Journal of Biogeography, 28, 775±794

Biogeography of the Antilles based on a parsimony analysis of orchid distributions J. Carlos Trejo-Torres and James D. Ackerman Department of Biology, University of Puerto Rico-Rõo Piedras, PO BOX 23360, San Juan, Puerto Rico 00931-3360

Abstract Aim We obtain biogeographical patterns based on the distributions of shared orchid species of the Caribbean. These patterns are used to de®ne biogeographical zones. We then analyse the concordance between the distributional patterns with ecological and physical features of the islands. Location We use orchid species recorded on 49 islands of the Greater, Lesser, and southern 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 best arrangements of shared taxa among areas. The analysis uses 356 shared orchid species of the 863 species recorded for studied areas. The methodology has been used to infer historical relationships among areas but we interpret the results as static or ecological patterns of biogeographical af®nities. Results Two kinds of island groupings are revealed. (1) Groups with common geology and geomorphology: the Bahama Archipelago, the Virgin Islands, the Cayman Islands and the southern Dutch Antilles. (2) An aggregation of distant islands with a heterogeneous 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 ecological features either from similar geomorphology or common physiography. The strong af®nity among islands considerably distant among each other is explained by the high vagility of dust-seeded orchids. Then, ¯oristic af®nities seem determined by ecological characteristics of islands rather than by dispersal barriers. We predict that other plant groups with dust-like diaspores and animals with good vagility should show comparable biogeographic patterns. Parsimony analysis of distributions (PAD) is an alternative methodology to multivariate analysis to compare biotas, and a graphic complement to quantitative methods producing numerical values.

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

between Asia and Australia. The Antilles are also signi®cant INTRODUCTION because they represent one of the two connections, by way of The Antilles, or West Indies, have been frequently studied by an island chain, between two major biogeographical realms: biogeographers. These islands make up one of the largest the Neartic and the Neotropic. The majority of studies on tropical archipelagos in the world, second only to the islands Antillean biogeography concern the distributional patterns or the cladistic biogeography of animals, especially verte- brates and insects (e.g. Liebherr, 1988; Woods, 1989; Correspondence: J. Carlos Trejo-Torres, Department of Biology, University of Puerto Rico-RõÂo Piedras, PO BOX 23360, San Juan, Puerto Rico 00931-3360. Page & Lydeard, 1994; Hedges, 1996). Surprisingly, there 1 E-mail: [email protected] are only a few publications on the phytogeography of this

Ó 2001 Blackwell Science Ltd 776 J. C. Trejo-Torres and J. D. Ackerman

region (Howard, 1973; Samek, 1988; Adams, 1989; Lavin, more generic name: parsimony analysis of distributions 1993; Borhidi, 1996), although the ¯oristics of the area have (PAD). The method excludes single-site species (including been actively studied for decades (Zanoni, 1986; Liogier, single-site endemics), and takes into account shared species 1996). whether endemic or not to the Antilles or adjacent areas. The Caribbean region has been divided phytogeographi- We substitute the term distribution for endemicity to avoid cally in three subregions by Samek (1988). These main divi- confusion. sions are: (1) Mexico to Panama, (2) Colombia to Venezuela, Our goal here is to distinguish biogeographical patterns in and (3) an insular subregion including the Antilles proper. the Antilles based on one plant family, Orchidaceae. This The southern Antilles, from Aruba to Tobago are considered group of plants, of which there are approximately 700 part of the Colombian±Venezuelan subregion. Borhidi species in the Antilles, is relatively well known taxonomi- (1996) joins both continental subregions, from Mexico to cally and geographically because there are relatively recent Venezuela, into a single one. While these authors consider orchid treatments for the Cayman Islands (Proctor, 1984, south Florida as part of the Antillean subregion, Gentry 1996), Puerto Rico (Ackerman, 1995) and the Bahamas (1982) included only the Florida Keys in the Antillean region. (Sauleda & Adams, 1982; Sauleda, 1992). Furthermore, a Distinguishing biotic regions, or de®ning biological treatment for the Greater Antilles is underway (Ackerman, boundaries, has been among the major concerns of bioge- 1997; in press). The homogeneous wind-dispersal mechan- ographers. Traditionally, this has been performed with ism (except in bird-dispersed Vanilla) of the dust-seeds subjective methods (e.g. Gentry, 1982; Samek, 1988; among the species of this family, makes orchids a good focal Borhidi, 1996). However, more objective or analytical group for the study of distribution. Assuming that most approaches have been developed for the analysis of distri- orchids have the same dispersal capacity, their distributional butional data of organisms (e.g. Gauch, 1982; McCoy et al., patterns may be explained in terms of other ecological, 1986; Patterson & Atmar, 1986; Rosen & Smith, 1988; geographical and historical factors. Rosen, 1988; Vargas, 1991; Real et al., 1992; Scheiner, The questions we address are: (1) what are the phyto- 1992; Worthen, 1996; Puente et al., 1998). Here, we use an geographical relationships among the Antillean islands alternative method based on a parsimonious analysis of taxa based on shared orchid species? (2) are these relationships distributions (Rosen & Smith, 1988). This method repre- affected when neighbouring continental regions are consid- sents a direct way to search for the biogeographical af®nities ered? (3) what is the phytogeographical regionalization of among areas (Connor, 1988; Vargas, 1991), for the detec- the Antilles based on orchids, and (4) how do the distribu- tion of areas of endemism (Morrone, 1994a; Cardoso da tional patterns of orchids match with geography, geology, 2Silva & Oren, 1996; Bates et al., 1998), and for the physiography and ecology of the region? Apart from the delimitation of biological boundaries (Posadas, 1996; biogeographical analysis per se, we discuss the use of Posadas et al., 1997; Morrone, 1998). The parsimony parsimony analysis of naturally de®ned areas (i.e. islands) analysis presented here is a tool for searching the most with extremely dissimilar number of species, and the use of parsimonious arrangement of shared species among areas, as single-site species to look at ®ner degrees of biogeographical a means of revealing the biogeographical af®nities in a differentiation. hierarchical pattern (Rosen & Smith, 1988; Brady, 1994). The analysis presented here was originally called parsi- METHODS mony analysis of endemicity (PAE), and was suggested by 3Rosen (1985, cited in Rosen & Smith, 1988) and developed A total of ®fty-two areas (Table 1) were included in the by Rosen & Smith (1988). It was also independently study. Data on the distribution of species were taken from suggested by Legendre (1986) and Connor (1988). Since literature and from unpublished sources (Table 2). Synony- then it has been employed in the study of extant taxa of mies and valid species names were standardized mainly New Zealand (Craw, 1988), Australia (Cracraft, 1991), according to J.D. Ackerman (unpublished data). Southeast Africa (Morrone, 1994a), the Patagonia A presence/absence matrix of the 863 orchid species (Posadas, 1996), the Andes (Morrone, 1994b; Posadas reported for the studied areas was constructed in MacClade et al., 1997), the Amazonia (Cardoso da Silva & Oren, 3.01 (Maddison & Maddison, 1992). Using this program, 1996; Bates et al., 1998), the Austral region (Craw, 1989; areas were entered in the place of taxa, while taxa were Morrone, 1998; Glasby & Alvarez, 1999), Mexico (Luna entered in the place of characters. In the matrix, presence et al., 1999), and the entire world (Conran, 1995). The was indicated with a `1' and absence with a `0'. Once we units of comparison that have been used are sites, quad- constructed the matrix, we ran analyses of parsimony using rants or sections of regions, biogeographical areas, or PAUP 4 (Swofford, 2000). A hypothetical outgroup area natural geographical areas (e.g. islands, continents, ocean with all 0s (no species) were used in the analyses to root the basins). We use entire islands or groups of them, as the trees. General heuristic searches were carried out to look for units of study. While the method has been mainly used for the most parsimonious trees, which indicate the ¯oristic discovering the historical relationships among areas, we af®nities among studied areas. We obtained consensus trees give a static or non-historical interpretation to the patterns (Strict, Majority Rule and Adams) when more than one obtained (Rosen, 1992; Posadas, 1996; Posadas et al., equally parsimonious tree were found. A description and a 1997). Although the method has been called PAE, we use a discussion of the parsimonious analysis methodology are

Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794 Biogeography of the Antilles based on a parsimony analysis of orchid distributions 777

Table 1 Antillean islands and neighbouring continental areas information in terms of shared species af®nities. We only included in the study. The Bahamas islands are combined following used these single-site (island or area) species for speci®c Correll & Correll (1982); names in italics are the ones used in ®gures ¯oristic comparisons. Also, Vanilla species were not consid- and appendix ered at all because they are animal-dispersed, an anomaly in the orchid family. Greater Antilles Virgin Islands Area relationships derived from PAD may be altered Cuba Anegada Hispaniola Culebra depending on the inclusion or exclusion of different areas (Dominican Republic & Haiti) just as is the case when using different ingroup/outgroup Jamaica St John taxa in cladistic studies. To test the robustness of our data, Puerto Rico St Thomas we conducted independent analyses of different subsets of Isla de la Juventud St Croix Mona Tortola areas. Cayman Brac Vieques Grand Cayman Virgin Gorda Little Cayman RESULTS Bahama Archipelago An analysis of all areas using the 356 shared orchid species Lesser Antilles Abacos ± Grand Bahama Antigua Andros ± Biminis produced 216 equally parsimonious trees. The three con- Barbados Cat sensus trees obtained from them produced entirely congruent Barbuda Crooked ± Mayaguana groupings (Fig. 1). The Strict consensus tree shows the Dominica Exumas groups found in all the most parsimonious trees. The Grenada Inaguas Guadeloupe Long ± Ragged Island Range Majority Rule tree shows the groups found in more than Martinique New Providence ± Eleutheras 50% of those trees. The Adams tree gives the highest Montserrat San Salvador ± Rum Cay possible resolution for data distribution. Nevis Turks and Caicos Four main clusters of areas can be identi®ed in the Saba consensus trees (Fig. 1): (1) Greater Antilles/Guianas±Trini- St BartheÂlemy St Eustatius Trinidad bank dad/larger Lesser Antilles/smaller Lesser Antilles/Virgin St Kitts Margarita Islands/Margarita-Tobago/other smaller Lesser Antilles, St Lucia Tobago (2) Yucatan/Florida/Isla de la Juventud/Bahamas/Mona/ St Martin Trinidad Anegada, (3) Cayman Islands, and (4) Aruba/CuracËao/ St Vincent Bonaire. Although these are well-de®ned aggregations, the Continental regions Southern Dutch Antilles polytomic base of the trees means that relationships among Florida (North America) Aruba them are not resolved. The arrangement of areas within the Yucatan (Central America) Bonaire Guianas: Guyana, Surinam and CuracËao groupings is highly in¯uenced by species numbers. French Guyana (South America) DISCUSSION Groupings of areas vs. physical factors Table 2 Primary data sources for the orchid species presence in the studied Caribbean regions We found two kinds of grouping patterns in the Antilles based on orchid species distributions. One pattern aggre- Area Data source gates islands that belong to single geological units while the Bahama Archipelago Sauleda & Adams (1982), Sauleda (1992) other pattern groups areas with different geological histories Cayman Islands Proctor (1984, 1996) (Fig. 2). Florida Wunderlin et al. (1996) In the ®rst pattern, the groupings are easy to understand Greater Antilles Ackerman, J. D. (unpublished data) as they correspond to islands belonging to de®ned geolo- Guianas Boggan et al. (1997) Isla de la Juventud Jennings (1917), Sauget & Barbier (1946) gical units. The Virgin Islands holds together as a group Lesser Antilles Garay & Sweet (1974), Nir, M. (except for Anegada). The islands of the Virgin Islands (unpublished) Bank, which excludes St Croix, were once part of a larger Margarita Hoyos (1985) volcanic island that included Puerto Rico. They are old Southern Dutch Antilles van Proosdij, A. (unpublished) islands, about 105 Ma (Donnelly, 1988) that separated Trinidad and Tobago Garay & Sweet (1974); Kenny (1988) Yucatan Carnevali, G. (unpublished) when 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 dry conditions (Ewel & Whitmore, 1973). Similarly, the Baha- found in Rosen & Smith (1988), Vargas (1991) and Rosen mas belong to a single geological platform, part of the (1988, 1992). North American plate. They are low-lying, sedimentary Only the informative species were taken into account in limestone areas. Many of the Bahamas were interconnected the analyses (Appendix 1). The uninformative species are in the past (Williams, 1989). Regarding the Cayman those found in only one area; therefore, they give no Islands, which are located south of Cuba on the Cayman

Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794 778 J. C. Trejo-Torres and J. D. Ackerman

Figure 1 Consensus trees of ¯oristic af®nities of the Antilles and some continental areas based on a parsimony analysis of 356 shared orchid species distributions. Consensus trees were obtained from 216 most parsimonious trees built by a general heuristic search (tree length ˆ 903, CI ˆ 0.395, RI ˆ 0.626, RC ˆ 0.247). Majority Rule and Adams trees are almost identical, except for placement of Anegada and basal areas within the Bahamas branch. Many basal areas of different groups in Majority Rule and Adams trees appear collapsed in the Strict tree. Numbers on branches of Majority Rule tree indicate the percentage of the most parsimonious trees that support the grouping; `+' after numbers indicates the same value for subsequent internodes. Numbers in parentheses after the area-names in the Majority Rule tree are numbers of 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.

Ridge, are small, low-lying islands with limestone sedimen- The second pattern is represented by areas differing in tary substrate to 36 Ma; no connection between these geological origin and that are separated by wide geograph- islands and other lands occurred in the past (Proctor, ical distances among themselves. One example is represented 1984). Lastly, Aruba, Bonaire and CuracËao, with volcanic by the Bahamas/Isla de la Juventud/Mona/Anegada group. and sedimentary substrates, are geologically related and As mentioned above, the Bahama Archipelago is composed part of the South Caribbean Island Chain (Bellizzia & of low-lying, sedimentary limestone islands located on the Dengo, 1990). They are also small, low-elevation islands North American plate. The Isla de la Juventud, which lies on (to 372 m) with volcanic and sedimentary substrates, and the western part of the Cuban Bank, is almost entirely low- have dry climates. In addition to their common origin, each lying, part calcareous and part metamorphic (CRAC, 1978). one of these island aggregates has a common geomorphol- Mona is a small, ¯at, calcareous island between two of the ogy and physiography with relatively homogeneous ecolog- Greater Antilles (Puerto Rico and Hispaniola). Lastly, ical conditions and low species richness. The anomalous Anegada is part of the Virgin Islands Bank; however, unlike ¯oristic connection of St Croix with the Virgin Islands the other Virgin Islands, it is ¯at, low-lying and calcareous group is understandable if we look at the physiography (D'Arcy, 1971). In spite of the separate geological rather than the geology. St Croix has an independent origin origin, these islands have a similar geomorphology and from the Virgin Islands Bank; nevertheless, they are physiography. The other example of the second pattern is physiographically similar. the Greater Antilles/Trinidad/larger Lesser Antilles/smaller

Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794 Biogeography of the Antilles based on a parsimony analysis of orchid distributions 779

Figure 2 Biogeographical patterns of af®nity of the Antilles based on a parsimonious analysis of 356 shared orchid species distri- butions. See Fig. 1 for detailed patterns of af®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 ˆ Anegada, B ˆ Barbados, C ˆ Cuba, CI ˆ Cayman Islands, H ˆ His- paniola, IJ ˆ Isla de la Juventud, J ˆ Jamaica, LLA ˆ Larger Lesser Antilles, M ˆ Mona, MT ˆ Margarita and Tobago, NBA ˆ North-western Bahama Archipelago, PR ˆ Puerto Rico, SBA ˆ South-eastern Bahama Archipelago, SDA ˆ Southern Dutch Antilles, SLA ˆ Smaller Lesser Antil- les, T ˆ Trinidad, VI ˆ Virgin Islands.

Eastern Antilles/Virgin Islands/Margarita±Tobago/other are affected by other factors such as prevailing winds and smaller Lesser Antilles group. The origin of the Greater hurricane tracks. Antilles, whether continental or oceanic, is under debate (Rosen, 1985; Iturralde-Vinent & MacPhee, 1999); how- Analysis of areas using different island data sets ever, they are quite old, ranging from 105 to 80 Ma (Donnelly, 1988), or perhaps 150 Ma (Lewis & Draper, Different analyses using subsets of areas (using the informative 1990). The Lesser Antilles ®rst emerged as an oceanic island species for each data set) show patterns compatible with the arc about 43 Ma, and later expanded 11 Ma to form the general analysis of all areas. Nevertheless, they do add north-western Lesser Antilles, from Dominica to Saba information about the relationships among some groupings (Coney, 1982; Maury et al., 1990). Finally, the islands of and areas. The following observations are based mainly on the Trinidad, Margarita and Tobago, belong to the Caribbean Majority Rule consensus trees (trees not presented). Mountain System that is part of the continental South 1 When we analyse only the islands, excluding the contin- American plate (Bellizzia & Dengo, 1990). There has been ental areas of Florida, the Guianas and the Yucatan, the no connection between these islands and the Lesser Antilles Bahamas split into two branches. The north-western (Donnelly, 1988). Trinidad lost connection with South areas (Andros/Abacos/Providence) form a group with the America some 5 Ma (Persad, 1985). Notwithstanding the Isla de la Juventud as basal. The Cayman Islands become differences in geology and age, most of the islands of this a sister clade of this Bahama group. Meanwhile, the rest grouping are volcanic, mountainous, have a variety of of the Bahama islands form another group with Mona climates and natural communities, and complex species still as basal. assemblages. 2 If we eliminate the Isla de la Juventud (and the contin- Orchid distribution patterns also seem to be in¯uenced by ental areas), the Bahamas are not divided into two groups their high vagility. On a regional scale (the Antilles) cohesive as before and form a monocladic group. Mona and clusters of islands spread over hundreds to thousands of Anegada remain as basal areas. kilometres (e.g. Bahamas±Anegada or the Greater Antilles± 3 When we eliminate the Bahamas (and the continental Eastern Antilles) re¯ect the high potential for orchid disper- areas), the Isla de la Juventud interestingly forms a pair sal, a capacity that may override the effects of geological with Margarita, and together a sister group of the history in de®ning the af®nities among those islands. Cayman Islands, all of them belong to an unresolved In general, distribution patterns of orchids are explainable clade. Meanwhile, Mona is left unresolved. according to the physical features of the islands. The similar 4 An analysis of the north-western areas, that is, the geomorphology and/or physiography of these islands pre- Yucatan and Florida, the Bahamas, the Greater Antilles, sumably results in similar ecological conditions that conse- the Isla de la Juventud, the Cayman Islands, Mona, and quently produce similar assemblages of species. However, the Virgin Islands, shows the Isla de la Juventud in an inter- geology clearly plays an important role in de®ning biogeo- mediate position between the Yucatan/Florida/Greater graphical af®nities at a more restricted geographical scale. Antilles and the Bahamas/Mona/Virgin Islands. The We do not know to what extent the biogeographical patterns Cayman Islands remain monocladic and again unresolved.

Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794 780 J. C. Trejo-Torres and J. D. Ackerman

5 We also evaluated distributions of just the Greater Islands with the Lesser Antilles in our analyses. Lastly, Antillean area, which includes the Isla de la Juventud, Tobago and Margarita form a pair within the Eastern Antilles the Cayman Islands, the Virgin Islands and Mona. This branch. The fact that these island groups appear as basal and analysis reveals the Virgin Islands and Mona as basal of pectinate with respect to the Greater Antilles, indicates a close the Greater Antilles in the order mentioned. Meanwhile af®nity among the islands of our Island Arc group. the Isla de la Juventud joins the Cayman Islands forming The Cayman Islands are always monocladic and usually an unresolved clade. occur as an independent clade. They have three endemic 6 An analysis of our Eastern Antilles, from the Virgin species [Dendrophylax fawcettii Rolfe, Encyclia kingsii Islands to Margarita, links Trinidad as basal to the larger (C. D. Adams) Nir, Myrmecophila albopurpurea (Strachan Lesser Antilles, particularly as the sister area of neigh- ex Fawcett) Ackerman; recent morphological and molecular bouring Grenada. Tobago and Margarita appear as sister data fail to distinguish T. caymanense (Moir) Braem from areas inserted within the Eastern Antilles group. The T. variegata (Swartz) Braem]. Among the species of restricted Virgin Islands remain monocladic. distribution two are shared with the Greater Antilles, Florida 7 After merging islands belonging to banks (de®ned by the and the Yucatan [Harrisella porrecta (Reichenbach f.) 200 m bathimetric line: Trinidad + Tobago +Margarita, Fawcett & Rendle, Tropidia polystachya (Swartz) Ames], Virgin Islands except St Croix, Antigua + Barbuda, other two are shared only with the Greater Antilles St Kitts + Nevis + St BartheÂlemy), the Trinidad bank [T. calochila (Cogniaux) Braem, T. variegata], and another remains as a sister and basal group of the Greater Antilles is shared only with Cuba [Myrmecophilla thomsoniana (as Trinidad alone). (Reichenbach f.) Rolfe]. These species in common suggest a close af®nity among the Cayman Islands, the Greater Points (1), (3) and (5) denote a close af®nity among the Antilles and the peninsulas of Yucatan and Florida. Isla de la Juventud, the Cayman Islands, and the Bahamas. The Bahamas are also an independent clade in the general analysis. They have ®ve endemic species [Encyclia caicensis Relationships among the groupings using exclusive (Sauleda & Adams), E. fehlingii (Sauleda) Sauleda & and endemic species Adams, E. gracilis (Lindley) Schlechter, E. inaguensis Nash ex Britton & Millspaugh, T. sasseri (Moir) Braem]. Of the The parsimony analysis produced consistent assemblages of species with the most restricted distribution, there is only areas. Nevertheless, the relationships among some of the one exclusively shared with Florida [T. bahamense (Nash ex groupings are not resolved. We attempt to understand these Britton & Millspaugh) Braem] while there are seven shared unde®ned af®nities by subjectively examining the number of exclusively with the Greater Antilles [Broughtonia lindenii endemic species, the number of exclusive species (single-site, Lindley, Domingoa haematochila (Reichenbach f.) Carabia, non-endemic species), and the presence of species of restric- Encyclia fucata (Lindley) Britton & Millspaugh, E. plicata ted distribution of every island or group. Are all the (Lindley) Schlechter, Tetramicra urbaniana Cogniaux, unresolved clusters of islands in the consensus trees, inde- T. gauntletii (Withner & Jesup) Braem, T. lucayana (Nash pendent biogeographical provinces? ex Britton & Millspaugh) Braem]. Thus, the relationships The Eastern Antilles, from the Virgin Islands to Margarita among the Bahamas and the Greater Antilles seems to be constitute a distinct branch with several minor subgroups. considerable. First, the larger Lesser Antilles from Guadeloupe to St Lucia is Lastly, Aruba, Bonaire and CuracËao form another inde- the most de®ned group, with seven endemic species [Ellean- pendent group. These islands have only three species thus dussii Cogniaux, Epidendrum discoidale Lindley, Max- reported, none of which is endemic. One occurs on different illaria guadalupensis Cogniaux, Pleurothallis dussii continental and insular areas [Polystachya foliosa (Hooker) Cogniaux, P. ophioglossoides (Jacquin) Garay, Pseudocen- Reichenbach f.], one held in common with Central and trum guadalupense Cogniaux, Stelis dussii Cogniaux]. Sec- South America [Brassavola nodosa (Linnaeus) Lindley], and ondly, the smaller Lesser Antilles (islands north of one shared only with Venezuela [Myrmecophila humboldtii Guadeloupe from Montserrat to St Martin) is a `group' that (Reichenbach f.) Rolfe]. Thus, the ABC islands show no splits along the Eastern Antilles branch (Figs 1 and 2). Some special af®nity to the rest of the Antilles. of these islands share two endemic species with the larger Lesser Antilles group (Epidendrum difforme Jacquin and Epidendrum patens Swartz). We recognize that this smaller Relationships with continental areas Lesser Antilles `group' is not cast in stone because basal The inclusion of the Guianas as a South American contin- species-poor areas are prone to changing af®nities. Thirdly, ental area was thought as a heuristic exercise. First, the the Virgin Islands, except Anegada, constitute a monocladic Guianas is a politically de®ned area rather than a biogeo- but weakly differentiated (no endemic species) group which is graphical region. Secondly, the Guianas is likely outside the ®rmly allied with the Lesser Antilles. Nevertheless, the Virgin Caribbean ¯oristic region and part of the Amazonian region Islands have two species that they share exclusively with (Gentry, 1982). If we were to include other South American neighbouring Puerto Rico [Psychilis macconnelliae Sauleda, regions in this study, no doubt the Guianas would shift Tolumnia prionochila (KraÈnzlin) Braem]. This connection af®nities dramatically. The Guianas form a sister group to was insuf®cient to alter the ¯oristic af®nities of the Virgin the Greater Antilles, which is indicative of the strong

Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794 Biogeography of the Antilles based on a parsimony analysis of orchid distributions 781 af®nities among insular and continental orchid ¯oras (as has Every main cluster has its own species-poor islands. We been put forth for the ¯ora in general by Samek, 1988; interpret the basal position of an area as indicative of Borhidi, 1996). ¯oristic subordination to the distal areas in the group. The Yucatan and Florida form a sister pair of the north- Otherwise, when the few species of an area do not de®ne its western Bahamas/Isla de la Juventud group. All these are af®liation, such an area turns out to be unresolved or sedimentary, calcareous, low-lying and ¯at areas. Samek unstable, being located at the base of the whole tree or at the (1988) and Borhidi (1996) de®ned South Florida together base of different branches, respectively. In other words, with the Bahamas as one province. On the other hand, either a high or a low number of species can de®ne Gentry (1982) excluded Florida (except for the Keys) from membership patterns among areas. Limiting the analysis to his Caribbean (insular) region. We consider Florida and areas with a similar number of species would unnecessarily the Yucatan as provinces not belonging to the Antillean restrict the use of this methodology for the Caribbean and subregion, which are supposedly part of the Continental many other regions. subregion of the Caribbean. Differences in species numbers within our island groups To de®ne more clearly the relationships among the insular can be converted into species±area curves. We have found Antilles and their neighbouring continental areas, more that the geologically and physiographically diverse montane thorough studies are necessary. Also, the inclusion of other islands have a different species±area curve than that for low- continental areas, working as distant sister areas in the lying islands, which are basically calcareous and physio- analyses, would be useful. For example, a sister continental graphically homogeneous (Ackerman et al., submitted). The area for the Yucatan biogeographical province (DuraÂn et al., calcareous group, comprised of nineteen islands (including 1998) could be a province in central-western Mexico. Both the Cayman Islands), has a z-value of 0.25 (however, r ˆ 0.4, outside provinces belong to the same ¯oristic Mexican- P ˆ 0.07), a typical value for islands (Rosenzweig, 1995). Central American Caribbean subregion (sensu Samek, 1988). Meanwhile, the montane island group, composed of twenty- six islands (our island arc group), has a z-value of 0.51 (r ˆ 0.89, P ˆ 0.0001). This is close to an interarchipelagic Islands with unresolved af®nity value found by Adler (1992, cited by Rosenzweig, 1995). The unresolved islands or island groupings found in some Moreover, if a species±area curve for this montane group is consensus trees could be explained in a number of ways. calculated with only the endemic species (using only nine First, it is possible that an incomplete record of the species islands that have single-island endemics), z ˆ 0.68 (r ˆ 0.88, for these islands could be hampering the resolution of their P ˆ 0.0019) suggesting that habitat diversity is important for af®nities. As we improved the species record for the smaller autochthonous speciation. The two z-values for the montane Lesser Antillean islands they became progressively resolved. group seem to indicate that the Island Arc group behaves as a We expect this trend to continue. Alternatively, the record cluster of provinces (Rosenzweig, 1995) as has been may be complete and these unresolved areas could be the proposed by Borhidi & MunÄ iz (1986). In contrast, the result of: (1) a lower number of species which would prevent endemic species±area curve for the relatively homogeneous their af®liation with other groups, (2) an equally strong calcareous group was not run because there are only three af®nity with more than one of the other groups, and (3) a islands (island banks indeed) with endemic species. non-hierarchical structure in the data (Ronquist, 1997; Glasby & Alvarez, 1999). One way an unresolved area or Phytogeographical regionalization of the Antilles group could be considered distinctive would be if it had a well-differentiated ¯ora (i.e. endemic or exclusive species). It The Antilles is among the principal phytogeographical is likely a combination of these factors that play a role in regions of the world (Gentry, 1982). It has been included unresolved cases. within the Caribbean region that extends to continental lands from Mexico to Venezuela (Samek, 1988; Borhidi, 1996). Our orchid data seem to support this view. The PAD, species richness and species±area curves phytogeographic sectorization of Samek (1988) and Borhidi The position of areas in tree branches is in¯uenced by species (1996) consider every one of the Greater Antilles as numbers. This pattern is not evident for the Antillean provinces. Major geographical aggregations of islands (i.e. archipelago as a whole and only becomes obvious within the the Bahamas, the Lesser Antilles) are also de®ned as major island assemblages. As we move from the tip to the provinces, while minor groups of islands (e.g. the Virgin base of the tree branches, species numbers tend to diminish. Islands, the Cayman Islands) are integrated with some of the This apparent numerical artefact has been stated as one of big islands. Apart from the recognition of these kind of the main problems resulting from the use of areas with divisions, we also examined af®nities among and within highly dissimilar number of species, and a comparable them in a tree-like hierarchical structure (Fig. 2). number of taxa per site has been recommended as a pre- 1 Every one of the major islands of the Greater Antilles requisite for PAE analyses (Rosen, 1988, 1992; Rosen & (i.e. Cuba, Hispaniola, Jamaica and Puerto Rico) is Smith, 1988; Vargas, 1991; Conran, 1995). What our study suf®ciently differentiated to be recognized as a separate shows is that even with very few species the af®liation of an province of the Antilles. island to a speci®c group occurs when the taxa permit it.

Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794 782 J. C. Trejo-Torres and J. D. Ackerman

2 The Lesser Antilles were taken as a single phytogeo- ¯ying birds and bats. In this case, area af®nities appear to be graphic province by Samek (1988) and Borhidi (1996). de®ned by physical factors, such as geomorphology, physi- Their orchid ¯ora is suf®ciently distinctive, supporting ography, climate, geography and island area. At the same this view. Our analysis places them as part of the Eastern time, it is plausible to predict that less vagile organisms, such Antilles group and at the same time subordinate to the as non-¯ying vertebrates and freshwater ®shes, will de®ne Greater Antillean main group. different patterns of area relationships, perhaps more in¯u- 3 The Virgin Islands were integrated as part of Puerto Rico enced by geological and geographical history of areas. by Samek (1988) and Borhidi (1996). However, our study The PAE/PAD is an alternative to the use of multivariate places them rather close to the Lesser Antilles, and phenetic methods for classi®cation of biotas (Rosen, 1988; altogether, as subordinate of the Greater Antilles. Vargas, 1991) and for comparing species assemblages. In 4 The Cayman Islands were considered part of Jamaica by fact, giving a static interpretation of area relationships, this Samek (1988) but as part of Cuba according to Borhidi is a method equivalent to multivariate ones (Rosen, 1988, (1996). Our analysis suggests that these islands have 1992). Some important differences between PAE/PAD and complex af®nities with the Greater Antilles, the Yucatan multivariate methods are: (1) The PAE/PAD uses parsimony and Florida continental areas, and also with the Bahamas. algorithms instead of similarity indexes. This means that the 5 The Bahama archipelago was one of the Caribbean hierarchy selected in this manner will be the best supported provinces distinguished by Samek (1988), Borhidi (1996), by evidence, for it maximizes congruence between data and and Gentry (1982), and this is also supported by our hierarchical patterns (Brady, 1994). (2) The PAE/PAD uses orchid data. Moreover, our study relates the Isla de la characters (taxa) selected a priori based on their informat- Juventud, Mona and Anegada (geographically and geo- iveness, that is, species found in all sites and single-site logically part of the Greater Antilles) as subordinates of species are eliminated as they give no information on the the Bahamian Province. The Isla de la Juventud shows a af®nities among areas (3). The PAE/PAD provides a more strong af®nity with the north-western Bahamas, while uni®ed methodology as multivariate methods use many Mona and Anegada are small islands with close af®nity to different similarity indices, which can also yield different the rest of the Bahamas. results. 6 Trinidad, Tobago and Margarita were considered part of The PAE/PAD can be seen as a complementary method. the Venezuela±Colombia Caribbean subregion by Samek While many community analyses are intended to describe (1988) and Borhidi (1996), while no clear af®liation for quantitative parameters, PAE/PAD is directed to describe Trinidad and Tobago can be obtained from Gentry composition patterns, based on the identity of species (1982). Our data support a closer af®nity of Trinidad (Worthen, 1996). Also, results of PAE/PAD in the form of with the Greater Antilles instead of the neighbouring trees with a hierarchical structure could be a graphic Lesser Antilles. Based on its differentiated orchid ¯ora complement to other methods which produce results that (approximately thirteen endemic species) we consider are single numerical values, such as indexes, scores or Trinidad as a ¯oristic province. On the other hand, exponents (i.e. nestedness analysis, Patterson & Atmar, Margarita and Tobago are linked with the Lesser Antilles± 1986). The hierarchical nature of the results does not Virgin Islands group rather than with Trinidad. There are necessarily mean that communities are organized that way. two potential problems with these islands. First, we After all, af®nities among communities may have an consider that the species record for these islands is not underlying reticulate structure (Ronquist, 1997). We view well updated, especially for Trinidad and Tobago whose PAE/PAD as only one way to represent structure in biogeo- most recent complete orchid ¯ora was published by graphical data. Schultes (1960). Secondly, the Venezuelan Guyana may The PAE/PAD has the potential as a method for under- be a more appropriate neighbouring continental area, standing ecological biogeographic problems. This is suppor- instead of our Guianas area. If this is true, the islands of ted by the Caribbean orchids for which composition patterns the Trinidad bank may be more related to the continent strikingly match physical and ecological factors. The use of than to the rest of the Antillean islands, as suggested by the method as a complement to vicariance and other histor- bat distribution patterns (Trejo-Torres & Rivera, unpub- ical biogeographic methods has been suggested (Cracraft, lished data) and by geological history. 41991). Because of its phenetic nature, patterns found 7 Aruba, Bonaire and CuracËao seem to be independent of through PAE/PAD could be re¯ecting ecological phenomena, the Antillean subregion and apparently part of the rather than historical ones. Nevertheless, until now, it has Continental Caribbean subregion. always been used under the historical point of view. Some of the aspects pointed out as disadvantages to the historical approach, do not exist in the ecological context. For Perspectives of PAD example, ¯oristic af®nities, biodiversity comparisons and The pattern we found for orchids is not necessarily the same composition patterns can be seen as phenetic phenomena. for the general ¯ora or for the entire biota of the region. We For the demarcation of biogeographical units PAE/PAD expect that phytogeographic relationships revealed by could be employed before searching for speci®c patterns in orchids might be similar to that of other highly vagile groups distributional data sets. This could avoid distortions from of organisms such as wind-dispersed plants, ¯ying insects, possible structure in the data caused by mixing areas

Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794 Biogeography of the Antilles based on a parsimony analysis of orchid distributions 783 belonging to different biotas. Within the same context, The highly compatible patterns obtained from different Morrone & Crisci (1995) suggested the use of panbiogeo- island sets and from different consensus trees indicate that graphic methods as an initial step to do other historical relationships of areas obtained are well supported by the biogeographic analyses. data. In other words, the groups identi®ed are not arti®cial, The PAE/PAD could prioritize conservation efforts at the but rather the result of their orchid-species af®nities. regional level (Posadas, 1996) by identifying areas or islands Whereas PAE/PAD is an appropriate methodology for the of high diversity. We have shown that such use can be study of geographical regions based on shared species, helpful for the Antilles, one of the world's biodiversity single-site species (the majority of which are endemics) are hotspots (Myers et al., 2000). However, conservation indicative of ®ner degrees of biogeographical differentiation. efforts often focus on unique species, particularly endemics We suggest using both data sets to analyse biogeographical (island endemics in our case), but PAD uses only shared af®nities, for it integrates differences and similarities. This species. The exclusive use of PAD in formulating conserva- approach provides a more thorough use of ¯oristic infor- tion strategies can thus be misleading. This is especially true mation for the analysis of biogeographical relationships. for species-poor islands that appear subordinated to species- The parsimony methodology seems to be appropriate for rich islands. Impoverished islands may thus be falsely the analysis of areas, or any other natural unit, with very interpreted as irrelevant for biodiversity conservation. For dissimilar number of species. The low number of species example, the tiny island of Mona lies subordinate to other found in a unit can de®ne its af®liation or can produce an islands of our calcareous group yet it is certainly worthy of unresolved ¯oristic af®nity. The lack of af®nity of an area to conservation efforts because Mona has its own set of any group may be either real or an artefact caused by an endemic plants and animals (including an orchid, Psychilis incomplete species record. monensis Sauleda). The PAE/PAD has great potential as an alternative, The PAE/PAD do have problems that need to be resolved complementary or ®rst-step method to other multivariate and these are the interpretation of: (1) the unresolved status and phylogenetic methodologies in biogeography. among the main groupings of areas, (2) the area relation- ships when using islands as well as continental regions, (3) ACKNOWLEDGMENTS the potential use of branch-lengths in the trees. Special attention should also be given when selecting data sets for The study was supported by NSF grants to JDA (DEB- organisms with diverse ecological characteristics (e.g. birds, 9505459, HRD-9353549, HRD-9628475). Arkelio Alicea, which could belong to different guilds: residents vs. migra- Kary GarcõÂa, Eileen Bravo, and Yanet Crespo were students tory, volant vs. non-volant). participating in the study under Alliance for Minorities Participation Program at UPR. We thank Steve Rehner logistic assistance in data analysis. We are grateful to Brian CONCLUSIONS Rosen, David Lees, Susan Arago n, Marta DõÂaz, and Juan According to our orchid data, the Antillean islands consti- J. Morrone who had made relevant comments on the tute a phytogeographical subregion of the Caribbean. The manuscript. Timothy Johnston assisted editing the manu- Lesser Antilles and the Bahamas are well-de®ned provinces. script and Celene Espadas, Guillermo Bianchi, and Ana Meanwhile, the Virgin Islands and the Cayman Islands are Porzecanski shared important references. GermaÂn Carnevali, poorly differentiated aggregates, perhaps to be considered as Andre van Proosdij, Mark Nir and Hagen Stenzel provided incipient provinces. All of them are subordinated to the main unpublished data. Greater Antillean group. Within islands groupings detected by PAD ordering of REFERENCES areas is highly in¯uenced by species numbers. The species- rich islands form cohesive groupings to which other species- Ackerman, J.D. 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Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794 786 J. C. Trejo-Torres and J. D. Ackerman

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 the biogeography of Caribbean bats. He worked on the community description, ¯oristics and taxonomy of the Yucatan forests and wetlands, 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 and evolution, particularly of orchids. He has published numerous papers on the evolution of deception pollination and the processes involved in the diversi®cation of the Orchidaceae. Among his current projects is the Orchid treatment for the Flora of the Greater Antilles.

Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794 Ó Appendix 1 Data matrix of the ®fty-two Caribbean areas and the 356 shared orchid species included in the study. Species names have been standardized by Ackerman. Species lcwl cec t 2001, Ltd Science Blackwell presence is indicated with a one, while absence with a zero. The Bahama islands are grouped following Correll & Correll (1982). See Table 1 for Bahama island groups and other geographical groupings lemy Âe ao Ëc

Species name/area name St Kitts Nevis St Eustatius Antigua Barbuda Montserrat Guadeloupe Dominica Martinique St Lucia St Vincent Grenada Barbados Trinidad Tobago Margarita Vieques Culebra Anegada Virgin Gorda Tortola St Thomas St John St Croix Aruba Cura Bonaire Guianas Yucatan Florida Puerto Rico Jamaica Hispaniola Cuba Isla Juventud Mona Grand Cayman Little Cayman Cayman Brac Turks Inaguas Crooked Long Salvador Exumas Cat Providence Andros Abacos St Martin Saba St Barth Aganisia pulchella 1000000000000000000000000000000000000010000000000000

ora fBiogeography of Journal Antillanorchis gundlachii 0000011000000000000000000000000000000000000000000000 Aspasia variegata 1000000000000000000000000000000000000010000000000000 Basiphyllaea corallicola 0011111000000000100011000000000000000000000000000000 Basiphyllaea sarcophylla 0000001100000000000000000000000000000000000000000000 Batemania colleyi 1000000000000000000000000000000000000010000000000000 Beloglottis costaricensis 1110010001000000000000000000000000000010000000000000 Bifrenaria aurantiaca 0010000000000000000000000000000000000010000000000000 Bifrenaria longicornis 1000000000000000000000000000000000000010000000000000

, Bletia patula 0011111000000000000000000000000101000010000000000000 28 Bletia purpurea 1110111100000000000111000000000000000000000000000000 775±794 , Brachionidium parvum 0001010000000000000000000000000110001000000000000000 Brachionidium sheringii 0001010000000000000000000000000000000000000000000000 igorpyo h nilsbsdo asmn nlsso rhddsrbtos787 distributions orchid of analysis parsimony a on based Antilles the of Biogeography Brachystele guayanensis 1100000000000000000000000000000000000010000000000000 Brassavola cucullata 1100000000000000000000111001101110011011000000100000 Brassavola nodosa 1000000000000000000000000000000000000000000000000111 Brassia caudata 1110111000000000100000000000000000000010000000000000 Brassia maculata 0100101000000000000000000000000000000000000000000000 Broughtonia domingensis 0000010010000000000000000000000000000000000000000000 Broughtonia lindenii 0000001100000011010111000000000000000000000000000000 Broughtonia ortgiesiana 0000001100000000000000000000000000000000000000000000 Bulbophyllum aristatum 0100011000000000000000000000000000000000000000000000 Bulbophyllum pachyrrhachis 1011111000000000000000000000000000000010000000000000 Calanthe calanthoides 0000111000000000000000000000000000000000000000000000 Calopogon tuberosus 0010001000000000000111000000000000000000000000000000 Campylocentrum fasciola 1101011000000000000000000000000000000010000000000000 Campylocentrum ®liforme 0001111000000000000000000000000000000000000000000000 Campylocentrum micranthum 1101111000000000000001000000000110000010000000000000 Campylocentrum pachyrrhizum 1111111000000000000000000000000000000010000000000000 Campylocentrum poeppigii 1100001000000000000000000000000000000000000000000000 Campylocentrum pygmaeum 0001001000000000000000000000000100000000000000000000 Catasetum cristatum 1000000000000000000000000000000000000010000000000000 Catasetum integerrimum 0100001000000000000000000000000000000000000000000000 Catasetum macrocarpum 1000000000000000000000000000000000000010000000000000 Caularthron bicornutum 1000000000000000000000000000000000000010000000000000 Cochleanthes ¯abelliformis 1001111000000000000000000000000000000010000000000000 Coelia triptera 0000101000000000000000000000000000000000000000000000 Comparettia falcata 1001111000000000000000000000000000000000000000000000 Coryanthes macrantha 1000000000000000000000000000000000000010000000000000 Coryanthes speciosa 1000000000000000000000000000000000000010000000000000 Corymborkis ¯ava 0000101000000000000000000000000100000000000000000000 Corymborkis forcipigera 0101011000000000000000000000000000000000000000000000 Cranichis diphylla 1000110000000000000000000000000000000000000000000000 Cranichis muscosa 0011111000000000000010000100001111111011000000000000 Cranichis ovata 0000010000000000000000000000000110101000000000000000 Appendix 1 continued Ackerman D. J. and Trejo-Torres C. J. 788 lemy Âe ao Ëc

Species name/area name St Kitts Nevis St Eustatius Antigua Barbuda Montserrat Guadeloupe Dominica Martinique St Lucia St Vincent Grenada Barbados Trinidad Tobago Margarita Vieques Culebra Anegada Virgin Gorda Tortola St Thomas St John St Croix Aruba Cura Bonaire Guianas Yucatan Florida Puerto Rico Jamaica Hispaniola Cuba Isla Juventud Mona Grand Cayman Little Cayman Cayman Brac Turks Inaguas Crooked Long Salvador Exumas Cat Providence Andros Abacos St Martin Saba St Barth Cranichis ricartii 0001011000000000000000000000000000000000000000000000 Cranichis tenui¯ora 0000011000000000000000000000000000000000000000000000 Cranichis tenuis 0001011000000000000000000000000000000000000000000000 Cranichis wageneri 0000111000000000000000000000000000000000000000000000 Cryptarrhena lunata 0000100000000000000000000000000000000010000000000000 Cyclopogon cranichoides 0111111001000000000000000000000000001110000000010000 Cyclopogon elatus 1011111001000000000000101001101111101110000001111000 Cyclopogon laxi¯orus 0000111000000000000000000000000000000000000000000000 Cyclopogon miradorense 0001011000000000000000000000000000000000000000000000 Cyrtopodium andersonii 1000000000000000000000100001101111111010000000000000 Cyrtopodium punctatum 0011011001000000000000000000000000000010000000000000 Dendrophylax barrettiae 0000110000000000000000000000000000000000000000000000 Dendrophylax lindenii 0010011000000000000000000000000000000000000000000000 Dendrophylax varius 0000011000000000000000000000000000000000000000000000 Dichaea glauca 0000111000000000000000000000000000000000000000000000 Dichaea graminoides 0000111000000000000000000000000000000010100000000000 Dichaea histrio 1000000000000000000000000000000000000000100000000000 Dichaea hookeri 1000000000000000000000000000000110010010000000000000 Dichaea hystricina 0001011000000000000000000000000110001010000000000000 Dichaea latifolia 0001111000000000000000000000000000000000000000000000 Dichaea morrisii 1000111000000000000000000000000111011010000000000000 Dichaea pendula 1001111000000000000000000000000101000000100000000000

Ó Dichaea picta 1000000000000000000000000000000000000010100000000000

lcwl cec t 2001, Ltd Science Blackwell Dichaea rendlei 1000000000000000000000000000000000000010000000000000 Dichaea trichocarpa 0000111000000000000000000000000000000000000000000000 Dilomilis elata 0000101000000000000000000000000000000000000000000000 Dilomilis montana 0001111000000000000000000000000000000000000000000000 Domingoa haematochila 0000011010000000001000000000000000000000000000000000 Domingoa nodosa 0000011000000000000000000000000000000000000000000000 Elleanthus caravata 1000000000000000000000000000000000011000000000000000 Elleanthus cephalotus 1000111000000000000000000000000101011010000000000000 Elleanthus cordidactylus 0001111000000000000000000000000000000000000000000000 Elleanthus dussii 0000000000000000000000000000000101000000000000000000 Elleanthus longibracteatus 0000100000000000000000000000000010001000000000000000 ora fBiogeography of Journal Eltroplectris calcarata 1011111001100000000111000000000000001000000000000000 Encyclia acutifolia 0000011000000000000000000000000000000000000000000000 Encyclia angustifolia 0000011000000000000000000000000000000000000000000000 Encyclia brevifolia 0000001100000000000000000000000000000000000000000000 Encyclia cordigera 1000000000000000000000000000000000000000100000000000 Encyclia fehlingii 0000000000000000000111000000000000000000000000000000 Encyclia fucata 0000001100000000000011000000000000000000000000000000 Encyclia gracilis 0000000000001111011111000000000000000000000000000000 ,

28 Encyclia gravida 0001111000000000000000000000000000000000000000000000

775±794 , Encyclia inaguensis 0000000000001100000000000000000000000000000000000000 Encyclia isochila 0001111000000000000000000000000000000000000000000000 Encyclia kingsii 0000000001010000000000000000000000000000000000000000 Encyclia nematocaulon 0100001000000000000000000000000000000000000000000000 Ó Appendix 1 continued lcwl cec t 2001, Ltd Science Blackwell lemy Âe ao Ëc

Species name/area name St Kitts Nevis St Eustatius Antigua Barbuda Montserrat Guadeloupe Dominica Martinique St Lucia St Vincent Grenada Barbados Trinidad Tobago Margarita Vieques Culebra Anegada Virgin Gorda Tortola St Thomas St John St Croix Aruba Cura Bonaire Guianas Yucatan Florida Puerto Rico Jamaica Hispaniola Cuba Isla Juventud Mona Grand Cayman Little Cayman Cayman Brac Turks Inaguas Crooked Long Salvador Exumas Cat Providence Andros Abacos St Martin Saba St Barth Encyclia oncidioides 1000000000000000000000000000000000000010000000000000 Encyclia plicata 0000001000000000000111000000000000000000000000000000 Encyclia rufa 0010001000001111111111000000000000000000000000000000

ora fBiogeography of Journal Encyclia serrulata 0000111000000000000000000000000000000000000000000000 Encyclia tampensis 0010001000000000000011000000000000000000000000000000 Epidendrum acunae 0010001000000000000000000000000000000000000000000000 Epidendrum anceps 1011111100000000000000110101100111111100100010110000 Epidendrum antillanum 0001010000000000000000010000000111010000000000000000 Epidendrum blancheanum 0000111000000000000000000000000000000000000000000000 Epidendrum carpophorum 1001111000000000000000000000000110111010000000000000 Epidendrum ciliare 1101111000000000000000111011100111111110111011111000

, Epidendrum compressum 1000000000000000000000000000000000000010000000000000 28 Epidendrum coronatum 1000000000000000000000000000000000000010000000000000 775±794 , Epidendrum cristatum 1000000000000000000000000000000000000010000000000000 Epidendrum dendrobioides 1000000000000000000000000000000111010000000000000000 Epidendrum difforme 0000000000000000000000000101000111111000000000000000 789 distributions orchid of analysis parsimony a on based Antilles the of Biogeography Epidendrum diffusum 0100101000000000000000000000000000000000000000000000 Epidendrum ¯oridense 0010001000000000000000000000000000000000000000000000 Epidendrum hartii 0000000000000000000000000000000000000011000000000000 Epidendrum ibaguense 1000000000000000000000000000000111010010000000000000 Epidendrum imbricatum 0000110000000000000000000000000000000000000000000000 Epidendrum isomerum 1100000000000000000000000000000000000000000000000000 Epidendrum jamaicense 0001111000000000000000000000000110000000000000000000 Epidendrum miserrimum 1001010000000000000000000000000111101000000000000000 Epidendrum mutelianum 0001100000000000000000000000000000000000000000000000 Epidendrum neoporpax 0000011000000000000000000000000000000000000000000000 Epidendrum nocturnum 1111111101000000000111000000000111011010100000000000 Epidendrum nutans 1000100000000000000000000000000000000000000000000000 Epidendrum orientale 0000101000000000000000000000000000000000000000000000 Epidendrum pallidi¯orum 0000001100000000000000000000001010011000000000000000 Epidendrum paranaense 0000110000000000000000000000000000000000000000000000 Epidendrum patens 0000000000000000000000000110000101000000000000000000 Epidendrum polygonatum 0000011000000000000000000000000000000000000000000000 Epidendrum portoricensis 0001011000000000000000000000000000000000000000000000 Epidendrum ramosum 1001111000000000000000000000000111111010000000000000 Epidendrum repens 0000111000000000000000000000000000000000000000000000 Epidendrum rigidum 1011111100000000000111000000000111111010100000000000 Epidendrum rivulare 0000111000000000000000000000000000000000000000000000 Epidendrum scapelligerum 0000101000000000000000000000000000000000000000000000 Epidendrum schlecterianum 1000000000000000000000000000000000000010000000000000 Epidendrum secundum 0001111000000000000000000000100111111000000000000000 Epidendrum strobiliferum 1110111000000000000000010000000111011010000000000000 Epidendrum vincentinum 1001111000000000000000000000000000000010000000000000 Epidendrum wrightii 0000011000000000000000000000000000000000000000000000 Epistephium ellipticum 1000000000000000000000000000000000000010000000000000 Epistephium parvi¯orum 1000000000000000000000000000000000000010000000000000 Erythrodes hirtella 1001111000000000000000000001000111111110000001000000 Appendix 1 continued Ackerman D. J. and Trejo-Torres C. J. 790 lemy Âe ao Ëc

Species name/area name St Kitts Nevis St Eustatius Antigua Barbuda Montserrat Guadeloupe Dominica Martinique St Lucia St Vincent Grenada Barbados Trinidad Tobago Margarita Vieques Culebra Anegada Virgin Gorda Tortola St Thomas St John St Croix Aruba Cura Bonaire Guianas Yucatan Florida Puerto Rico Jamaica Hispaniola Cuba Isla Juventud Mona Grand Cayman Little Cayman Cayman Brac Turks Inaguas Crooked Long Salvador Exumas Cat Providence Andros Abacos St Martin Saba St Barth Erythrodes major 1000010000000000000000000000000000000000000000000000 Erythrodes plantaginea 0001111000000000000000010101001111011011000000000000 Eulophia alta 1111111000000000000001000000100111111011000001000000 Eulophia ecristata 0010001000000000000000000000000000000000000000000000 Eurystyles ananassocomos 0001111000000000000000000000000000000000000000000000 Fuertesiella pterichoides 0000011000000000000000000000000000000000000000000000 Galeandra beyrichii 0011111000000000000000000000000000000000000000000000 Govenia utriculata 0011111000000000000011000000000000000000000000000000 Habenaria alata 0001111100000000000010000000100111111011000000100000 Habenaria amal®tana 1001000000000000000000000000100100010000000000000000 Habenaria bicornis 0000001100000000000000000000000000000000000000000000 Habenaria distans 0111111000000000000000000000000000000000000000000000 Habenaria dussii 0000000000000000000000000000000100010000000000000000 Habenaria eustachya 0001111000000000000000000000000100000000000000000000 Habenaria ¯oribunda 1010111100000000000011000000000100000000000000000000 Habenaria leprieurii 1000000000000000000000000000000000000010000000000000 Habenaria mesodactyla 0100000000000000000000000000000000000010000000000000 Habenaria monorrhiza 0101111000000000000000000000100111111111100001101000 Habenaria pauci¯ora 0100000000000000000000000000000000000010000000000000 Habenaria quinqueseta 1010111000000000000011000000000000000000000000000000 Habenaria repens 1111111000000000000000000000000000000010000000000000 Hapalorchis lineatus 0001111000000000000000000000000000000000000000000000

Ó Harrisella porrecta 0111111101000000000000000000000000000000000000000000

lcwl cec t 2001, Ltd Science Blackwell Helleriella punctulata 0001000000000000000000000000000100001000000000000000 Homalopetalum vomeriforme 0000101000000000000000000000000000000000000000000000 Huntleya lucida 1000000000000000000000000000000000000010100000000000 Huntleya meleagris 1000000000000000000000000000000000000010000000000000 Ionopsis satyrioides 1101111000000000100000000000000001000010000000000000 Ionopsis utricularoides 1111111110000000100000001000000101110010000000100000 Isochilus linearis 1001111000000000000000000000001111010010000000000000 Jacquinella globosa 1101111000000000000000010101000111111010100000000000 Jacquinella teretifolia 1001111000000000000000000000000000000000000000000000 Kegeliella houtteana 1000000000000000000000000000000000000010000000000000 Koellensteinia graminea 1000000000000000000000000000000000000010000000000000 ora fBiogeography of Journal Leochilus carinatus 0000001000000000000000000000000000000010000000000000 Leochilus labiatus 0000111000000000100000000000000111010011000000000000 Leochilus puertoricensis 0001000000000000000000000000000100010000000000000000 Leochilus scriptus 0100001000000000000000000000000000000010000000000000 Lepanthopsis anthoctenium 0000011000000000000000000000000000000000000000000000 Lepanthopsis melanantha 0011111000000000000000000000000000000000000000000000 Lepanthopsis melanantha 0011111000000000000000000000000000000000000000000000 Lepanthopsis microlepanthes 0000111000000000000000000000000000000000000000000000 ,

28 Leucohyle subulata 1000101000000000000000000000000000000010000000000000

775±794 , Liparis nervosa 1111111000000000000000000000110111011010000001100000 Liparis neuroglossa 0000110000000000000000000000000000000000000000000000 Liparis saundersiana 0001111000000000000000000000000000000000000000000000 Liparis vexillifera 0001111000000000000000000000000000000010000000000000 Ó Appendix 1 continued lcwl cec t 2001, Ltd Science Blackwell lemy Âe ao Ëc

Species name/area name Guianas Yucatan Florida Puerto Rico Jamaica Hispaniola Cuba Isla Juventud Mona Grand Cayman Little Cayman Cayman Brac Turks Inaguas Crooked Long Salvador Exumas Cat Providence Andros Abacos St Martin Saba St Barth St Kitts Nevis St Eustatius Antigua Barbuda Montserrat Guadeloupe Dominica Martinique St Lucia St Vincent Grenada Barbados Trinidad Tobago Margarita Vieques Culebra Anegada Virgin Gorda Tortola St Thomas St John St Croix Aruba Cura Bonaire Liparis viridipurpurea 0000111000000000000000000000000000000000000000000000 Lockhartia elegans 1000000000000000000000000000000000000010000000000000 Lophiasis maculata 1110101100000000000000000000000011111000100000000000

ora fBiogeography of Journal Lycaste barringtoniae 0001111000000000000000000000000000000000000000000000 Macradenia lutescens 1010111100000000000000000000000000000010000000000000 Malaxis domingensis 0000011000000000000000000000000000000000000000000000 Malaxis hispanolae 0000011000000000000000000000000000000000000000000000 Malaxis integra 0000110000000000000000000000000000000000000000000000 Malaxis major 0001001000000000000000000000000110000000000000000000 Malaxis massonii 0001111000000000000000000100000111000010000000000000 Malaxis spicata 0011111000000010101011000000000000000000000000000000

, Malaxis umbelli¯ora 1000111000000000000000000100000110010010000000000000 28 Malaxis unifolia 0010111000000000000000000000000000000000000000000000 775±794 , Maxilaria discolor 1000000000000000000000000000000000000010000000000000 Maxillaria acutifolia 1001111000000000000000000000000000000000000000000000 Maxillaria adendrobium 0000111000000000000000000000000000000000000000000000 791 distributions orchid of analysis parsimony a on based Antilles the of Biogeography Maxillaria alba 1000101000000000000000000000000111010010000000000000 Maxillaria brachybulbon 1000000000000000000000000000000000000010000000000000 Maxillaria camaridii 1000000000000000000000000000000000000010000000000000 Maxillaria coccinea 0001010000000000000000010100000111011010000000000000 Maxillaria crassifolia 1110101100000000000000000000000000000000000000000000 Maxillaria in¯exa 0000111000000000000000000000000110001000000000000000 Maxillaria liparophylla 1000000000000000000000000000000000000010000000000000 Maxillaria parvi¯ora 1011111000000000000000000000000000001010000000000000 Maxillaria reichenheimana 1000000000000000000000000000000000000010000000000000 Maxillaria rufescens 1000000000000000000000000000000000000010000000000000 Maxillaria uncata 1100000000000000000000000000000000000000000000000000 Maxillaria variabilis 0100000000000000000000000000000000000010000000000000 Mesadenus lucayanus 0111011010001011111111000000100000000000000100100000 Myrmecophila thomsoniana 0000001001110000000000000000000000000000000000000000 Nidema ottonis 1001111000000000000000000000000000000010000000000000 Notylia angustifolia 1000000000000000000000000000000000000010000000000000 Octomeria apiculata 0000000000000000000000000000000100001011000000000000 Octomeria graminifolia 1000000000000000000000000000000111110010000000000000 Octomeria tridentata 0000001000000000000000000000000000000010000000000000 Oeceoclades maculata 1111111010000000000110000000000000000000000000010000 Oerstedella verrucosa 0000101000000000000000000000000000000000000000000000 Oncidium altissimum 0001010000000000000000000000000011010000000000100000 Oncidium ampliatum 0001000000000000000000000000000111111010000000000000 Oncidium cebolleta 1100000000000000000000000000000101100010000000000000 Oncidium ensatum 0110001100000010000111000000000000000000000000000000 Oncidium meirax 0001010000000000000000000000000100000000000000000000 Ornithocephalus gladiatus 1000000000000000000000000000000000001010000000000000 Otostylis brachystalix 1000000000000000000000000000000000000010000000000000 Palmorchis pubescens 1000000000000000000000000000000000000010000000000000 Paphinia cristata 1000000000000000000000000000000000000010000000000000 Pelexia adnata 0101111000000000000010000000000000000000000000000000 Appendix 1 continued Ackerman D. J. and Trejo-Torres C. J. 792 lemy Âe ao Ëc

Species name/area name St Kitts Nevis St Eustatius Antigua Barbuda Montserrat Guadeloupe Dominica Martinique St Lucia St Vincent Grenada Barbados Trinidad Tobago Margarita Vieques Culebra Anegada Virgin Gorda Tortola St Thomas St John St Croix Aruba Cura Bonaire Guianas Yucatan Florida Puerto Rico Jamaica Hispaniola Cuba Isla Juventud Mona Grand Cayman Little Cayman Cayman Brac Turks Inaguas Crooked Long Salvador Exumas Cat Providence Andros Abacos St Martin Saba St Barth Peristeria cerina 1000000000000000000000000000000000000010000000000000 Peristeria pendula 1000000000000000000000000000000000000010000000000000 Pinelia leochilus 0000011000000000000000000000000000000000000000000000 Platanthera replicata 0000001100000000000000000000000000000000000000000000 Platystele ovalifolia 1100001000000000000000000000000000000000000000000000 Platystele stenostachia 1100000000000000000000000000000000000000000000000000 Platythelys querceticola 0010111000000000000011000000000101000000000000000000 Pleurothallis ophioglossoides 0000000000000000000000000000000111010000100000000000 Pleurothallis acutissima 1000000000000000000000000000000000000010000000000000 Pleurothallis appendiculata 0001010000000000000000000000000000000000000000000000 Pleurothallis aristata 1001111000000000000000000000000111100000000000000000 Pleurothallis brighamii 0100111000000000000000000000000000000000000000000000 Pleurothallis consimilis 1000000000000000000000000000000000000010000000000000 Pleurothallis corniculata 1000111000000000000000000000000000000000000000000000 Pleurothallis discoidea 1000000000000000000000000000000000000010100000000000 Pleurothallis domingensis 0001111000000000000000000000000100000000000000000000 Pleurothallis dussii 0000000000000000000000000000000110000000000000000000 Pleurothallis erosa 0000011000000000000000000000000000000000000000000000 Pleurothallis gelida 0011111000000000000000000000000000000000000000000000 Pleurothallis grobyi 1100011000000000000000000000000000000000000000000000 Pleurothallis helenae 0000110000000000000000000000000000000000000000000000 Pleurothallis hymenantha 1000001000000000000000000000000000000000000000000000

Ó Pleurothallis imraei 1000010000000000000000000000000111000000000000000000

lcwl cec t 2001, Ltd Science Blackwell Pleurothallis lanceola 0000110000000000000000000000000010000000000000000000 Pleurothallis laxa 0000110000000000000000000000000000000000000000000000 Pleurothallis mazei 0000010000000000000000000000000100000000000000000000 Pleurothallis miguelii 0000010000000000000000000000000000001010000000000000 Pleurothallis monophylla 1000100000000000000000000000000000000000000000000000 Pleurothallis mornicola 0000010000000000000000000000000000001010000000000000 Pleurothallis murex 0000011000000000000000000000000000000000000000000000 Pleurothallis nummularia 0000101000000000000000000000000000000000000000000000 Pleurothallis oblongifolia 0001111000000000000000000000000010000000000000000000 Pleurothallis obovata 1001011000000000000000000000000000000000000000000000 Pleurothallis polygonoides 1000000000000000000000000000000000000010000000000000 ora fBiogeography of Journal Pleurothallis pruinosa 1001111000000000000000000000000111101010000000000000 Pleurothallis pubescens 1001010000000000000000000000000000000000000000000000 Pleurothallis racemi¯ora 0001111000000000000000000000000000000000000000000000 Pleurothallis revoluta 1000000000000000000000000000000000000000100000000000 Pleurothallis ruscifolia 1001111000000000000000000000000111011010100000000000 Pleurothallis sclerophylla 1000000000000000000000000000000000000000100000000000 Pleurothallis sertularioides 0100101000000000000000000000000000000000000000000000 Pleurothallis sicaria 1000000000000000000000000000000000000010000000000000 ,

28 Pleurothallis testaefolia 0000111000000000000000000000000001100000000000000000

775±794 , Pleurothallis tribuloides 0100111000000000000000000000000000000000000000000000 Pleurothallis trichophora 0000011000000000000000000000000000000000000000000000 Pleurothallis trichostata 0001010000000000000000000000000000000000000000000000 Pleurothallis velaticaulis 0000000000000000000000000000000000000010100000000000 Ó Appendix 1 continued lcwl cec t 2001, Ltd Science Blackwell lemy Âe ao Ëc

Species name/area name Guianas Yucatan Florida Puerto Rico Jamaica Hispaniola Cuba Isla Juventud Mona Grand Cayman Little Cayman Cayman Brac Turks Inaguas Crooked Long Salvador Exumas Cat Providence Andros Abacos St Martin Saba St Barth St Kitts Nevis St Eustatius Antigua Barbuda Montserrat Guadeloupe Dominica Martinique St Lucia St Vincent Grenada Barbados Trinidad Tobago Margarita Vieques Culebra Anegada Virgin Gorda Tortola St Thomas St John St Croix Aruba Cura Bonaire Pleurothallis wilsonii 0001111000000000000000000000000100000000000000000000 Pogonia grandi¯ora 1000000000000000000000000000000000000010000000000000 Pogonia tenuis 1000000000000000000000000000000000000010000000000000

ora fBiogeography of Journal Polystachya concreta 1011111100100000000111000101001111111010100000100000 Polystachya foliosa 1101111000000000000011000000001000001010000010100010 Ponthieva brittoniae 0010001000000000000110000000000000000000000000000000 Ponthieva diptera 1000111000000000000000000000000000000000000000000000 Ponthieva harrisii 0000110000000000000000000000000000000000000000000000 Ponthieva pauci¯ora 0000111000000000000000000000000000000000000000000000 Ponthieva petiolata 0000011000000000000000000100000111111000000000000000 Ponthieva racemosa 0111111000000000000000000000000000000000000001110000

, Ponthieva ventricosa 0001111000000000000000000000000000000000000000000000 28 Prescottia oligantha 1011111000000000000110000000000111001000000001110000 775±794 , Prescottia stachyodes 1101111000000000000000000001010111010010000000010000 Prosthechea boothiana 0110011101110000000011000000000000000000000000000000 Prosthechea cochleata 1111111101110000000111000000000110000000110000001000 793 distributions orchid of analysis parsimony a on based Antilles the of Biogeography Prosthechea fragans 0000111000000000000000000000000011011011100000000000 Prosthechea pygmaea 1111111000000000000000000000000010000010000000000000 Prosthechea vespa 1000011000000000000000000000000000000010000000000000 Pseudocentrum minus 0000110000000000000000000000000000000000000000000000 Psilochilus macrophyllus 1001111000000000000000000000000110111010000000000000 Psychilis kraenzlinii 0001000000000000000000110000000000000000000000000000 Psychilis macconnelliae 0001000000000000000000000000000000000000011111111000 Psygmorchis pusilla 1100000000000000000000000000000000010010000000000000 Reichenbachanthus re¯exus 1000011000000000000000000000000000001010000000000000 Rodriguezia lanceolata 1000000000000000000000000000000000010011000000000000 Sarcoglottis sceptrodes 0100000000000000100000000000000000000000000000000000 Scaphyglottis fusiformis 1000000000000000000000000000000000000010000000000000 Scaphyglottis modesta 1001011000000000000000000000000110011010000000000000 Scaphyglottis prolifera 1100000000000000000000000000000000000000000000000000 Schiedeella amesiana 0000011000000000000000000000000000000000000000000000 Schomburgkia ®mbriata 1000000000000000000000000000000000000010000000000000 Schomburgkia humboldtii 0000000000000000000000000000000000000000000000000011 Schomburgkia lysonsii 0000101000000000000000000000000000000000000000000000 Sobralia fragrans 1100000000000000000000000000000000000000000000000000 Spiranthes torta 0111111100000001000111000000100111101010000000100000 Stanhopea grandi¯ora 1000000000000000000000000000000000000010000000000000 Stelis muscifera 1000000000000000000000000000000000000010000000000000 Stelis perpusilli¯ora 0001010000000000000000000000000110000010000000000000 Stelis pygmaea 1001010000000000000000000000000111010000000000000000 Stelis scabrida 0000100000000000000000000010000111110000000000000000 Stelis trigoni¯orum 0000111000000000000000000000000000000000000000000000 Stenia pallida 1000000000000000000000000000000000000010000000000000 Stenorrhynchos lanceolatum 1011111100000000000111001000010000000110000000100000 Stenorrhynchos speciosum 0000111000000000000000000000000000000000000000000000 Stenorrhynchos squamulosum 0000101000000000000000000000000000000000000000000000 Tetramicra bulbosa 0000110000000000000000000000000000000000000000000000 Appendix 1 continued Ackerman D. J. and Trejo-Torres C. J. 794 lemy Âe ao Ëc

Species name/area name St Kitts Nevis St Eustatius Antigua Barbuda Montserrat Guadeloupe Dominica Martinique St Lucia St Vincent Grenada Barbados Trinidad Tobago Margarita Vieques Culebra Anegada Virgin Gorda Tortola St Thomas St John St Croix Aruba Cura Bonaire Guianas Yucatan Florida Puerto Rico Jamaica Hispaniola Cuba Isla Juventud Mona Grand Cayman Little Cayman Cayman Brac Turks Inaguas Crooked Long Salvador Exumas Cat Providence Andros Abacos St Martin Saba St Barth Tetramicra canaliculata 0001010000000000000000001001110101000110000110111000 Tetramicra eulophiae 0000011100000000000000000000000000000000000000000000 Tetramicra parvi¯ora 0000110000000000000000000000000000000000000000000000 Tetramicra urbaniana 0000001000000000000110000000000000000000000000000000 Tolumnia bahamense 0010000000000000000011000000000000000000000000000000 Tolumnia calochila 0000011001000000000000000000000000000000000000000000 Tolumnia gauntletii 0000100000000000000110000000000000000000000000000000 Tolumnia lyrata 0000001100000000000000000000000000000000000000000000 Tolumnia prionochila 0001000000000000000000000000000000000000011111110000 Tolumnia sasseri 0000000000000000000011000000000000000000000000000000 Tolumnia urophyllum 0000010000000000000000001011111110000000000000000000 Tolumnia variegata 0001011011000000000000000000000000000000011011111000 Trichopilia fragans 0000011000000000000000000000000000000000000000000000 Trichosalpinx ciliaris 1100000000000000000000000000000000000000000000000000 Trichosalpinx dura 1001111000000000000000000000000101001011100000000000 Trigonidium egertonianum 1100000000000000000000000000000000000000000000000000 Triphora amazonica 1011010000000000000000000000000000000010000000000000 Triphora gentianoides 0110111000000000000000000000000000000000000000000000 Triphora hassleriana 0001010000000000000000000000000000000000000000000000 Triphora surinamensis 1001110000000000000000000000000110000010000000000000 Triphora yucatanensis 0110000000000000000000000000000000000000000000000000 Trizeuxis falcata 1000000000000000000000000000000000000010000000000000

Ó Tropidia polystachya 0110111001000000000000000000000000000000000000000000

lcwl cec t 2001, Ltd Science Blackwell Wullschlaegellia aphylla 0000101000000000000000000000000000000010000000000000 Wullschlaegellia calcarata 1001010000000000000000000000000110000000000000000000 Xylobium colleyi 1000000000000000000000000000000000000010000000000000 Xylobium foveatum 1000100000000000000000000000000000000000000000000000 Xylobium palmifolium 1000111000000000000000000000000010011010000000000000 Zootrophion atropurpureum 0000111000000000000000000000000000000000000000000000 ora fBiogeography of Journal , 28 775±794 ,