characters. In the matrix, presencewas indicated with a `1' and absence with a `0'. Once weconstructed the matrix, we ran analyses of parsimony usingPAUP 4 (Swofford, 2000). A hypothetical outgroup areawith all 0s (no species) were used in the analyses to root thetrees. General heuristic searches were carried out to look forthe most parsimonious trees, which indicate the ¯oristicaf®nities among studied areas. We obtained consensus trees(Strict, Majority Rule and Adams) when more than oneequally parsimonious tree were found. A description and adiscussion of the parsimonious analysis methodology are

Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794

776 J. C. Trejo-Torres and J. D. Ackerman

found in Rosen & Smith (1988), Vargas (1991) and Rosen(1988, 1992).

Only the informative species were taken into account inthe analyses (Appendix 1). The uninformative species arethose found in only one area; therefore, they give no

information in terms of shared species af®nities. We onlyused these single-site (island or area) species for speci®c¯oristic comparisons. Also, Vanilla species were not consid-ered at all because they are animal-dispersed, an anomaly inthe orchid family.

Area relationships derived from PAD may be altereddepending on the inclusion or exclusion of different areasjust as is the case when using different ingroup/outgrouptaxa in cladistic studies. To test the robustness of our data,we conducted independent analyses of different subsets ofareas.

RESULTS

An analysis of all areas using the 356 shared orchid speciesproduced 216 equally parsimonious trees. The three con-sensus trees obtained from them produced entirely congruentgroupings (Fig. 1). The Strict consensus tree shows thegroups found in all the most parsimonious trees. TheMajority Rule tree shows the groups found in more than50% of those trees. The Adams tree gives the highestpossible resolution for data distribution.

Four main clusters of areas can be identi®ed in theconsensus trees (Fig. 1): (1) Greater Antilles/Guianas±Trini-dad/larger Lesser Antilles/smaller Lesser Antilles/VirginIslands/Margarita-Tobago/other smaller Lesser Antilles,(2) Yucatan/Florida/Isla de la Juventud/Bahamas/Mona/Anegada, (3) Cayman Islands, and (4) Aruba/CuracËao/Bonaire. Although these are well-de®ned aggregations, thepolytomic base of the trees means that relationships amongthem are not resolved. The arrangement of areas within thegroupings is highly in¯uenced by species numbers.

DISCUSSION

Groupings of areas vs. physical factors

We found two kinds of grouping patterns in the Antillesbased on orchid species distributions. One pattern aggre-gates islands that belong to single geological units while theother pattern groups areas with different geological histories(Fig. 2).

In the ®rst pattern, the groupings are easy to understandas they correspond to islands belonging to de®ned geolo-gical units. The Virgin Islands holds together as a group(except for Anegada). The islands of the Virgin IslandsBank, which excludes St Croix, were once part of a largervolcanic island that included Puerto Rico. They are oldislands, about 105 Ma (Donnelly, 1988) that separatedwhen the sea level rose after the last Pleistocene glaciation,some 18 Ma ago. They are small, low-mountain islands(to 521 m) with hot climates ranging from moist to dryconditions (Ewel & Whitmore, 1973). Similarly, the Baha-mas belong to a single geological platform, part of theNorth American plate. They are low-lying, sedimentarylimestone areas. Many of the Bahamas were interconnectedin the past (Williams, 1989). Regarding the CaymanIslands, which are located south of Cuba on the Cayman

Table 2 Primary data sources for the orchid species presence inthe studied Caribbean regions

Area Data source

Bahama Archipelago Sauleda & Adams (1982), Sauleda (1992)

Cayman Islands Proctor (1984, 1996)

Florida Wunderlin et al. (1996)

Greater Antilles Ackerman, J. D. (unpublished data)

Guianas Boggan et al. (1997)

Isla de la Juventud Jennings (1917), Sauget & Barbier (1946)

Lesser Antilles Garay & Sweet (1974), Nir, M.

(unpublished)

Margarita Hoyos (1985)

Southern Dutch Antilles van Proosdij, A. (unpublished)

Trinidad and Tobago Garay & Sweet (1974); Kenny (1988)

Yucatan Carnevali, G. (unpublished)

Table 1 Antillean islands and neighbouring continental areasincluded in the study. The Bahamas islands are combined followingCorrell & Correll (1982); names in italics are the ones used in ®guresand appendix

Greater Antilles Virgin IslandsCuba Anegada

Hispaniola

(Dominican Republic & Haiti)

Culebra

Jamaica St John

Puerto Rico St Thomas

Isla de la Juventud St Croix

Mona Tortola

Cayman Brac Vieques

Grand Cayman Virgin Gorda

Little Cayman

Bahama Archipelago

Lesser Antilles Abacos ± Grand Bahama

Antigua Andros ± Biminis

Barbados Cat

Barbuda Crooked ± Mayaguana

Dominica Exumas

Grenada Inaguas

Guadeloupe Long ± Ragged Island Range

Martinique New Providence ± Eleutheras

Montserrat San Salvador ± Rum Cay

Nevis Turks and Caicos

Saba

St BartheÂlemy

St Eustatius Trinidad bank

St Kitts Margarita

St Lucia Tobago

St Martin Trinidad

St Vincent

Continental regions Southern Dutch Antilles

Florida (North America) Aruba

Yucatan (Central America) Bonaire

Guianas: Guyana, Surinam and

French Guyana (South America)

CuracËao


Biogeography of the Antilles based on a parsimony analysis of orchid distributions 777

Ridge, are small, low-lying islands with limestone sedimen-tary substrate to 36 Ma; no connection between theseislands and other lands occurred in the past (Proctor,1984). Lastly, Aruba, Bonaire and CuracËao, with volcanicand sedimentary substrates, are geologically related andpart of the South Caribbean Island Chain (Bellizzia &Dengo, 1990). They are also small, low-elevation islands(to 372 m) with volcanic and sedimentary substrates, andhave dry climates. In addition to their common origin, eachone of these island aggregates has a common geomorphol-ogy and physiography with relatively homogeneous ecolog-ical conditions and low species richness. The anomalous¯oristic connection of St Croix with the Virgin Islandsgroup is understandable if we look at the physiographyrather than the geology. St Croix has an independent originfrom the Virgin Islands Bank; nevertheless, they arephysiographically similar.

The second pattern is represented by areas differing ingeological origin and that are separated by wide geograph-ical distances among themselves. One example is representedby the Bahamas/Isla de la Juventud/Mona/Anegada group.As mentioned above, the Bahama Archipelago is composedof low-lying, sedimentary limestone islands located on theNorth American plate. The Isla de la Juventud, which lies onthe western part of the Cuban Bank, is almost entirely low-lying, part calcareous and part metamorphic (CRAC, 1978).Mona is a small, ¯at, calcareous island between two of theGreater Antilles (Puerto Rico and Hispaniola). Lastly,Anegada is part of the Virgin Islands Bank; however, unlikethe other Virgin Islands, it is ¯at, low-lying and calcareous(D'Arcy, 1971). In spite of the separate geologicalorigin, these islands have a similar geomorphology andphysiography. The other example of the second pattern isthe Greater Antilles/Trinidad/larger Lesser Antilles/smaller

Figure 1 Consensus trees of ¯oristic af®nities of the Antilles and some continental areas based on a parsimony analysis of 356 sharedorchid species distributions. Consensus trees were obtained from 216 most parsimonious trees built by a general heuristic search (treelength � 903, CI � 0.395, RI � 0.626, RC � 0.247). Majority Rule and Adams trees are almost identical, except for placement of Anegada andbasal areas within the Bahamas branch. Many basal areas of different groups in Majority Rule and Adams trees appear collapsed in the Stricttree. Numbers on branches of Majority Rule tree indicate the percentage of the most parsimonious trees that support the grouping; +̀' afternumbers indicates the same value for subsequent internodes. Numbers in parentheses after the area-names in the Majority Rule tree are numbersof shared species. Relationships among areas are interpreted through branch connections and not in terms of vicinity in tree branches.Polytomies indicate unresolved af®nities among areas. Names in bold call the attention of some area af®nities.



Eastern Antilles/Virgin Islands/Margarita±Tobago/othersmaller Lesser Antilles group. The origin of the GreaterAntilles, whether continental or oceanic, is under debate(Rosen, 1985; Iturralde-Vinent & MacPhee, 1999); how-ever, they are quite old, ranging from 105 to 80 Ma(Donnelly, 1988), or perhaps 150 Ma (Lewis & Draper,1990). The Lesser Antilles ®rst emerged as an oceanic islandarc about 43 Ma, and later expanded 11 Ma to form thenorth-western Lesser Antilles, from Dominica to Saba(Coney, 1982; Maury et al., 1990). Finally, the islands ofTrinidad, Margarita and Tobago, belong to the CaribbeanMountain System that is part of the continental SouthAmerican plate (Bellizzia & Dengo, 1990). There has beenno connection between these islands and the Lesser Antilles(Donnelly, 1988). Trinidad lost connection with SouthAmerica some 5 Ma (Persad, 1985). Notwithstanding thedifferences in geology and age, most of the islands of thisgrouping are volcanic, mountainous, have a variety ofclimates and natural communities, and complex speciesassemblages.

Orchid distribution patterns also seem to be in¯uenced bytheir high vagility. On a regional scale (the Antilles) cohesiveclusters of islands spread over hundreds to thousands ofkilometres (e.g. Bahamas±Anegada or the Greater Antilles±Eastern Antilles) re¯ect the high potential for orchid disper-sal, a capacity that may override the effects of geologicalhistory in de®ning the af®nities among those islands.

In general, distribution patterns of orchids are explainableaccording to the physical features of the islands. The similargeomorphology and/or physiography of these islands pre-sumably results in similar ecological conditions that conse-quently produce similar assemblages of species. However,geology clearly plays an important role in de®ning biogeo-graphical af®nities at a more restricted geographical scale.We do not know to what extent the biogeographical patterns

are affected by other factors such as prevailing winds andhurricane tracks.

Analysis of areas using different island data sets

Different analyses using subsets of areas (using the informativespecies for each data set) show patterns compatible with thegeneral analysis of all areas. Nevertheless, they do addinformation about the relationships among some groupingsand areas. The following observations are based mainly on theMajority Rule consensus trees (trees not presented).

1 When we analyse only the islands, excluding the contin-ental areas of Florida, the Guianas and the Yucatan, theBahamas split into two branches. The north-westernareas (Andros/Abacos/Providence) form a group with theIsla de la Juventud as basal. The Cayman Islands becomea sister clade of this Bahama group. Meanwhile, the restof the Bahama islands form another group with Monastill as basal.

2 If we eliminate the Isla de la Juventud (and the contin-ental areas), the Bahamas are not divided into two groupsas before and form a monocladic group. Mona andAnegada remain as basal areas.

3 When we eliminate the Bahamas (and the continentalareas), the Isla de la Juventud interestingly forms a pairwith Margarita, and together a sister group of theCayman Islands, all of them belong to an unresolvedclade. Meanwhile, Mona is left unresolved.

4 An analysis of the north-western areas, that is, theYucatan and Florida, the Bahamas, the Greater Antilles,the Isla de la Juventud, the Cayman Islands, Mona, andthe Virgin Islands, shows the Isla de la Juventud in an inter-mediate position between the Yucatan/Florida/GreaterAntilles and the Bahamas/Mona/Virgin Islands. TheCayman Islands remain monocladic and again unresolved.

Figure 2 Biogeographical patterns of af®nityof the Antilles based on a parsimoniousanalysis of 356 shared orchid species distri-butions. See Fig. 1 for detailed patterns ofaf®nity and single-island names. Dotted are-as: continental areas; short-dashed lines:island groups; long-dashed lines: relation-ships with continental areas; question marks:unresolved group or controversial af®nity;arrow: presumed af®liation. Island or group-ing names: A � Anegada, B � Barbados,C � Cuba, CI � Cayman Islands, H � His-paniola, IJ � Isla de la Juventud, J �Jamaica, LLA � Larger Lesser Antilles,M �Mona, MT �Margarita and Tobago,NBA � North-western Bahama Archipelago,PR � Puerto Rico, SBA � South-easternBahama Archipelago, SDA � SouthernDutch Antilles, SLA � Smaller Lesser Antil-les, T � Trinidad, VI � Virgin Islands.



5 We also evaluated distributions of just the GreaterAntillean area, which includes the Isla de la Juventud,the Cayman Islands, the Virgin Islands and Mona. Thisanalysis reveals the Virgin Islands and Mona as basal ofthe Greater Antilles in the order mentioned. Meanwhilethe Isla de la Juventud joins the Cayman Islands formingan unresolved clade.

6 An analysis of our Eastern Antilles, from the VirginIslands to Margarita, links Trinidad as basal to the largerLesser Antilles, particularly as the sister area of neigh-bouring Grenada. Tobago and Margarita appear as sisterareas inserted within the Eastern Antilles group. TheVirgin Islands remain monocladic.

7 After merging islands belonging to banks (de®ned by the200 m bathimetric line: Trinidad+Tobago+Margarita,Virgin Islands except St Croix, Antigua+Barbuda,St Kitts+Nevis+ St BartheÂlemy), the Trinidad bankremains as a sister and basal group of the Greater Antilles(as Trinidad alone).

Points (1), (3) and (5) denote a close af®nity among theIsla de la Juventud, the Cayman Islands, and the Bahamas.

Relationships among the groupings using exclusive

and endemic species

The parsimony analysis produced consistent assemblages ofareas. Nevertheless, the relationships among some of thegroupings are not resolved. We attempt to understand theseunde®ned af®nities by subjectively examining the number ofendemic species, the number of exclusive species (single-site,non-endemic species), and the presence of species of restric-ted distribution of every island or group. Are all theunresolved clusters of islands in the consensus trees, inde-pendent biogeographical provinces?

The Eastern Antilles, from the Virgin Islands to Margaritaconstitute a distinct branch with several minor subgroups.First, the larger Lesser Antilles from Guadeloupe to St Lucia isthe most de®ned group, with seven endemic species [Ellean-thus dussii Cogniaux, Epidendrum discoidale Lindley, Max-illaria guadalupensis Cogniaux, Pleurothallis dussiiCogniaux, P. ophioglossoides (Jacquin) Garay, Pseudocen-trum guadalupense Cogniaux, Stelis dussii Cogniaux]. Sec-ondly, the smaller Lesser Antilles (islands north ofGuadeloupe from Montserrat to St Martin) is a `group' thatsplits along the Eastern Antilles branch (Figs 1 and 2). Someof these islands share two endemic species with the largerLesser Antilles group (Epidendrum difforme Jacquin andEpidendrum patens Swartz). We recognize that this smallerLesser Antilles `group' is not cast in stone because basalspecies-poor areas are prone to changing af®nities. Thirdly,the Virgin Islands, except Anegada, constitute a monocladicbut weakly differentiated (no endemic species) group which is®rmly allied with the Lesser Antilles. Nevertheless, the VirginIslands have two species that they share exclusively withneighbouring Puerto Rico [Psychilis macconnelliae Sauleda,Tolumnia prionochila (KraÈnzlin) Braem]. This connectionwas insuf®cient to alter the ¯oristic af®nities of the Virgin

Islands with the Lesser Antilles in our analyses. Lastly,Tobago and Margarita form a pair within the Eastern Antillesbranch. The fact that these island groups appear as basal andpectinate with respect to the Greater Antilles, indicates a closeaf®nity among the islands of our Island Arc group.

The Cayman Islands are always monocladic and usuallyoccur as an independent clade. They have three endemicspecies [Dendrophylax fawcettii Rolfe, Encyclia kingsii(C. D. Adams) Nir, Myrmecophila albopurpurea (Strachanex Fawcett) Ackerman; recent morphological and moleculardata fail to distinguish T. caymanense (Moir) Braem fromT. variegata (Swartz) Braem]. Among the species of restricteddistribution two are shared with the Greater Antilles, Floridaand the Yucatan [Harrisella porrecta (Reichenbach f.)Fawcett & Rendle, Tropidia polystachya (Swartz) Ames],other two are shared only with the Greater Antilles[T. calochila (Cogniaux) Braem, T. variegata], and anotheris shared only with Cuba [Myrmecophilla thomsoniana(Reichenbach f.) Rolfe]. These species in common suggest aclose af®nity among the Cayman Islands, the GreaterAntilles and the peninsulas of Yucatan and Florida.

The Bahamas are also an independent clade in the generalanalysis. They have ®ve endemic species [Encyclia caicensis(Sauleda & Adams), E. fehlingii (Sauleda) Sauleda &Adams, E. gracilis (Lindley) Schlechter, E. inaguensis Nashex Britton & Millspaugh, T. sasseri (Moir) Braem]. Of thespecies with the most restricted distribution, there is onlyone exclusively shared with Florida [T. bahamense (Nash exBritton & Millspaugh) Braem] while there are seven sharedexclusively with the Greater Antilles [Broughtonia lindeniiLindley, Domingoa haematochila (Reichenbach f.) Carabia,Encyclia fucata (Lindley) Britton & Millspaugh, E. plicata(Lindley) Schlechter, Tetramicra urbaniana Cogniaux,T. gauntletii (Withner & Jesup) Braem, T. lucayana (Nashex Britton & Millspaugh) Braem]. Thus, the relationshipsamong the Bahamas and the Greater Antilles seems to beconsiderable.

Lastly, Aruba, Bonaire and CuracËao form another inde-pendent group. These islands have only three speciesreported, none of which is endemic. One occurs on differentcontinental and insular areas [Polystachya foliosa (Hooker)Reichenbach f.], one held in common with Central andSouth America [Brassavola nodosa (Linnaeus) Lindley], andone shared only with Venezuela [Myrmecophila humboldtii(Reichenbach f.) Rolfe]. Thus, the ABC islands show nospecial af®nity to the rest of the Antilles.

Relationships with continental areas

The inclusion of the Guianas as a South American contin-ental area was thought as a heuristic exercise. First, theGuianas is a politically de®ned area rather than a biogeo-graphical region. Secondly, the Guianas is likely outside theCaribbean ¯oristic region and part of the Amazonian region(Gentry, 1982). If we were to include other South Americanregions in this study, no doubt the Guianas would shiftaf®nities dramatically. The Guianas form a sister group tothe Greater Antilles, which is indicative of the strong



af®nities among insular and continental orchid ¯oras (as hasbeen put forth for the ¯ora in general by Samek, 1988;Borhidi, 1996).

The Yucatan and Florida form a sister pair of the north-western Bahamas/Isla de la Juventud group. All these aresedimentary, calcareous, low-lying and ¯at areas. Samek(1988) and Borhidi (1996) de®ned South Florida togetherwith the Bahamas as one province. On the other hand,Gentry (1982) excluded Florida (except for the Keys) fromhis Caribbean (insular) region. We consider Florida andthe Yucatan as provinces not belonging to the Antilleansubregion, which are supposedly part of the Continentalsubregion of the Caribbean.

To de®ne more clearly the relationships among the insularAntilles and their neighbouring continental areas, morethorough studies are necessary. Also, the inclusion of othercontinental areas, working as distant sister areas in theanalyses, would be useful. For example, a sister continentalarea for the Yucatan biogeographical province (DuraÂn et al.,1998) could be a province in central-western Mexico. Bothoutside provinces belong to the same ¯oristic Mexican-Central American Caribbean subregion (sensu Samek, 1988).

Islands with unresolved af®nity

The unresolved islands or island groupings found in someconsensus trees could be explained in a number of ways.First, it is possible that an incomplete record of the speciesfor these islands could be hampering the resolution of theiraf®nities. As we improved the species record for the smallerLesser Antillean islands they became progressively resolved.We expect this trend to continue. Alternatively, the recordmay be complete and these unresolved areas could be theresult of: (1) a lower number of species which would preventtheir af®liation with other groups, (2) an equally strongaf®nity with more than one of the other groups, and (3) anon-hierarchical structure in the data (Ronquist, 1997;Glasby & Alvarez, 1999). One way an unresolved area orgroup could be considered distinctive would be if it had awell-differentiated ¯ora (i.e. endemic or exclusive species). Itis likely a combination of these factors that play a role inunresolved cases.

PAD, species richness and species±area curves

The position of areas in tree branches is in¯uenced by speciesnumbers. This pattern is not evident for the Antilleanarchipelago as a whole and only becomes obvious within themajor island assemblages. As we move from the tip to thebase of the tree branches, species numbers tend to diminish.This apparent numerical artefact has been stated as one ofthe main problems resulting from the use of areas withhighly dissimilar number of species, and a comparablenumber of taxa per site has been recommended as a pre-requisite for PAE analyses (Rosen, 1988, 1992; Rosen &Smith, 1988; Vargas, 1991; Conran, 1995). What our studyshows is that even with very few species the af®liation of anisland to a speci®c group occurs when the taxa permit it.

Every main cluster has its own species-poor islands. Weinterpret the basal position of an area as indicative of¯oristic subordination to the distal areas in the group.Otherwise, when the few species of an area do not de®ne itsaf®liation, such an area turns out to be unresolved orunstable, being located at the base of the whole tree or at thebase of different branches, respectively. In other words,either a high or a low number of species can de®nemembership patterns among areas. Limiting the analysis toareas with a similar number of species would unnecessarilyrestrict the use of this methodology for the Caribbean andmany other regions.

Differences in species numbers within our island groupscan be converted into species±area curves. We have foundthat the geologically and physiographically diverse montaneislands have a different species±area curve than that for low-lying islands, which are basically calcareous and physio-graphically homogeneous (Ackerman et al., submitted). Thecalcareous group, comprised of nineteen islands (includingthe Cayman Islands), has a z-value of 0.25 (however, r � 0.4,P � 0.07), a typical value for islands (Rosenzweig, 1995).Meanwhile, the montane island group, composed of twenty-six islands (our island arc group), has a z-value of 0.51(r � 0.89, P � 0.0001). This is close to an interarchipelagicvalue found by Adler (1992, cited by Rosenzweig, 1995).Moreover, if a species±area curve for this montane group iscalculated with only the endemic species (using only nineislands that have single-island endemics), z � 0.68 (r � 0.88,P � 0.0019) suggesting that habitat diversity is important forautochthonous speciation. The two z-values for the montanegroup seem to indicate that the Island Arc group behaves as acluster of provinces (Rosenzweig, 1995) as has beenproposed by Borhidi & MunÄ iz (1986). In contrast, theendemic species±area curve for the relatively homogeneouscalcareous group was not run because there are only threeislands (island banks indeed) with endemic species.

Phytogeographical regionalization of the Antilles

The Antilles is among the principal phytogeographicalregions of the world (Gentry, 1982). It has been includedwithin the Caribbean region that extends to continentallands from Mexico to Venezuela (Samek, 1988; Borhidi,1996). Our orchid data seem to support this view. Thephytogeographic sectorization of Samek (1988) and Borhidi(1996) consider every one of the Greater Antilles asprovinces. Major geographical aggregations of islands (i.e.the Bahamas, the Lesser Antilles) are also de®ned asprovinces, while minor groups of islands (e.g. the VirginIslands, the Cayman Islands) are integrated with some of thebig islands. Apart from the recognition of these kind ofdivisions, we also examined af®nities among and withinthem in a tree-like hierarchical structure (Fig. 2).

1 Every one of the major islands of the Greater Antilles(i.e. Cuba, Hispaniola, Jamaica and Puerto Rico) issuf®ciently differentiated to be recognized as a separateprovince of the Antilles.



2 The Lesser Antilles were taken as a single phytogeo-graphic province by Samek (1988) and Borhidi (1996).Their orchid ¯ora is suf®ciently distinctive, supportingthis view. Our analysis places them as part of the EasternAntilles group and at the same time subordinate to theGreater Antillean main group.

3 The Virgin Islands were integrated as part of Puerto Ricoby Samek (1988) and Borhidi (1996). However, our studyplaces them rather close to the Lesser Antilles, andaltogether, as subordinate of the Greater Antilles.

4 The Cayman Islands were considered part of Jamaica bySamek (1988) but as part of Cuba according to Borhidi(1996). Our analysis suggests that these islands havecomplex af®nities with the Greater Antilles, the Yucatanand Florida continental areas, and also with the Bahamas.

5 The Bahama archipelago was one of the Caribbeanprovinces distinguished by Samek (1988), Borhidi (1996),and Gentry (1982), and this is also supported by ourorchid data. Moreover, our study relates the Isla de laJuventud, Mona and Anegada (geographically and geo-logically part of the Greater Antilles) as subordinates ofthe Bahamian Province. The Isla de la Juventud shows astrong af®nity with the north-western Bahamas, whileMona and Anegada are small islands with close af®nity tothe rest of the Bahamas.

6 Trinidad, Tobago and Margarita were considered part ofthe Venezuela±Colombia Caribbean subregion by Samek(1988) and Borhidi (1996), while no clear af®liation forTrinidad and Tobago can be obtained from Gentry(1982). Our data support a closer af®nity of Trinidadwith the Greater Antilles instead of the neighbouringLesser Antilles. Based on its differentiated orchid ¯ora(approximately thirteen endemic species) we considerTrinidad as a ¯oristic province. On the other hand,Margarita and Tobago are linked with the Lesser Antilles±Virgin Islands group rather than with Trinidad. There aretwo potential problems with these islands. First, weconsider that the species record for these islands is notwell updated, especially for Trinidad and Tobago whosemost recent complete orchid ¯ora was published bySchultes (1960). Secondly, the Venezuelan Guyana maybe a more appropriate neighbouring continental area,instead of our Guianas area. If this is true, the islands ofthe Trinidad bank may be more related to the continentthan to the rest of the Antillean islands, as suggested bybat distribution patterns (Trejo-Torres & Rivera, unpub-lished data) and by geological history.

7 Aruba, Bonaire and CuracËao seem to be independent ofthe Antillean subregion and apparently part of theContinental Caribbean subregion.

Perspectives of PAD

The pattern we found for orchids is not necessarily the samefor the general ¯ora or for the entire biota of the region. Weexpect that phytogeographic relationships revealed byorchids might be similar to that of other highly vagile groupsof organisms such as wind-dispersed plants, ¯ying insects,

¯ying birds and bats. In this case, area af®nities appear to bede®ned by physical factors, such as geomorphology, physi-ography, climate, geography and island area. At the sametime, it is plausible to predict that less vagile organisms, suchas non-¯ying vertebrates and freshwater ®shes, will de®nedifferent patterns of area relationships, perhaps more in¯u-enced by geological and geographical history of areas.

The PAE/PAD is an alternative to the use of multivariatephenetic methods for classi®cation of biotas (Rosen, 1988;Vargas, 1991) and for comparing species assemblages. Infact, giving a static interpretation of area relationships, thisis a method equivalent to multivariate ones (Rosen, 1988,1992). Some important differences between PAE/PAD andmultivariate methods are: (1) The PAE/PAD uses parsimonyalgorithms instead of similarity indexes. This means that thehierarchy selected in this manner will be the best supportedby evidence, for it maximizes congruence between data andhierarchical patterns (Brady, 1994). (2) The PAE/PAD usescharacters (taxa) selected a priori based on their informat-iveness, that is, species found in all sites and single-sitespecies are eliminated as they give no information on theaf®nities among areas (3). The PAE/PAD provides a moreuni®ed methodology as multivariate methods use manydifferent similarity indices, which can also yield differentresults.

The PAE/PAD can be seen as a complementary method.While many community analyses are intended to describequantitative parameters, PAE/PAD is directed to describecomposition patterns, based on the identity of species(Worthen, 1996). Also, results of PAE/PAD in the form oftrees with a hierarchical structure could be a graphiccomplement to other methods which produce results thatare single numerical values, such as indexes, scores orexponents (i.e. nestedness analysis, Patterson & Atmar,1986). The hierarchical nature of the results does notnecessarily mean that communities are organized that way.After all, af®nities among communities may have anunderlying reticulate structure (Ronquist, 1997). We viewPAE/PAD as only one way to represent structure in biogeo-graphical data.

The PAE/PAD has the potential as a method for under-standing ecological biogeographic problems. This is suppor-ted by the Caribbean orchids for which composition patternsstrikingly match physical and ecological factors. The use ofthe method as a complement to vicariance and other histor-ical biogeographic methods has been suggested (Cracraft,199144 ). Because of its phenetic nature, patterns foundthrough PAE/PAD could be re¯ecting ecological phenomena,rather than historical ones. Nevertheless, until now, it hasalways been used under the historical point of view. Some ofthe aspects pointed out as disadvantages to the historicalapproach, do not exist in the ecological context. Forexample, ¯oristic af®nities, biodiversity comparisons andcomposition patterns can be seen as phenetic phenomena.

For the demarcation of biogeographical units PAE/PADcould be employed before searching for speci®c patterns indistributional data sets. This could avoid distortions frompossible structure in the data caused by mixing areas



belonging to different biotas. Within the same context,Morrone & Crisci (1995) suggested the use of panbiogeo-graphic methods as an initial step to do other historicalbiogeographic analyses.

The PAE/PAD could prioritize conservation efforts at theregional level (Posadas, 1996) by identifying areas or islandsof high diversity. We have shown that such use can behelpful for the Antilles, one of the world's biodiversityhotspots (Myers et al., 2000). However, conservationefforts often focus on unique species, particularly endemics(island endemics in our case), but PAD uses only sharedspecies. The exclusive use of PAD in formulating conserva-tion strategies can thus be misleading. This is especially truefor species-poor islands that appear subordinated to species-rich islands. Impoverished islands may thus be falselyinterpreted as irrelevant for biodiversity conservation. Forexample, the tiny island of Mona lies subordinate to otherislands of our calcareous group yet it is certainly worthy ofconservation efforts because Mona has its own set ofendemic plants and animals (including an orchid, Psychilismonensis Sauleda).

The PAE/PAD do have problems that need to be resolvedand these are the interpretation of: (1) the unresolved statusamong the main groupings of areas, (2) the area relation-ships when using islands as well as continental regions, (3)the potential use of branch-lengths in the trees. Specialattention should also be given when selecting data sets fororganisms with diverse ecological characteristics (e.g. birds,which could belong to different guilds: residents vs. migra-tory, volant vs. non-volant).

CONCLUSIONS

According to our orchid data, the Antillean islands consti-tute a phytogeographical subregion of the Caribbean. TheLesser Antilles and the Bahamas are well-de®ned provinces.Meanwhile, the Virgin Islands and the Cayman Islands arepoorly differentiated aggregates, perhaps to be considered asincipient provinces. All of them are subordinated to the mainGreater Antillean group.

Within islands groupings detected by PAD ordering ofareas is highly in¯uenced by species numbers. The species-rich islands form cohesive groupings to which other species-poor islands unite as subordinates. These subordinate islandsare liable to changing af®nities.

Orchid species show three core zones of high diversityin the Antilles: the Greater Antilles (Hispaniola, Cuba,Jamaica, Puerto Rico); the larger Lesser Antilles (Guade-loupe, Dominica, Martinique, St Vincent, Grenada,St Lucia); and the north-western Bahamas (Andros, Abacos,Grand Bahama, New Providence, Eleuthera).

The patterns we found for orchid distributions in theAntilles are in¯uenced primarily by physical characteristicsof areas. That is, areas with common ecological featureseither because of similar physiographical conditions orcommon geomorphology form tight clusters, despite theirdifferences in geology or the long distances among them.

The highly compatible patterns obtained from differentisland sets and from different consensus trees indicate thatrelationships of areas obtained are well supported by thedata. In other words, the groups identi®ed are not arti®cial,but rather the result of their orchid-species af®nities.

Whereas PAE/PAD is an appropriate methodology for thestudy of geographical regions based on shared species,single-site species (the majority of which are endemics) areindicative of ®ner degrees of biogeographical differentiation.We suggest using both data sets to analyse biogeographicalaf®nities, for it integrates differences and similarities. Thisapproach provides a more thorough use of ¯oristic infor-mation for the analysis of biogeographical relationships.

The parsimony methodology seems to be appropriate forthe analysis of areas, or any other natural unit, with verydissimilar number of species. The low number of speciesfound in a unit can de®ne its af®liation or can produce anunresolved ¯oristic af®nity. The lack of af®nity of an area toany group may be either real or an artefact caused by anincomplete species record.

The PAE/PAD has great potential as an alternative,complementary or ®rst-step method to other multivariateand phylogenetic methodologies in biogeography.

ACKNOWLEDGMENTS

The study was supported by NSF grants to JDA (DEB-9505459, HRD-9353549, HRD-9628475). Arkelio Alicea,Kary GarcõÂa, Eileen Bravo, and Yanet Crespo were studentsparticipating in the study under Alliance for MinoritiesParticipation Program at UPR. We thank Steve Rehnerlogistic assistance in data analysis. We are grateful to BrianRosen, David Lees, Susan AragoÂn, Marta DõÂaz, and JuanJ. Morrone who had made relevant comments on themanuscript. Timothy Johnston assisted editing the manu-script and Celene Espadas, Guillermo Bianchi, and AnaPorzecanski shared important references. GermaÂn Carnevali,Andre van Proosdij, Mark Nir and Hagen Stenzel providedunpublished data.

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BIOSKETCHES

J. Carlos Trejo-Torres is a graduate student at the University of Puerto Rico-RõÂo Piedras working under the guidance of James D.Ackerman. He is studying composition patterns in the limestone forest of the Greater Antilles. He is also working on thebiogeography of Caribbean bats. He worked on the community description, ¯oristics and taxonomy of the Yucatan forests andwetlands, especially the endemic ¯ora.

James D. Ackerman is a biologist at the University of Puerto Rico-RõÂo Piedras. He has interests in plant ecology, systematics andevolution, particularly of orchids. He has published numerous papers on the evolution of deception pollination and the processesinvolved in the diversi®cation of the Orchidaceae. Among his current projects is the Orchid treatment for the Flora of the GreaterAntilles.



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00

00

00

00

00

00

0

Ell

eanth

us

longi

bra

ctea

tus

00

00

10

00

00

00

00

00

00

00

00

00

00

00

00

00

10

00

10

00

00

00

00

00

00

00

Elt

rople

ctri

sca

lcar

ata

10

11

11

10

01

10

00

00

00

01

11

00

00

00

00

00

00

00

10

00

00

00

00

00

00

00

Ency

clia

acuti

foli

a0

00

00

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Ency

clia

angu

stif

oli

a0

00

00

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Ency

clia

bre

vifo

lia

00

00

00

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Ency

clia

cord

iger

a1

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

01

00

00

00

00

00

0

Ency

clia

fehli

ngi

i0

00

00

00

00

00

00

00

00

00

11

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Ency

clia

fuca

ta0

00

00

01

10

00

00

00

00

00

01

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Ency

clia

grac

ilis

00

00

00

00

00

00

11

11

01

11

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Ency

clia

grav

ida

00

01

11

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Ency

clia

inag

uen

sis

00

00

00

00

00

00

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Ency

clia

isoch

ila

00

01

11

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Ency

clia

kin

gsii

00

00

00

00

01

01

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Ency

clia

nem

atoca

ulo

n0

10

00

01

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0



Ap

pen

dix

1co

nti

nued

Spec

ies

nam

e/are

anam

e

Guianas

Yucatan

Florida

PuertoRico

Jamaica

Hispaniola

Cuba

IslaJuventud

Mona

GrandCayman

LittleCayman

CaymanBrac

Turks

Inaguas

Crooked

Long

Salvador

Exumas

Cat

Providence

Andros

Abacos

StMartin

Saba

StBarthÂ elemy

StKitts

Nevis

StEustatius

Antigua

Barbuda

Montserrat

Guadeloupe

Dominica

Martinique

StLucia

StVincent

Grenada

Barbados

Trinidad

Tobago

Margarita

Vieques

Culebra

Anegada

VirginGorda

Tortola

StThomas

StJohn

StCroix

Aruba

CuraË cao

Bonaire

Ency

clia

onci

dio

ides

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

10

00

00

00

00

00

00

Ency

clia

pli

cata

00

00

00

10

00

00

00

00

00

01

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Ency

clia

rufa

00

10

00

10

00

00

11

11

11

11

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Ency

clia

serr

ula

ta0

00

01

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Ency

clia

tam

pen

sis

00

10

00

10

00

00

00

00

00

00

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Epid

endru

mac

unae

00

10

00

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Epid

endru

man

ceps

10

11

11

11

00

00

00

00

00

00

00

11

01

01

10

01

11

11

11

00

10

00

10

11

00

00

Epid

endru

man

till

anum

00

01

01

00

00

00

00

00

00

00

00

01

00

00

00

01

11

01

00

00

00

00

00

00

00

00

Epid

endru

mbla

nch

eanum

00

00

11

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Epid

endru

mca

rpophoru

m1

00

11

11

00

00

00

00

00

00

00

00

00

00

00

00

11

01

11

01

00

00

00

00

00

00

0

Epid

endru

mci

liar

e1

10

11

11

00

00

00

00

00

00

00

01

11

01

11

00

11

11

11

11

01

11

01

11

11

00

0

Epid

endru

mco

mpre

ssum

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

10

00

00

00

00

00

00

Epid

endru

mco

ronat

um

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

10

00

00

00

00

00

00

Epid

endru

mcr

ista

tum

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

10

00

00

00

00

00

00

Epid

endru

mden

dro

bio

ides

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

01

11

01

00

00

00

00

00

00

00

00

Epid

endru

mdif

form

e0

00

00

00

00

00

00

00

00

00

00

00

00

10

10

00

11

11

11

00

00

00

00

00

00

00

0

Epid

endru

mdif

fusu

m0

10

01

01

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Epid

endru

m¯ori

den

se0

01

00

01

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Epid

endru

mhar

tii

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

11

00

00

00

00

00

00

Epid

endru

mib

aguen

se1

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

11

10

10

01

00

00

00

00

00

00

0

Epid

endru

mim

bri

catu

m0

00

01

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Epid

endru

mis

om

erum

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Epid

endru

mja

mai

cense

00

01

11

10

00

00

00

00

00

00

00

00

00

00

00

01

10

00

00

00

00

00

00

00

00

00

Epid

endru

mm

iser

rim

um

10

01

01

00

00

00

00

00

00

00

00

00

00

00

00

01

11

10

10

00

00

00

00

00

00

00

Epid

endru

mm

ute

lian

um

00

01

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Epid

endru

mneo

porp

ax0

00

00

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Epid

endru

mnoct

urn

um

11

11

11

11

01

00

00

00

00

01

11

00

00

00

00

01

11

01

10

10

10

00

00

00

00

00

Epid

endru

mnuta

ns

10

00

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Epid

endru

mori

enta

le0

00

01

01

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Epid

endru

mpal

lidi¯

oru

m0

00

00

01

10

00

00

00

00

00

00

00

00

00

00

01

01

00

11

00

00

00

00

00

00

00

0

Epid

endru

mpar

anae

nse

00

00

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Epid

endru

mpat

ens

00

00

00

00

00

00

00

00

00

00

00

00

01

10

00

01

01

00

00

00

00

00

00

00

00

00

Epid

endru

mpoly

gonat

um

00

00

01

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Epid

endru

mport

ori

censi

s0

00

10

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Epid

endru

mra

mosu

m1

00

11

11

00

00

00

00

00

00

00

00

00

00

00

00

11

11

11

01

00

00

00

00

00

00

0

Epid

endru

mre

pen

s0

00

01

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Epid

endru

mri

gidum

10

11

11

11

00

00

00

00

00

01

11

00

00

00

00

01

11

11

10

10

10

00

00

00

00

00

Epid

endru

mri

vula

re0

00

01

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Epid

endru

msc

apel

lige

rum

00

00

10

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Epid

endru

msc

hle

cter

ianum

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

10

00

00

00

00

00

00

Epid

endru

mse

cundum

00

01

11

10

00

00

00

00

00

00

00

00

00

00

10

01

11

11

10

00

00

00

00

00

00

00

Epid

endru

mst

robil

ifer

um

11

10

11

10

00

00

00

00

00

00

00

01

00

00

00

01

11

01

10

10

00

00

00

00

00

00

Epid

endru

mvi

nce

nti

num

10

01

11

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

10

00

00

00

00

00

00

Epid

endru

mw

righ

tii

00

00

01

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Epis

tephiu

mel

lipti

cum

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

10

00

00

00

00

00

00

Epis

tephiu

mpar

vi¯oru

m1

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

01

00

00

00

00

00

00

0

Ery

thro

des

hir

tell

a1

00

11

11

00

00

00

00

00

00

00

00

00

00

10

00

11

11

11

11

00

00

00

10

00

00

0



Ap

pen

dix

1co

nti

nued

Spec

ies

nam

e/are

anam

e

Guianas

Yucatan

Florida

PuertoRico

Jamaica

Hispaniola

Cuba

IslaJuventud

Mona

GrandCayman

LittleCayman

CaymanBrac

Turks

Inaguas

Crooked

Long

Salvador

Exumas

Cat

Providence

Andros

Abacos

StMartin

Saba

StBarthÂ elemy

StKitts

Nevis

StEustatius

Antigua

Barbuda

Montserrat

Guadeloupe

Dominica

Martinique

StLucia

StVincent

Grenada

Barbados

Trinidad

Tobago

Margarita

Vieques

Culebra

Anegada

VirginGorda

Tortola

StThomas

StJohn

StCroix

Aruba

CuraË cao

Bonaire

Ery

thro

des

maj

or

10

00

01

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Ery

thro

des

pla

nta

ginea

00

01

11

10

00

00

00

00

00

00

00

01

01

01

00

11

11

01

10

11

00

00

00

00

00

00

Eulo

phia

alta

11

11

11

10

00

00

00

00

00

00

01

00

00

00

10

01

11

11

10

11

00

00

01

00

00

00

Eulo

phia

ecri

stat

a0

01

00

01

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Eury

styl

esan

anas

soco

mos

00

01

11

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Fuer

tesi

ella

pte

rich

oid

es0

00

00

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Gal

eandra

bey

rich

ii0

01

11

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Gove

nia

utr

icula

ta0

01

11

11

00

00

00

00

00

00

01

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Hab

enar

iaal

ata

00

01

11

11

00

00

00

00

00

00

10

00

00

00

10

01

11

11

10

11

00

00

00

10

00

00

Hab

enar

iaam

al®ta

na

10

01

00

00

00

00

00

00

00

00

00

00

00

00

10

01

00

01

00

00

00

00

00

00

00

00

Hab

enar

iabic

orn

is0

00

00

01

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Hab

enar

iadis

tans

01

11

11

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Hab

enar

iaduss

ii0

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

10

00

10

00

00

00

00

00

00

00

0

Hab

enar

iaeu

stac

hya

00

01

11

10

00

00

00

00

00

00

00

00

00

00

00

01

00

00

00

00

00

00

00

00

00

00

Hab

enar

ia¯ori

bunda

10

10

11

11

00

00

00

00

00

00

11

00

00

00

00

01

00

00

00

00

00

00

00

00

00

00

Hab

enar

iale

pri

euri

i1

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

01

00

00

00

00

00

00

0

Hab

enar

iam

esodac

tyla

01

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

10

00

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Ap

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00

11

Schom

burg

kia

lyso

nsi

i0

00

01

01

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Sobra

lia

frag

rans

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Spir

anth

esto

rta

01

11

11

11

00

00

00

01

00

01

11

00

00

00

10

01

11

10

10

10

00

00

00

10

00

00

Stan

hopea

gran

di¯

ora

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

10

00

00

00

00

00

00

Stel

ism

usc

ifer

a1

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

01

00

00

00

00

00

00

0

Stel

isper

pusi

lli¯

ora

00

01

01

00

00

00

00

00

00

00

00

00

00

00

00

01

10

00

00

10

00

00

00

00

00

00

Stel

ispyg

mae

a1

00

10

10

00

00

00

00

00

00

00

00

00

00

00

00

11

10

10

00

00

00

00

00

00

00

0

Stel

issc

abri

da

00

00

10

00

00

00

00

00

00

00

00

00

00

10

00

01

11

11

00

00

00

00

00

00

00

00

Stel

istr

igoni¯

oru

m0

00

01

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Sten

iapal

lida

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

10

00

00

00

00

00

00

Sten

orr

hyn

chos

lance

ola

tum

10

11

11

11

00

00

00

00

00

01

11

00

10

00

01

00

00

00

01

10

00

00

00

10

00

00

Sten

orr

hyn

chos

spec

iosu

m0

00

01

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Sten

orr

hyn

chos

squam

ulo

sum

00

00

10

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Tet

ram

icra

bulb

osa

00

00

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00



Ap

pen

dix

1co

nti

nued

Spec

ies

nam

e/are

anam

e

Guianas

Yucatan

Florida

PuertoRico

Jamaica

Hispaniola

Cuba

IslaJuventud

Mona

GrandCayman

LittleCayman

CaymanBrac

Turks

Inaguas

Crooked

Long

Salvador

Exumas

Cat

Providence

Andros

Abacos

StMartin

Saba

StBarthÂ elemy

StKitts

Nevis

StEustatius

Antigua

Barbuda

Montserrat

Guadeloupe

Dominica

Martinique

StLucia

StVincent

Grenada

Barbados

Trinidad

Tobago

Margarita

Vieques

Culebra

Anegada

VirginGorda

Tortola

StThomas

StJohn

StCroix

Aruba

CuraË cao

Bonaire

Tet

ram

icra

canal

icula

ta0

00

10

10

00

00

00

00

00

00

00

00

01

00

11

10

10

10

00

11

00

00

11

01

11

00

0

Tet

ram

icra

eulo

phia

e0

00

00

11

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Tet

ram

icra

par

vi¯ora

00

00

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Tet

ram

icra

urb

ania

na

00

00

00

10

00

00

00

00

00

01

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Tolu

mnia

bah

amen

se0

01

00

00

00

00

00

00

00

00

01

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Tolu

mnia

calo

chil

a0

00

00

11

00

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Tolu

mnia

gauntl

etii

00

00

10

00

00

00

00

00

00

01

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Tolu

mnia

lyra

ta0

00

00

01

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Tolu

mnia

pri

onoch

ila

00

01

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

01

11

11

11

00

00

Tolu

mnia

sass

eri

00

00

00

00

00

00

00

00

00

00

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Tolu

mnia

uro

phyl

lum

00

00

01

00

00

00

00

00

00

00

00

00

10

11

11

11

10

00

00

00

00

00

00

00

00

00

Tolu

mnia

vari

egat

a0

00

10

11

01

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

11

01

11

11

00

0

Tri

chopil

iafr

agan

s0

00

00

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Tri

chosa

lpin

xci

liar

is1

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Tri

chosa

lpin

xdura

10

01

11

10

00

00

00

00

00

00

00

00

00

00

00

01

01

00

10

11

10

00

00

00

00

00

Tri

gonid

ium

eger

tonia

num

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Tri

phora

amaz

onic

a1

01

10

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

01

00

00

00

00

00

00

0

Tri

phora

genti

anoid

es0

11

01

11

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

0

Tri

phora

has

sler

iana

00

01

01

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Tri

phora

suri

nam

ensi

s1

00

11

10

00

00

00

00

00

00

00

00

00

00

00

00

11

00

00

01

00

00

00

00

00

00

0

Tri

phora

yuca

tanen

sis

01

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Tri

zeuxis

falc

ata

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

10

00

00

00

00

00

00

Tro

pid

iapoly

stac

hya

01

10

11

10

01

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Wull

schla

egel

lia

aphyl

la0

00

01

01

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

01

00

00

00

00

00

00

0

Wull

schla

egel

lia

calc

arat

a1

00

10

10

00

00

00

00

00

00

00

00

00

00

00

00

11

00

00

00

00

00

00

00

00

00

0

Xyl

obiu

mco

lley

i1

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

01

00

00

00

00

00

00

0

Xyl

obiu

mfo

veat

um

10

00

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

Xyl

obiu

mpal

mif

oli

um

10

00

11

10

00

00

00

00

00

00

00

00

00

00

00

00

10

01

10

10

00

00

00

00

00

00

Zootr

ophio

nat

ropurp

ure

um

00

00

11

10

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00

00



biogeography of the antilles based on a parsimony...

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