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.
INTRODUCTION
The Antilles, or West Indies, have been frequently studied by
biogeographers. These islands make up one of the largest
tropical archipelagos in the world, second only to the islands
Correspondence: J. Carlos Trejo-Torres, Department of Biology, University of
Puerto Rico-RõÂo Piedras, PO BOX 23360, San Juan, Puerto Rico 00931-3360.
1 E-mail: [email protected]
Ó 2001 Blackwell Science Ltd
between Asia and Australia. The Antilles are also signi®cant
because they represent one of the two connections, by way of
an island chain, between two major biogeographical realms:
the Neartic and the Neotropic. The majority of studies on
Antillean biogeography concern the distributional patterns
or the cladistic biogeography of animals, especially vertebrates and insects (e.g. Liebherr, 1988; Woods, 1989;
Page & Lydeard, 1994; Hedges, 1996). Surprisingly, there
are only a few publications on the phytogeography of this
776 J. C. Trejo-Torres and J. D. Ackerman
region (Howard, 1973; Samek, 1988; Adams, 1989; Lavin,
1993; Borhidi, 1996), although the ¯oristics of the area have
been actively studied for decades (Zanoni, 1986; Liogier,
1996).
The Caribbean region has been divided phytogeographically in three subregions by Samek (1988). These main divisions 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 considered
part of the Colombian±Venezuelan subregion. Borhidi
(1996) joins both continental subregions, from Mexico to
Venezuela, into a single one. While these authors consider
south Florida as part of the Antillean subregion, Gentry
(1982) included only the Florida Keys in the Antillean region.
Distinguishing biotic regions, or de®ning biological
boundaries, has been among the major concerns of biogeographers. Traditionally, this has been performed with
subjective methods (e.g. Gentry, 1982; Samek, 1988;
Borhidi, 1996). However, more objective or analytical
approaches have been developed for the analysis of distributional 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 an
alternative method based on a parsimonious analysis of taxa
distributions (Rosen & Smith, 1988). This method represents a direct way to search for the biogeographical af®nities
among areas (Connor, 1988; Vargas, 1991), for the detection of areas of endemism (Morrone, 1994a; Cardoso da
2 Silva & Oren, 1996; Bates et al., 1998), and for the
delimitation of biological boundaries (Posadas, 1996;
Posadas et al., 1997; Morrone, 1998). The parsimony
analysis presented here is a tool for searching the most
parsimonious arrangement of shared species among areas, as
a means of revealing the biogeographical af®nities in a
hierarchical pattern (Rosen & Smith, 1988; Brady, 1994).
The analysis presented here was originally called parsimony analysis of endemicity (PAE), and was suggested by
3 Rosen (1985, cited in Rosen & Smith, 1988) and developed
by Rosen & Smith (1988). It was also independently
suggested by Legendre (1986) and Connor (1988). Since
then it has been employed in the study of extant taxa of
New Zealand (Craw, 1988), Australia (Cracraft, 1991),
Southeast Africa (Morrone, 1994a), the Patagonia
(Posadas, 1996), the Andes (Morrone, 1994b; Posadas
et al., 1997), the Amazonia (Cardoso da Silva & Oren,
1996; Bates et al., 1998), the Austral region (Craw, 1989;
Morrone, 1998; Glasby & Alvarez, 1999), Mexico (Luna
et al., 1999), and the entire world (Conran, 1995). The
units of comparison that have been used are sites, quadrants or sections of regions, biogeographical areas, or
natural geographical areas (e.g. islands, continents, ocean
basins). We use entire islands or groups of them, as the
units of study. While the method has been mainly used for
discovering the historical relationships among areas, we
give a static or non-historical interpretation to the patterns
obtained (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 (including
single-site endemics), and takes into account shared species
whether endemic or not to the Antilles or adjacent areas.
We substitute the term distribution for endemicity to avoid
confusion.
Our goal here is to distinguish biogeographical patterns in
the Antilles based on one plant family, Orchidaceae. This
group of plants, of which there are approximately 700
species in the Antilles, is relatively well known taxonomically and geographically because there are relatively recent
orchid treatments for the Cayman Islands (Proctor, 1984,
1996), Puerto Rico (Ackerman, 1995) and the Bahamas
(Sauleda & Adams, 1982; Sauleda, 1992). Furthermore, a
treatment for the Greater Antilles is underway (Ackerman,
1997; in press). The homogeneous wind-dispersal mechanism (except in bird-dispersed Vanilla) of the dust-seeds
among the species of this family, makes orchids a good focal
group for the study of distribution. Assuming that most
orchids have the same dispersal capacity, their distributional
patterns may be explained in terms of other ecological,
geographical and historical factors.
The questions we address are: (1) what are the phytogeographical relationships among the Antillean islands
based on shared orchid species? (2) are these relationships
affected when neighbouring continental regions are considered? (3) what is the phytogeographical regionalization of
the Antilles based on orchids, and (4) how do the distributional patterns of orchids match with geography, geology,
physiography and ecology of the region? Apart from the
biogeographical analysis per se, we discuss the use of
parsimony analysis of naturally de®ned areas (i.e. islands)
with extremely dissimilar number of species, and the use of
single-site species to look at ®ner degrees of biogeographical
differentiation.
METHODS
A total of ®fty-two areas (Table 1) were included in the
study. Data on the distribution of species were taken from
literature and from unpublished sources (Table 2). Synonymies and valid species names were standardized mainly
according to J.D. Ackerman (unpublished data).
A presence/absence matrix of the 863 orchid species
reported for the studied areas was constructed in MacClade
3.01 (Maddison & Maddison, 1992). Using this program,
areas were entered in the place of taxa, while taxa were
entered in the place of characters. In the matrix, presence
was indicated with a `1' and absence with a `0'. Once we
constructed the matrix, we ran analyses of parsimony using
PAUP 4 (Swofford, 2000). A hypothetical outgroup area
with all 0s (no species) were used in the analyses to root the
trees. General heuristic searches were carried out to look for
the most parsimonious trees, which indicate the ¯oristic
af®nities among studied areas. We obtained consensus trees
(Strict, Majority Rule and Adams) when more than one
equally parsimonious tree were found. A description and 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
included in the study. The Bahamas islands are combined following
Correll & Correll (1982); names in italics are the ones used in ®gures
and appendix
Greater Antilles
Cuba
Hispaniola
(Dominican Republic & Haiti)
Jamaica
Puerto Rico
Isla de la Juventud
Mona
Cayman Brac
Grand Cayman
Little Cayman
Lesser Antilles
Antigua
Barbados
Barbuda
Dominica
Grenada
Guadeloupe
Martinique
Montserrat
Nevis
Saba
St BartheÂlemy
St Eustatius
St Kitts
St Lucia
St Martin
St Vincent
Virgin Islands
Anegada
Culebra
St John
St Thomas
St Croix
Tortola
Vieques
Virgin Gorda
Bahama Archipelago
Abacos ± Grand Bahama
Andros ± Biminis
Cat
Crooked ± Mayaguana
Exumas
Inaguas
Long ± Ragged Island Range
New Providence ± Eleutheras
San Salvador ± Rum Cay
Turks and Caicos
Trinidad bank
Margarita
Tobago
Trinidad
Continental regions
Florida (North America)
Yucatan (Central America)
Guianas: Guyana, Surinam and
French Guyana (South America)
Southern Dutch Antilles
Aruba
Bonaire
CuracËao
information in terms of shared species af®nities. We only
used these single-site (island or area) species for speci®c
¯oristic comparisons. Also, Vanilla species were not considered at all because they are animal-dispersed, an anomaly in
the orchid family.
Area relationships derived from PAD may be altered
depending on the inclusion or exclusion of different areas
just as is the case when using different ingroup/outgroup
taxa in cladistic studies. To test the robustness of our data,
we conducted independent analyses of different subsets of
areas.
RESULTS
An analysis of all areas using the 356 shared orchid species
produced 216 equally parsimonious trees. The three consensus trees obtained from them produced entirely congruent
groupings (Fig. 1). The Strict consensus tree shows the
groups found in all the most parsimonious trees. The
Majority Rule tree shows the groups found in more than
50% of those trees. The Adams tree gives the highest
possible resolution for data distribution.
Four main clusters of areas can be identi®ed in the
consensus trees (Fig. 1): (1) Greater Antilles/Guianas±Trinidad/larger Lesser Antilles/smaller Lesser Antilles/Virgin
Islands/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, the
polytomic base of the trees means that relationships among
them are not resolved. The arrangement of areas within the
groupings is highly in¯uenced by species numbers.
DISCUSSION
Table 2 Primary data sources for the orchid species presence in
the studied Caribbean regions
Area
Data source
Bahama Archipelago
Cayman Islands
Florida
Greater Antilles
Guianas
Isla de la Juventud
Lesser Antilles
Sauleda & Adams (1982), Sauleda (1992)
Proctor (1984, 1996)
Wunderlin et al. (1996)
Ackerman, J. D. (unpublished data)
Boggan et al. (1997)
Jennings (1917), Sauget & Barbier (1946)
Garay & Sweet (1974), Nir, M.
(unpublished)
Hoyos (1985)
van Proosdij, A. (unpublished)
Garay & Sweet (1974); Kenny (1988)
Carnevali, G. (unpublished)
Margarita
Southern Dutch Antilles
Trinidad and Tobago
Yucatan
found in Rosen & Smith (1988), Vargas (1991) and Rosen
(1988, 1992).
Only the informative species were taken into account in
the analyses (Appendix 1). The uninformative species are
those found in only one area; therefore, they give no
Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794
Groupings of areas vs. physical factors
We found two kinds of grouping patterns in the Antilles
based on orchid species distributions. One pattern aggregates islands that belong to single geological units while the
other pattern groups areas with different geological histories
(Fig. 2).
In the ®rst pattern, the groupings are easy to understand
as they correspond to islands belonging to de®ned geological units. The Virgin Islands holds together as a group
(except for Anegada). The islands of the Virgin Islands
Bank, which excludes St Croix, were once part of a larger
volcanic island that included Puerto Rico. They are old
islands, about 105 Ma (Donnelly, 1988) that separated
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 Bahamas belong to a single geological platform, part of the
North American plate. They are low-lying, sedimentary
limestone areas. Many of the Bahamas were interconnected
in the past (Williams, 1989). Regarding the Cayman
Islands, which are located south of Cuba on the Cayman
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 sedimentary substrate to 36 Ma; no connection between these
islands and other lands occurred in the past (Proctor,
1984). Lastly, Aruba, Bonaire and CuracËao, with volcanic
and sedimentary substrates, are geologically related and
part of the South Caribbean Island Chain (Bellizzia &
Dengo, 1990). They are also small, low-elevation islands
(to 372 m) with volcanic and sedimentary substrates, and
have dry climates. In addition to their common origin, each
one of these island aggregates has a common geomorphology and physiography with relatively homogeneous ecological conditions and low species richness. The anomalous
¯oristic connection of St Croix with the Virgin Islands
group is understandable if we look at the physiography
rather than the geology. St Croix has an independent origin
from the Virgin Islands Bank; nevertheless, they are
physiographically similar.
The second pattern is represented by areas differing in
geological origin and that are separated by wide geographical distances among themselves. One example is represented
by the Bahamas/Isla de la Juventud/Mona/Anegada group.
As mentioned above, the Bahama Archipelago is composed
of low-lying, sedimentary limestone islands located on the
North American plate. The Isla de la Juventud, which lies on
the western part of the Cuban Bank, is almost entirely lowlying, part calcareous and part metamorphic (CRAC, 1978).
Mona is a small, ¯at, calcareous island between two of the
Greater Antilles (Puerto Rico and Hispaniola). Lastly,
Anegada is part of the Virgin Islands Bank; however, unlike
the other Virgin Islands, it is ¯at, low-lying and calcareous
(D'Arcy, 1971). In spite of the separate geological
origin, these islands have a similar geomorphology and
physiography. The other example of the second pattern is
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 distributions. See Fig. 1 for detailed patterns of
af®nity and single-island names. Dotted areas: continental areas; short-dashed lines:
island groups; long-dashed lines: relationships with continental areas; question marks:
unresolved group or controversial af®nity;
arrow: presumed af®liation. Island or grouping names: A ˆ Anegada, B ˆ Barbados,
C ˆ Cuba, CI ˆ Cayman Islands, H ˆ Hispaniola, 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 Antilles, T ˆ Trinidad, VI ˆ Virgin Islands.
Eastern Antilles/Virgin Islands/Margarita±Tobago/other
smaller Lesser Antilles group. The origin of the Greater
Antilles, whether continental or oceanic, is under debate
(Rosen, 1985; Iturralde-Vinent & MacPhee, 1999); however, 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 island
arc about 43 Ma, and later expanded 11 Ma to form the
north-western Lesser Antilles, from Dominica to Saba
(Coney, 1982; Maury et al., 1990). Finally, the islands of
Trinidad, Margarita and Tobago, belong to the Caribbean
Mountain System that is part of the continental South
American plate (Bellizzia & Dengo, 1990). There has been
no connection between these islands and the Lesser Antilles
(Donnelly, 1988). Trinidad lost connection with South
America some 5 Ma (Persad, 1985). Notwithstanding the
differences in geology and age, most of the islands of this
grouping are volcanic, mountainous, have a variety of
climates and natural communities, and complex species
assemblages.
Orchid distribution patterns also seem to be in¯uenced by
their high vagility. On a regional scale (the Antilles) cohesive
clusters of islands spread over hundreds to thousands of
kilometres (e.g. Bahamas±Anegada or the Greater Antilles±
Eastern Antilles) re¯ect the high potential for orchid dispersal, a capacity that may override the effects of geological
history in de®ning the af®nities among those islands.
In general, distribution patterns of orchids are explainable
according to the physical features of the islands. The similar
geomorphology and/or physiography of these islands presumably results in similar ecological conditions that consequently produce similar assemblages of species. However,
geology clearly plays an important role in de®ning biogeographical af®nities at a more restricted geographical scale.
We do not know to what extent the biogeographical patterns
Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794
are affected by other factors such as prevailing winds and
hurricane tracks.
Analysis of areas using different island data sets
Different analyses using subsets of areas (using the informative
species for each data set) show patterns compatible with the
general analysis of all areas. Nevertheless, they do add
information about the relationships among some groupings
and areas. The following observations are based mainly on the
Majority Rule consensus trees (trees not presented).
1 When we analyse only the islands, excluding the continental areas of Florida, the Guianas and the Yucatan, the
Bahamas split into two branches. The north-western
areas (Andros/Abacos/Providence) form a group with the
Isla de la Juventud as basal. The Cayman Islands become
a sister clade of this Bahama group. Meanwhile, the rest
of the Bahama islands form another group with Mona
still as basal.
2 If we eliminate the Isla de la Juventud (and the continental areas), the Bahamas are not divided into two groups
as before and form a monocladic group. Mona and
Anegada remain as basal areas.
3 When we eliminate the Bahamas (and the continental
areas), the Isla de la Juventud interestingly forms a pair
with Margarita, and together a sister group of the
Cayman Islands, all of them belong to an unresolved
clade. Meanwhile, Mona is left unresolved.
4 An analysis of the north-western areas, that is, the
Yucatan and Florida, the Bahamas, the Greater Antilles,
the Isla de la Juventud, the Cayman Islands, Mona, and
the Virgin Islands, shows the Isla de la Juventud in an intermediate position between the Yucatan/Florida/Greater
Antilles and the Bahamas/Mona/Virgin Islands. The
Cayman Islands remain monocladic and again unresolved.
780 J. C. Trejo-Torres and J. D. Ackerman
5 We also evaluated distributions of just the Greater
Antillean area, which includes the Isla de la Juventud,
the Cayman Islands, the Virgin Islands and Mona. This
analysis reveals the Virgin Islands and Mona as basal of
the Greater Antilles in the order mentioned. Meanwhile
the Isla de la Juventud joins the Cayman Islands forming
an unresolved clade.
6 An analysis of our Eastern Antilles, from the Virgin
Islands to Margarita, links Trinidad as basal to the larger
Lesser Antilles, particularly as the sister area of neighbouring Grenada. Tobago and Margarita appear as sister
areas inserted within the Eastern Antilles group. The
Virgin Islands remain monocladic.
7 After merging islands belonging to banks (de®ned by the
200 m bathimetric line: Trinidad + Tobago +Margarita,
Virgin Islands except St Croix, Antigua + Barbuda,
St Kitts + Nevis + St BartheÂlemy), the Trinidad bank
remains as a sister and basal group of the Greater Antilles
(as Trinidad alone).
Points (1), (3) and (5) denote a close af®nity among the
Isla de la Juventud, the Cayman Islands, and the Bahamas.
Relationships among the groupings using exclusive
and endemic species
The parsimony analysis produced consistent assemblages of
areas. Nevertheless, the relationships among some of the
groupings are not resolved. We attempt to understand these
unde®ned af®nities by subjectively examining the number of
endemic species, the number of exclusive species (single-site,
non-endemic species), and the presence of species of restricted distribution of every island or group. Are all the
unresolved clusters of islands in the consensus trees, independent biogeographical provinces?
The Eastern Antilles, from the Virgin Islands to Margarita
constitute a distinct branch with several minor subgroups.
First, the larger Lesser Antilles from Guadeloupe to St Lucia is
the most de®ned group, with seven endemic species [Elleanthus dussii Cogniaux, Epidendrum discoidale Lindley, Maxillaria guadalupensis Cogniaux, Pleurothallis dussii
Cogniaux, P. ophioglossoides (Jacquin) Garay, Pseudocentrum guadalupense Cogniaux, Stelis dussii Cogniaux]. Secondly, the smaller Lesser Antilles (islands north of
Guadeloupe from Montserrat to St Martin) is a `group' that
splits along the Eastern Antilles branch (Figs 1 and 2). Some
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
Lesser Antilles `group' is not cast in stone because basal
species-poor areas are prone to changing af®nities. Thirdly,
the Virgin Islands, except Anegada, constitute a monocladic
but weakly differentiated (no endemic species) group which is
®rmly allied with the Lesser Antilles. Nevertheless, the Virgin
Islands have two species that they share exclusively with
neighbouring Puerto Rico [Psychilis macconnelliae Sauleda,
Tolumnia prionochila (KraÈnzlin) Braem]. This connection
was 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 Antilles
branch. The fact that these island groups appear as basal and
pectinate with respect to the Greater Antilles, indicates a close
af®nity among the islands of our Island Arc group.
The Cayman Islands are always monocladic and usually
occur as an independent clade. They have three endemic
species [Dendrophylax fawcettii Rolfe, Encyclia kingsii
(C. D. Adams) Nir, Myrmecophila albopurpurea (Strachan
ex Fawcett) Ackerman; recent morphological and molecular
data fail to distinguish T. caymanense (Moir) Braem from
T. variegata (Swartz) Braem]. Among the species of restricted
distribution two are shared with the Greater Antilles, Florida
and 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 another
is shared only with Cuba [Myrmecophilla thomsoniana
(Reichenbach f.) Rolfe]. These species in common suggest a
close af®nity among the Cayman Islands, the Greater
Antilles and the peninsulas of Yucatan and Florida.
The Bahamas are also an independent clade in the general
analysis. They have ®ve endemic species [Encyclia caicensis
(Sauleda & Adams), E. fehlingii (Sauleda) Sauleda &
Adams, E. gracilis (Lindley) Schlechter, E. inaguensis Nash
ex Britton & Millspaugh, T. sasseri (Moir) Braem]. Of the
species with the most restricted distribution, there is only
one exclusively shared with Florida [T. bahamense (Nash ex
Britton & Millspaugh) Braem] while there are seven shared
exclusively with the Greater Antilles [Broughtonia lindenii
Lindley, Domingoa haematochila (Reichenbach f.) Carabia,
Encyclia fucata (Lindley) Britton & Millspaugh, E. plicata
(Lindley) Schlechter, Tetramicra urbaniana Cogniaux,
T. gauntletii (Withner & Jesup) Braem, T. lucayana (Nash
ex Britton & Millspaugh) Braem]. Thus, the relationships
among the Bahamas and the Greater Antilles seems to be
considerable.
Lastly, Aruba, Bonaire and CuracËao form another independent group. These islands have only three species
reported, none of which is endemic. One occurs on different
continental and insular areas [Polystachya foliosa (Hooker)
Reichenbach f.], one held in common with Central and
South America [Brassavola nodosa (Linnaeus) Lindley], and
one shared only with Venezuela [Myrmecophila humboldtii
(Reichenbach f.) Rolfe]. Thus, the ABC islands show no
special af®nity to the rest of the Antilles.
Relationships with continental areas
The inclusion of the Guianas as a South American continental area was thought as a heuristic exercise. First, the
Guianas is a politically de®ned area rather than a biogeographical region. Secondly, the Guianas is likely outside the
Caribbean ¯oristic region and part of the Amazonian region
(Gentry, 1982). If we were to include other South American
regions in this study, no doubt the Guianas would shift
af®nities dramatically. The Guianas form a sister group to
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
been put forth for the ¯ora in general by Samek, 1988;
Borhidi, 1996).
The Yucatan and Florida form a sister pair of the northwestern Bahamas/Isla de la Juventud group. All these are
sedimentary, calcareous, low-lying and ¯at areas. Samek
(1988) and Borhidi (1996) de®ned South Florida together
with the Bahamas as one province. On the other hand,
Gentry (1982) excluded Florida (except for the Keys) from
his Caribbean (insular) region. We consider Florida and
the Yucatan as provinces not belonging to the Antillean
subregion, which are supposedly part of the Continental
subregion of the Caribbean.
To de®ne more clearly the relationships among the insular
Antilles and their neighbouring continental areas, more
thorough studies are necessary. Also, the inclusion of other
continental areas, working as distant sister areas in the
analyses, would be useful. For example, a sister continental
area for the Yucatan biogeographical province (DuraÂn et al.,
1998) could be a province in central-western Mexico. Both
outside provinces belong to the same ¯oristic MexicanCentral American Caribbean subregion (sensu Samek, 1988).
Islands with unresolved af®nity
The unresolved islands or island groupings found in some
consensus trees could be explained in a number of ways.
First, it is possible that an incomplete record of the species
for these islands could be hampering the resolution of their
af®nities. As we improved the species record for the smaller
Lesser Antillean islands they became progressively resolved.
We expect this trend to continue. Alternatively, the record
may be complete and these unresolved areas could be the
result of: (1) a lower number of species which would prevent
their af®liation with other groups, (2) an equally strong
af®nity with more than one of the other groups, and (3) a
non-hierarchical structure in the data (Ronquist, 1997;
Glasby & Alvarez, 1999). One way an unresolved area or
group could be considered distinctive would be if it had a
well-differentiated ¯ora (i.e. endemic or exclusive species). It
is likely a combination of these factors that play a role in
unresolved cases.
PAD, species richness and species±area curves
The position of areas in tree branches is in¯uenced by species
numbers. This pattern is not evident for the Antillean
archipelago as a whole and only becomes obvious within the
major island assemblages. As we move from the tip to the
base of the tree branches, species numbers tend to diminish.
This apparent numerical artefact has been stated as one of
the main problems resulting from the use of areas with
highly dissimilar number of species, and a comparable
number of taxa per site has been recommended as a prerequisite for PAE analyses (Rosen, 1988, 1992; Rosen &
Smith, 1988; Vargas, 1991; Conran, 1995). What our study
shows is that even with very few species the af®liation of an
island to a speci®c group occurs when the taxa permit it.
Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794
Every main cluster has its own species-poor islands. We
interpret 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 its
af®liation, such an area turns out to be unresolved or
unstable, being located at the base of the whole tree or at the
base of different branches, respectively. In other words,
either a high or a low number of species can de®ne
membership patterns among areas. Limiting the analysis to
areas with a similar number of species would unnecessarily
restrict the use of this methodology for the Caribbean and
many other regions.
Differences in species numbers within our island groups
can be converted into species±area curves. We have found
that the geologically and physiographically diverse montane
islands have a different species±area curve than that for lowlying islands, which are basically calcareous and physiographically homogeneous (Ackerman et al., submitted). The
calcareous group, comprised of nineteen islands (including
the Cayman Islands), has a z-value of 0.25 (however, r ˆ 0.4,
P ˆ 0.07), a typical value for islands (Rosenzweig, 1995).
Meanwhile, the montane island group, composed of twentysix 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
value found by Adler (1992, cited by Rosenzweig, 1995).
Moreover, if a species±area curve for this montane group is
calculated with only the endemic species (using only nine
islands that have single-island endemics), z ˆ 0.68 (r ˆ 0.88,
P ˆ 0.0019) suggesting that habitat diversity is important for
autochthonous speciation. The two z-values for the montane
group seem to indicate that the Island Arc group behaves as a
cluster of provinces (Rosenzweig, 1995) as has been
proposed by Borhidi & MunÄiz (1986). In contrast, the
endemic species±area curve for the relatively homogeneous
calcareous group was not run because there are only three
islands (island banks indeed) with endemic species.
Phytogeographical regionalization of the Antilles
The Antilles is among the principal phytogeographical
regions of the world (Gentry, 1982). It has been included
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
phytogeographic sectorization of Samek (1988) and Borhidi
(1996) consider every one of the Greater Antilles as
provinces. Major geographical aggregations of islands (i.e.
the Bahamas, the Lesser Antilles) are also de®ned as
provinces, while minor groups of islands (e.g. the Virgin
Islands, the Cayman Islands) are integrated with some of the
big islands. Apart from the recognition of these kind of
divisions, we also examined af®nities among and within
them 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) is
suf®ciently differentiated to be recognized as a separate
province of the Antilles.
782 J. C. Trejo-Torres and J. D. Ackerman
2 The Lesser Antilles were taken as a single phytogeographic province by Samek (1988) and Borhidi (1996).
Their orchid ¯ora is suf®ciently distinctive, supporting
this view. Our analysis places them as part of the Eastern
Antilles group and at the same time subordinate to the
Greater Antillean main group.
3 The Virgin Islands were integrated as part of Puerto Rico
by Samek (1988) and Borhidi (1996). However, our study
places them rather close to the Lesser Antilles, and
altogether, as subordinate of the Greater Antilles.
4 The Cayman Islands were considered part of Jamaica by
Samek (1988) but as part of Cuba according to Borhidi
(1996). Our analysis suggests that these islands have
complex af®nities with the Greater Antilles, the Yucatan
and Florida continental areas, and also with the Bahamas.
5 The Bahama archipelago was one of the Caribbean
provinces distinguished by Samek (1988), Borhidi (1996),
and Gentry (1982), and this is also supported by our
orchid data. Moreover, our study relates the Isla de la
Juventud, Mona and Anegada (geographically and geologically part of the Greater Antilles) as subordinates of
the Bahamian Province. The Isla de la Juventud shows a
strong af®nity with the north-western Bahamas, while
Mona and Anegada are small islands with close af®nity to
the rest of the Bahamas.
6 Trinidad, Tobago and Margarita were considered part of
the Venezuela±Colombia Caribbean subregion by Samek
(1988) and Borhidi (1996), while no clear af®liation for
Trinidad and Tobago can be obtained from Gentry
(1982). Our data support a closer af®nity of Trinidad
with the Greater Antilles instead of the neighbouring
Lesser Antilles. Based on its differentiated orchid ¯ora
(approximately thirteen endemic species) we consider
Trinidad 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 are
two potential problems with these islands. First, we
consider that the species record for these islands is not
well updated, especially for Trinidad and Tobago whose
most recent complete orchid ¯ora was published by
Schultes (1960). Secondly, the Venezuelan Guyana may
be a more appropriate neighbouring continental area,
instead of our Guianas area. If this is true, the islands of
the Trinidad bank may be more related to the continent
than to the rest of the Antillean islands, as suggested by
bat distribution patterns (Trejo-Torres & Rivera, unpublished data) and by geological history.
7 Aruba, Bonaire and CuracËao seem to be independent of
the Antillean subregion and apparently part of the
Continental Caribbean subregion.
Perspectives of PAD
The pattern we found for orchids is not necessarily the same
for the general ¯ora or for the entire biota of the region. We
expect that phytogeographic relationships revealed by
orchids might be similar to that of other highly vagile groups
of organisms such as wind-dispersed plants, ¯ying insects,
¯ying birds and bats. In this case, area af®nities appear to be
de®ned by physical factors, such as geomorphology, physiography, climate, geography and island area. At the same
time, it is plausible to predict that less vagile organisms, such
as non-¯ying vertebrates and freshwater ®shes, will de®ne
different patterns of area relationships, perhaps more in¯uenced by geological and geographical history of areas.
The PAE/PAD is an alternative to the use of multivariate
phenetic methods for classi®cation of biotas (Rosen, 1988;
Vargas, 1991) and for comparing species assemblages. In
fact, giving a static interpretation of area relationships, this
is a method equivalent to multivariate ones (Rosen, 1988,
1992). Some important differences between PAE/PAD and
multivariate methods are: (1) The PAE/PAD uses parsimony
algorithms instead of similarity indexes. This means that the
hierarchy selected in this manner will be the best supported
by evidence, for it maximizes congruence between data and
hierarchical patterns (Brady, 1994). (2) The PAE/PAD uses
characters (taxa) selected a priori based on their informativeness, that is, species found in all sites and single-site
species are eliminated as they give no information on the
af®nities among areas (3). The PAE/PAD provides a more
uni®ed methodology as multivariate methods use many
different similarity indices, which can also yield different
results.
The PAE/PAD can be seen as a complementary method.
While many community analyses are intended to describe
quantitative parameters, PAE/PAD is directed to describe
composition patterns, based on the identity of species
(Worthen, 1996). Also, results of PAE/PAD in the form of
trees with a hierarchical structure could be a graphic
complement to other methods which produce results that
are single numerical values, such as indexes, scores or
exponents (i.e. nestedness analysis, Patterson & Atmar,
1986). The hierarchical nature of the results does not
necessarily mean that communities are organized that way.
After all, af®nities among communities may have an
underlying reticulate structure (Ronquist, 1997). We view
PAE/PAD as only one way to represent structure in biogeographical data.
The PAE/PAD has the potential as a method for understanding ecological biogeographic problems. This is supported by the Caribbean orchids for which composition patterns
strikingly match physical and ecological factors. The use of
the method as a complement to vicariance and other historical biogeographic methods has been suggested (Cracraft,
4 1991). Because of its phenetic nature, patterns found
through PAE/PAD could be re¯ecting ecological phenomena,
rather than historical ones. Nevertheless, until now, it has
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
example, ¯oristic af®nities, biodiversity comparisons and
composition patterns can be seen as phenetic phenomena.
For the demarcation of biogeographical units PAE/PAD
could be employed before searching for speci®c patterns in
distributional data sets. This could avoid distortions from
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,
Morrone & Crisci (1995) suggested the use of panbiogeographic methods as an initial step to do other historical
biogeographic analyses.
The PAE/PAD could prioritize conservation efforts at the
regional level (Posadas, 1996) by identifying areas or islands
of high diversity. We have shown that such use can be
helpful for the Antilles, one of the world's biodiversity
hotspots (Myers et al., 2000). However, conservation
efforts often focus on unique species, particularly endemics
(island endemics in our case), but PAD uses only shared
species. The exclusive use of PAD in formulating conservation strategies can thus be misleading. This is especially true
for species-poor islands that appear subordinated to speciesrich islands. Impoverished islands may thus be falsely
interpreted as irrelevant for biodiversity conservation. For
example, the tiny island of Mona lies subordinate to other
islands of our calcareous group yet it is certainly worthy of
conservation efforts because Mona has its own set of
endemic plants and animals (including an orchid, Psychilis
monensis Sauleda).
The PAE/PAD do have problems that need to be resolved
and these are the interpretation of: (1) the unresolved status
among the main groupings of areas, (2) the area relationships when using islands as well as continental regions, (3)
the potential use of branch-lengths in the trees. Special
attention should also be given when selecting data sets for
organisms with diverse ecological characteristics (e.g. birds,
which could belong to different guilds: residents vs. migratory, volant vs. non-volant).
CONCLUSIONS
According to our orchid data, the Antillean islands constitute a phytogeographical subregion of the Caribbean. The
Lesser Antilles and the Bahamas are well-de®ned provinces.
Meanwhile, the Virgin Islands and the Cayman Islands are
poorly differentiated aggregates, perhaps to be considered as
incipient provinces. All of them are subordinated to the main
Greater Antillean group.
Within islands groupings detected by PAD ordering of
areas is highly in¯uenced by species numbers. The speciesrich islands form cohesive groupings to which other speciespoor islands unite as subordinates. These subordinate islands
are liable to changing af®nities.
Orchid species show three core zones of high diversity
in the Antilles: the Greater Antilles (Hispaniola, Cuba,
Jamaica, Puerto Rico); the larger Lesser Antilles (Guadeloupe, 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 the
Antilles are in¯uenced primarily by physical characteristics
of areas. That is, areas with common ecological features
either because of similar physiographical conditions or
common geomorphology form tight clusters, despite their
differences in geology or the long distances among them.
Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794
The highly compatible patterns obtained from different
island sets and from different consensus trees indicate that
relationships of areas obtained are well supported by the
data. 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 the
study of geographical regions based on shared species,
single-site species (the majority of which are endemics) are
indicative of ®ner degrees of biogeographical differentiation.
We suggest using both data sets to analyse biogeographical
af®nities, for it integrates differences and similarities. This
approach provides a more thorough use of ¯oristic information for the analysis of biogeographical relationships.
The parsimony methodology seems to be appropriate for
the analysis of areas, or any other natural unit, with very
dissimilar number of species. The low number of species
found in a unit can de®ne its af®liation or can produce an
unresolved ¯oristic af®nity. The lack of af®nity of an area to
any group may be either real or an artefact caused by an
incomplete species record.
The PAE/PAD has great potential as an alternative,
complementary or ®rst-step method to other multivariate
and phylogenetic methodologies in biogeography.
ACKNOWLEDGMENTS
The study was supported by NSF grants to JDA (DEB9505459, HRD-9353549, HRD-9628475). Arkelio Alicea,
Kary GarcõÂa, Eileen Bravo, and Yanet Crespo were students
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
Rosen, David Lees, Susan AragoÂn, Marta DõÂaz, and Juan
J. Morrone who had made relevant comments on the
manuscript. Timothy Johnston assisted editing the manuscript and Celene Espadas, Guillermo Bianchi, and Ana
Porzecanski shared important references. GermaÂn Carnevali,
Andre van Proosdij, Mark Nir and Hagen Stenzel provided
unpublished data.
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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
1
0
1
0
0
1
1
0
1
0
1
0
0
1
1
1
1
0
0
0
0
0
1
0
0
1
0
1
1
1
0
1
0
1
1
1
0
1
1
1
0
0
1
0
0
Aganisia pulchella
Antillanorchis gundlachii
Aspasia variegata
Basiphyllaea corallicola
Basiphyllaea sarcophylla
Batemania colleyi
Beloglottis costaricensis
Bifrenaria aurantiaca
Bifrenaria longicornis
Bletia patula
Bletia purpurea
Brachionidium parvum
Brachionidium sheringii
Brachystele guayanensis
Brassavola cucullata
Brassavola nodosa
Brassia caudata
Brassia maculata
Broughtonia domingensis
Broughtonia lindenii
Broughtonia ortgiesiana
Bulbophyllum aristatum
Bulbophyllum pachyrrhachis
Calanthe calanthoides
Calopogon tuberosus
Campylocentrum fasciola
Campylocentrum ®liforme
Campylocentrum micranthum
Campylocentrum pachyrrhizum
Campylocentrum poeppigii
Campylocentrum pygmaeum
Catasetum cristatum
Catasetum integerrimum
Catasetum macrocarpum
Caularthron bicornutum
Cochleanthes ¯abelliformis
Coelia triptera
Comparettia falcata
Coryanthes macrantha
Coryanthes speciosa
Corymborkis ¯ava
Corymborkis forcipigera
Cranichis diphylla
Cranichis muscosa
Cranichis ovata
0
0
0
0
0
0
1
0
0
0
1
0
0
1
1
0
1
1
0
0
0
1
0
0
0
1
0
1
1
1
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
Guianas
Yucatan
Species name/area name
0
0
0
1
0
0
0
0
0
1
0
1
1
0
0
0
0
0
0
0
0
0
1
0
0
1
1
1
1
0
1
0
0
0
0
1
0
1
0
0
0
1
0
1
0
0
0
0
1
0
0
0
0
0
1
1
0
0
0
0
0
1
1
0
0
0
0
1
1
0
0
1
1
1
0
0
0
0
0
0
1
1
1
0
0
1
0
1
1
0
Florida
Puerto Rico
Jamaica
0
0
0
1
0
0
1
1
0
1
1
0
0
0
0
0
1
0
0
0
0
0
1
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
1
1
0
0
0
0
1
1
0
0
0
0
0
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1
0
1
0
0
1
1
1
0
0
1
1
0
1
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Hispaniola
Cuba
Isla Juventud
Mona
0
1
0
1
0
0
1
0
0
1
1
1
1
0
0
0
1
0
1
0
0
1
1
1
0
1
1
1
1
0
0
0
0
0
0
1
0
1
0
0
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grand Cayman
Little Cayman
Cayman Brac
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Turks
Inaguas
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Crooked
Long
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Salvador
Exumas
Cat
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Providence
Andros
Abacos
St Martin
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Saba
St Barth Âelemy
St Kitts
Nevis
St Eustatius
Antigua
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
Barbuda
Montserrat
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
Guadeloupe
Dominica
Martinique
St Lucia
St Vincent
0
0
0
0
0
0
0
0
0
1
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
1
1
1
1
1
0
0
0
1
1
0
1
0
0
0
0
0
1
0
0
1
0
1
1
0
0
1
0
1
1
1
0
0
1
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grenada
Barbados
Trinidad
Tobago
Margarita
Vieques
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Culebra
Anegada
Virgin Gorda
Tortola
St Thomas
St John
St Croix
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Aruba
Cura Ëcao
Bonaire
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
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
Biogeography of the Antilles based on a parsimony analysis of orchid distributions 787
Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
1
1
0
0
1
1
1
1
0
0
0
0
0
1
1
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
Species name/area name
Cranichis ricartii
Cranichis tenui¯ora
Cranichis tenuis
Cranichis wageneri
Cryptarrhena lunata
Cyclopogon cranichoides
Cyclopogon elatus
Cyclopogon laxi¯orus
Cyclopogon miradorense
Cyrtopodium andersonii
Cyrtopodium punctatum
Dendrophylax barrettiae
Dendrophylax lindenii
Dendrophylax varius
Dichaea glauca
Dichaea graminoides
Dichaea histrio
Dichaea hookeri
Dichaea hystricina
Dichaea latifolia
Dichaea morrisii
Dichaea pendula
Dichaea picta
Dichaea rendlei
Dichaea trichocarpa
Dilomilis elata
Dilomilis montana
Domingoa haematochila
Domingoa nodosa
Elleanthus caravata
Elleanthus cephalotus
Elleanthus cordidactylus
Elleanthus dussii
Elleanthus longibracteatus
Eltroplectris calcarata
Encyclia acutifolia
Encyclia angustifolia
Encyclia brevifolia
Encyclia cordigera
Encyclia fehlingii
Encyclia fucata
Encyclia gracilis
Encyclia gravida
Encyclia inaguensis
Encyclia isochila
Encyclia kingsii
Encyclia nematocaulon
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
Guianas
Yucatan
Appendix 1 continued
1
0
1
0
0
1
1
0
1
0
1
0
0
0
0
0
0
0
1
1
0
1
0
0
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
1
1
1
1
1
0
0
0
1
0
0
1
1
0
0
0
1
1
1
0
0
1
1
1
0
0
0
1
1
0
1
1
0
0
0
0
0
0
0
1
0
1
0
0
Florida
Puerto Rico
Jamaica
0
0
0
0
0
1
1
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
0
1
1
1
1
0
1
0
1
1
1
1
0
0
1
1
1
1
0
0
1
1
1
1
1
0
1
1
0
0
1
1
1
1
0
0
1
0
1
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Hispaniola
Cuba
Isla Juventud
Mona
1
1
1
1
0
1
1
1
1
0
1
1
1
1
1
1
0
0
1
1
1
1
0
0
1
0
1
1
1
0
1
1
0
0
1
1
1
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
Grand Cayman
Little Cayman
Cayman Brac
0
0
0
0
0
1
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
Turks
Inaguas
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
Crooked
Long
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
Salvador
Exumas
Cat
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Providence
Andros
Abacos
St Martin
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Saba
St Barth Âelemy
St Kitts
Nevis
St Eustatius
Antigua
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Barbuda
Montserrat
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
1
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Guadeloupe
Dominica
Martinique
St Lucia
St Vincent
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
1
1
0
1
1
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
1
1
0
0
0
0
1
0
1
1
0
1
0
1
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grenada
Barbados
Trinidad
Tobago
Margarita
Vieques
0
0
0
0
0
1
1
0
0
1
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
1
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Culebra
Anegada
Virgin Gorda
Tortola
St Thomas
St John
St Croix
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Aruba
Cura Ëcao
Bonaire
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
788 J. C. Trejo-Torres and J. D. Ackerman
Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794
1
0
0
0
0
0
1
0
0
1
1
1
1
1
1
0
0
0
0
1
0
1
0
1
0
0
1
1
0
0
0
0
0
0
1
0
1
0
0
1
0
1
1
0
1
1
1
Species name/area name
Encyclia oncidioides
Encyclia plicata
Encyclia rufa
Encyclia serrulata
Encyclia tampensis
Epidendrum acunae
Epidendrum anceps
Epidendrum antillanum
Epidendrum blancheanum
Epidendrum carpophorum
Epidendrum ciliare
Epidendrum compressum
Epidendrum coronatum
Epidendrum cristatum
Epidendrum dendrobioides
Epidendrum difforme
Epidendrum diffusum
Epidendrum ¯oridense
Epidendrum hartii
Epidendrum ibaguense
Epidendrum imbricatum
Epidendrum isomerum
Epidendrum jamaicense
Epidendrum miserrimum
Epidendrum mutelianum
Epidendrum neoporpax
Epidendrum nocturnum
Epidendrum nutans
Epidendrum orientale
Epidendrum pallidi¯orum
Epidendrum paranaense
Epidendrum patens
Epidendrum polygonatum
Epidendrum portoricensis
Epidendrum ramosum
Epidendrum repens
Epidendrum rigidum
Epidendrum rivulare
Epidendrum scapelligerum
Epidendrum schlecterianum
Epidendrum secundum
Epidendrum strobiliferum
Epidendrum vincentinum
Epidendrum wrightii
Epistephium ellipticum
Epistephium parvi¯orum
Erythrodes hirtella
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
Guianas
Yucatan
Appendix 1 continued
0
0
0
0
0
0
1
1
0
1
1
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
1
0
0
0
0
0
0
1
1
0
1
0
0
0
1
0
1
0
0
0
1
0
0
0
1
0
0
1
0
1
1
1
0
0
0
0
0
1
0
0
0
1
0
1
0
1
0
1
1
1
0
1
0
0
0
1
1
1
1
1
0
1
1
1
0
0
0
1
Florida
Puerto Rico
Jamaica
0
0
1
0
1
1
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
1
1
1
1
1
1
0
1
1
1
0
0
0
0
0
1
1
0
0
0
0
1
0
0
1
1
0
1
1
0
0
1
1
1
1
1
1
1
0
1
1
1
1
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Hispaniola
Cuba
Isla Juventud
Mona
0
0
0
1
0
0
1
1
1
1
1
0
0
0
0
0
0
0
0
0
1
0
1
1
0
1
1
0
0
0
1
0
1
1
1
1
1
1
0
0
1
1
1
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grand Cayman
Little Cayman
Cayman Brac
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Turks
Inaguas
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Crooked
Long
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Salvador
Exumas
Cat
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Providence
Andros
Abacos
St Martin
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
Saba
St Barth Âelemy
St Kitts
Nevis
St Eustatius
Antigua
0
0
0
0
0
0
1
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Barbuda
Montserrat
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
1
1
0
0
0
1
1
0
0
0
1
0
0
1
1
0
0
1
0
0
1
0
0
0
0
1
0
1
0
0
0
1
1
0
0
0
0
1
0
0
0
0
0
0
1
1
0
0
1
0
0
0
1
1
0
0
0
1
0
0
0
1
0
0
1
0
0
0
0
1
0
0
1
0
1
0
0
0
1
1
0
0
0
0
1
0
0
0
0
0
0
1
0
0
1
1
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
1
1
0
1
1
0
0
0
1
1
0
0
0
1
0
0
0
0
0
0
1
0
0
1
0
0
0
0
1
0
1
0
0
0
1
1
0
0
0
0
1
Guadeloupe
Dominica
Martinique
St Lucia
St Vincent
0
0
0
0
0
0
1
1
0
1
1
0
0
0
1
1
0
0
0
1
0
0
1
1
0
0
1
0
0
0
0
1
0
0
1
0
1
0
0
0
1
1
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
1
1
1
1
1
0
0
0
0
1
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
1
0
0
1
0
1
1
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grenada
Barbados
Trinidad
Tobago
Margarita
Vieques
0
0
0
0
0
0
1
0
0
1
1
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
0
0
0
1
0
1
0
0
0
1
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Culebra
Anegada
Virgin Gorda
Tortola
St Thomas
St John
St Croix
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Aruba
Cura Ëcao
Bonaire
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Biogeography of the Antilles based on a parsimony analysis of orchid distributions 789
Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794
1
0
1
0
0
0
0
0
0
1
0
0
0
0
1
1
0
0
0
1
1
0
0
0
0
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
1
1
0
0
0
Species name/area name
Erythrodes major
Erythrodes plantaginea
Eulophia alta
Eulophia ecristata
Eurystyles ananassocomos
Fuertesiella pterichoides
Galeandra beyrichii
Govenia utriculata
Habenaria alata
Habenaria amal®tana
Habenaria bicornis
Habenaria distans
Habenaria dussii
Habenaria eustachya
Habenaria ¯oribunda
Habenaria leprieurii
Habenaria mesodactyla
Habenaria monorrhiza
Habenaria pauci¯ora
Habenaria quinqueseta
Habenaria repens
Hapalorchis lineatus
Harrisella porrecta
Helleriella punctulata
Homalopetalum vomeriforme
Huntleya lucida
Huntleya meleagris
Ionopsis satyrioides
Ionopsis utricularoides
Isochilus linearis
Jacquinella globosa
Jacquinella teretifolia
Kegeliella houtteana
Koellensteinia graminea
Leochilus carinatus
Leochilus labiatus
Leochilus puertoricensis
Leochilus scriptus
Lepanthopsis anthoctenium
Lepanthopsis melanantha
Lepanthopsis melanantha
Lepanthopsis microlepanthes
Leucohyle subulata
Liparis nervosa
Liparis neuroglossa
Liparis saundersiana
Liparis vexillifera
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
1
1
1
0
1
0
1
0
0
0
0
1
1
0
1
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
Guianas
Yucatan
Appendix 1 continued
0
1
1
0
1
0
1
1
1
1
0
1
0
1
0
0
0
1
0
0
1
1
1
1
0
0
0
1
1
1
1
1
0
0
0
0
1
0
0
1
1
0
0
1
0
1
1
0
1
1
0
1
0
1
1
1
0
0
1
0
1
1
0
0
1
0
1
1
1
1
0
1
0
0
1
1
1
1
1
0
0
0
1
0
0
0
1
1
1
1
1
1
1
1
Florida
Puerto Rico
Jamaica
0
0
1
1
0
0
1
1
0
0
0
1
0
0
1
0
0
0
0
1
1
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
1
0
0
1
0
0
0
0
1
1
1
1
1
1
1
1
0
1
1
0
1
1
0
0
1
0
1
1
1
1
0
1
0
0
1
1
1
1
1
0
0
1
1
0
1
1
1
1
1
1
1
0
1
1
0
0
0
0
0
0
0
0
1
0
1
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Hispaniola
Cuba
Isla Juventud
Mona
1
1
1
0
1
1
1
1
1
0
0
1
0
1
1
0
0
1
0
1
1
1
1
0
0
0
0
1
1
1
1
1
0
0
0
1
0
0
1
1
1
1
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grand Cayman
Little Cayman
Cayman Brac
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Turks
Inaguas
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Crooked
Long
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Salvador
Exumas
Cat
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Providence
Andros
Abacos
St Martin
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
Saba
St Barth Âelemy
St Kitts
Nevis
St Eustatius
Antigua
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Barbuda
Montserrat
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
1
1
1
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
1
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
1
1
0
0
0
0
0
0
1
0
0
0
Guadeloupe
Dominica
Martinique
St Lucia
St Vincent
0
1
1
0
0
0
0
0
1
1
0
0
1
1
1
0
0
1
0
0
0
0
0
1
0
0
0
0
1
1
1
0
0
0
0
1
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
0
1
1
1
1
0
1
0
0
0
0
1
1
1
1
1
1
0
1
1
1
1
0
1
0
0
0
0
1
1
0
0
1
0
1
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grenada
Barbados
Trinidad
Tobago
Margarita
Vieques
0
1
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Culebra
Anegada
Virgin Gorda
Tortola
St Thomas
St John
St Croix
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Aruba
Cura Ëcao
Bonaire
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
790 J. C. Trejo-Torres and J. D. Ackerman
Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794
0
1
1
0
1
0
0
0
0
0
0
1
0
1
1
0
1
1
1
0
1
0
1
1
1
1
1
0
0
0
1
1
0
1
0
1
0
0
0
1
0
0
1
1
1
1
0
Species name/area name
Liparis viridipurpurea
Lockhartia elegans
Lophiasis maculata
Lycaste barringtoniae
Macradenia lutescens
Malaxis domingensis
Malaxis hispanolae
Malaxis integra
Malaxis major
Malaxis massonii
Malaxis spicata
Malaxis umbelli¯ora
Malaxis unifolia
Maxilaria discolor
Maxillaria acutifolia
Maxillaria adendrobium
Maxillaria alba
Maxillaria brachybulbon
Maxillaria camaridii
Maxillaria coccinea
Maxillaria crassifolia
Maxillaria in¯exa
Maxillaria liparophylla
Maxillaria parvi¯ora
Maxillaria reichenheimana
Maxillaria rufescens
Maxillaria uncata
Maxillaria variabilis
Mesadenus lucayanus
Myrmecophila thomsoniana
Nidema ottonis
Notylia angustifolia
Octomeria apiculata
Octomeria graminifolia
Octomeria tridentata
Oeceoclades maculata
Oerstedella verrucosa
Oncidium altissimum
Oncidium ampliatum
Oncidium cebolleta
Oncidium ensatum
Oncidium meirax
Ornithocephalus gladiatus
Otostylis brachystalix
Palmorchis pubescens
Paphinia cristata
Pelexia adnata
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
1
1
0
0
0
0
0
0
1
0
0
0
1
1
0
0
0
0
0
1
Guianas
Yucatan
Appendix 1 continued
0
0
0
1
0
0
0
0
1
1
1
0
0
0
1
0
0
0
0
1
0
0
0
1
0
0
0
0
1
0
1
0
0
0
0
1
0
1
1
0
0
1
0
0
0
0
1
1
0
1
1
1
0
0
1
0
1
1
1
1
0
1
1
1
0
0
0
1
1
0
1
0
0
0
0
0
0
1
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
1
Florida
Puerto Rico
Jamaica
0
0
1
0
1
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
1
0
1
1
1
1
1
0
1
1
1
1
1
0
1
1
1
0
0
0
1
1
0
1
0
0
0
0
1
1
1
0
0
0
1
1
1
0
0
0
1
0
0
0
0
0
1
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
Hispaniola
Cuba
Isla Juventud
Mona
1
0
0
1
1
1
1
1
0
1
1
1
1
0
1
1
0
0
0
1
0
1
0
1
0
0
0
0
1
0
1
0
0
0
0
1
0
1
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grand Cayman
Little Cayman
Cayman Brac
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Turks
Inaguas
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Crooked
Long
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Salvador
Exumas
Cat
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Providence
Andros
Abacos
St Martin
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Saba
St Barth Âelemy
St Kitts
Nevis
St Eustatius
Antigua
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Barbuda
Montserrat
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
1
0
1
0
0
0
0
1
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
1
0
0
0
0
0
0
0
0
Guadeloupe
Dominica
Martinique
St Lucia
St Vincent
0
0
0
0
0
0
0
0
1
1
0
1
0
0
0
0
1
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
1
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
1
0
1
0
1
0
0
1
1
1
1
0
0
1
1
1
1
0
1
0
0
1
1
1
1
1
0
0
0
1
1
0
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grenada
Barbados
Trinidad
Tobago
Margarita
Vieques
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Culebra
Anegada
Virgin Gorda
Tortola
St Thomas
St John
St Croix
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Aruba
Cura Ëcao
Bonaire
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Biogeography of the Antilles based on a parsimony analysis of orchid distributions 791
Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794
1
1
0
0
1
1
0
0
1
0
1
0
1
1
1
0
0
0
0
1
0
1
1
0
0
0
0
1
0
0
0
0
1
1
1
1
0
1
1
1
0
1
0
0
0
0
0
Species name/area name
Peristeria cerina
Peristeria pendula
Pinelia leochilus
Platanthera replicata
Platystele ovalifolia
Platystele stenostachia
Platythelys querceticola
Pleurothallis ophioglossoides
Pleurothallis acutissima
Pleurothallis appendiculata
Pleurothallis aristata
Pleurothallis brighamii
Pleurothallis consimilis
Pleurothallis corniculata
Pleurothallis discoidea
Pleurothallis domingensis
Pleurothallis dussii
Pleurothallis erosa
Pleurothallis gelida
Pleurothallis grobyi
Pleurothallis helenae
Pleurothallis hymenantha
Pleurothallis imraei
Pleurothallis lanceola
Pleurothallis laxa
Pleurothallis mazei
Pleurothallis miguelii
Pleurothallis monophylla
Pleurothallis mornicola
Pleurothallis murex
Pleurothallis nummularia
Pleurothallis oblongifolia
Pleurothallis obovata
Pleurothallis polygonoides
Pleurothallis pruinosa
Pleurothallis pubescens
Pleurothallis racemi¯ora
Pleurothallis revoluta
Pleurothallis ruscifolia
Pleurothallis sclerophylla
Pleurothallis sertularioides
Pleurothallis sicaria
Pleurothallis testaefolia
Pleurothallis tribuloides
Pleurothallis trichophora
Pleurothallis trichostata
Pleurothallis velaticaulis
0
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
Guianas
Yucatan
Appendix 1 continued
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
1
1
1
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
1
1
0
1
0
1
0
0
1
0
1
0
0
1
1
0
0
1
0
0
1
1
0
0
1
0
1
0
1
0
1
0
1
1
0
0
0
Florida
Puerto Rico
Jamaica
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
1
0
0
0
1
1
0
1
0
1
0
1
1
1
0
1
0
0
0
0
0
0
0
1
1
1
1
0
1
0
1
0
1
0
1
0
1
1
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Hispaniola
Cuba
Isla Juventud
Mona
0
0
1
0
0
0
1
0
0
1
1
1
0
1
0
1
0
1
1
1
1
0
1
1
1
1
1
0
1
1
0
1
1
0
1
1
1
0
1
0
0
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grand Cayman
Little Cayman
Cayman Brac
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Turks
Inaguas
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Crooked
Long
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Salvador
Exumas
Cat
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Providence
Andros
Abacos
St Martin
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Saba
St Barth Âelemy
St Kitts
Nevis
St Eustatius
Antigua
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Barbuda
Montserrat
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
1
0
0
0
0
0
1
1
0
0
0
0
0
0
0
1
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
Guadeloupe
Dominica
Martinique
St Lucia
St Vincent
0
0
0
0
0
0
1
1
0
0
1
0
0
0
0
1
1
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
1
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
1
1
0
0
0
1
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grenada
Barbados
Trinidad
Tobago
Margarita
Vieques
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Culebra
Anegada
Virgin Gorda
Tortola
St Thomas
St John
St Croix
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Aruba
Cura Ëcao
Bonaire
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
792 J. C. Trejo-Torres and J. D. Ackerman
Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794
0
1
1
1
1
0
1
0
0
0
0
0
1
1
0
1
0
1
1
0
1
0
0
1
1
1
0
1
1
1
0
1
0
0
1
0
1
1
0
1
0
0
1
1
0
0
0
Species name/area name
Pleurothallis wilsonii
Pogonia grandi¯ora
Pogonia tenuis
Polystachya concreta
Polystachya foliosa
Ponthieva brittoniae
Ponthieva diptera
Ponthieva harrisii
Ponthieva pauci¯ora
Ponthieva petiolata
Ponthieva racemosa
Ponthieva ventricosa
Prescottia oligantha
Prescottia stachyodes
Prosthechea boothiana
Prosthechea cochleata
Prosthechea fragans
Prosthechea pygmaea
Prosthechea vespa
Pseudocentrum minus
Psilochilus macrophyllus
Psychilis kraenzlinii
Psychilis macconnelliae
Psygmorchis pusilla
Reichenbachanthus re¯exus
Rodriguezia lanceolata
Sarcoglottis sceptrodes
Scaphyglottis fusiformis
Scaphyglottis modesta
Scaphyglottis prolifera
Schiedeella amesiana
Schomburgkia ®mbriata
Schomburgkia humboldtii
Schomburgkia lysonsii
Sobralia fragrans
Spiranthes torta
Stanhopea grandi¯ora
Stelis muscifera
Stelis perpusilli¯ora
Stelis pygmaea
Stelis scabrida
Stelis trigoni¯orum
Stenia pallida
Stenorrhynchos lanceolatum
Stenorrhynchos speciosum
Stenorrhynchos squamulosum
Tetramicra bulbosa
0
0
0
0
1
0
0
0
0
0
1
0
0
1
1
1
0
1
0
0
0
0
0
1
0
0
1
0
0
1
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
Guianas
Yucatan
Appendix 1 continued
1
0
0
1
1
0
0
0
0
0
1
1
1
1
0
1
0
1
0
0
1
1
1
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
1
1
0
0
0
1
0
0
0
1
0
0
1
1
0
1
1
1
0
1
1
1
1
0
1
1
1
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
1
1
0
1
1
1
1
Florida
Puerto Rico
Jamaica
0
0
0
1
0
1
0
0
0
0
1
0
1
0
1
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
1
0
0
1
1
1
1
0
1
1
1
1
1
1
1
1
1
1
1
0
1
0
0
0
1
0
0
0
1
0
1
0
0
1
0
1
0
0
0
0
0
1
0
1
1
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Hispaniola
Cuba
Isla Juventud
Mona
1
0
0
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
1
0
0
0
1
0
1
0
0
0
0
1
0
0
1
1
0
1
0
1
1
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grand Cayman
Little Cayman
Cayman Brac
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Turks
Inaguas
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
Crooked
Long
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Salvador
Exumas
Cat
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
1
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Providence
Andros
Abacos
St Martin
0
0
0
1
0
1
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
Saba
St Barth Âelemy
St Kitts
Nevis
St Eustatius
Antigua
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Barbuda
Montserrat
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
1
1
0
1
1
1
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
1
1
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
1
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
1
0
0
1
0
0
0
1
0
0
1
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
Guadeloupe
Dominica
Martinique
St Lucia
St Vincent
1
0
0
1
0
0
0
0
0
1
0
0
1
1
0
1
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
1
1
1
0
0
0
0
0
0
0
0
1
0
0
1
1
1
0
1
0
0
1
1
1
0
1
1
0
0
1
0
0
0
1
1
1
1
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grenada
Barbados
Trinidad
Tobago
Margarita
Vieques
0
0
0
1
1
0
0
0
0
1
0
0
1
0
0
0
1
0
0
0
1
0
0
0
1
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Culebra
Anegada
Virgin Gorda
Tortola
St Thomas
St John
St Croix
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Aruba
Cura Ëcao
Bonaire
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Biogeography of the Antilles based on a parsimony analysis of orchid distributions 793
Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
0
0
1
0
1
0
0
1
1
1
1
0
Species name/area name
Tetramicra canaliculata
Tetramicra eulophiae
Tetramicra parvi¯ora
Tetramicra urbaniana
Tolumnia bahamense
Tolumnia calochila
Tolumnia gauntletii
Tolumnia lyrata
Tolumnia prionochila
Tolumnia sasseri
Tolumnia urophyllum
Tolumnia variegata
Trichopilia fragans
Trichosalpinx ciliaris
Trichosalpinx dura
Trigonidium egertonianum
Triphora amazonica
Triphora gentianoides
Triphora hassleriana
Triphora surinamensis
Triphora yucatanensis
Trizeuxis falcata
Tropidia polystachya
Wullschlaegellia aphylla
Wullschlaegellia calcarata
Xylobium colleyi
Xylobium foveatum
Xylobium palmifolium
Zootrophion atropurpureum
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
1
0
0
1
0
1
0
0
0
0
0
0
Guianas
Yucatan
Appendix 1 continued
1
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
1
0
1
1
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
1
0
0
1
0
1
0
0
1
1
0
0
1
1
1
Florida
Puerto Rico
Jamaica
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
1
0
1
0
0
0
0
0
0
0
1
0
1
0
1
0
1
0
0
0
1
1
0
1
0
0
1
0
0
0
0
1
1
0
0
0
1
1
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Hispaniola
Cuba
Isla Juventud
Mona
1
1
1
0
0
1
0
0
0
0
1
1
1
0
1
0
1
1
1
1
0
0
1
0
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grand Cayman
Little Cayman
Cayman Brac
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Turks
Inaguas
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Crooked
Long
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Salvador
Exumas
Cat
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Providence
Andros
Abacos
St Martin
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Saba
St Barth Âelemy
St Kitts
Nevis
St Eustatius
Antigua
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Barbuda
Montserrat
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
Guadeloupe
Dominica
Martinique
St Lucia
St Vincent
1
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
1
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
1
0
1
0
1
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Grenada
Barbados
Trinidad
Tobago
Margarita
Vieques
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Culebra
Anegada
Virgin Gorda
Tortola
St Thomas
St John
St Croix
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Aruba
Cura Ëcao
Bonaire
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
794 J. C. Trejo-Torres and J. D. Ackerman
Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 775±794