Biological Journal of the Linnean Society. 101, 288-322

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Biological Journal of the Linnean Society. 101, 288-322
Biological Journal of the Linnean Society, 2010, 101, 288–322. With 9 figures
Molecular systematics of Selenops spiders (Araneae:
Selenopidae) from North and Central America:
implications for Caribbean biogeography
SARAH C. CREWS1,2* and ROSEMARY G. GILLESPIE1
1
University of California Berkeley, Department of Environmental Sciences Policy and Management,
137 Mulford Hall, Berkeley, CA 94720-3114, USA
2
Berkeley City College, Department of Science and Biotechnology, 2050 Center Street, Berkeley, CA
94704, USA
Received 16 February 2010; revised 3 May 2010; accepted for publication 3 May 2010
bij_1494
288..322
The Caribbean region includes a geologically complex mix of islands, which have served as a backdrop for some
significant studies of biogeography, mostly with vertebrates. Here, we use the tropical/subtropical spider genus
Selenops (Selenopidae) to obtain a finer resolution of the role of geology in shaping patterns of species diversity.
We obtained a broad geographic sample from over 200 localities from both the islands and American mainland.
DNA sequence data were generated for three mitochondrial genes and one nuclear gene for eleven outgroup taxa
and nearly 60 selenopid species. Phylogenetic analysis of the data revealed several biogeographic patterns common
to other lineages that have diversified in the region, the most significant being: (1) a distinct biogeographic break
between Northern and Southern Lesser Antilles, although with a slight shift in the location of the disjunction; (2)
diversification within the islands of Jamaica and Hispaniola; (3) higher diversity of species in the Greater Antilles
relative to the Lesser Antilles. However, a strikingly unique pattern in Caribbean Selenops is that Cuban species
are not basal in the Caribbean clade. Analyses to test competing hypotheses of vicariance and dispersal support
colonization through GAARlandia, an Eocene–Oligocene land span extending from South America to the Greater
Antilles, rather than over-water dispersal. © 2010 The Linnean Society of London, Biological Journal of the
Linnean Society, 2010, 101, 288–322.
ADDITIONAL KEYWORDS: Bayesian phylogenetics – island biogeography – likelihood analysis of
geographic range evolution.
INTRODUCTION
Remote islands form the basis for many biological
studies because of their ability to act as a laboratory,
with repeated sets of ecological and/or evolutionary
experiments occurring within a circumscribed time
frame (Cronk, 1997; Losos et al., 1998; Gillespie &
Roderick, 2002; Gillespie, 2004; Ricklefs & Bermingham, 2008). While the Hawaiian Islands have served
as a model system for processes of in situ diversification, the long history of studies on the biota of the
Caribbean has provided some of the most important
*Corresponding author. E-mail: [email protected]
288
insights into the complex interaction between colonization and diversification. In particular, the Caribbean has served as the setting for the establishment
of most of the central tenets in the equilibrium theory
of island biogeography (Munroe, 1948), the arguments being formulated independently by MacArthur
and Wilson (1963, 1967) much later (Lomolino &
Brown, 2009). More recent research on the islands
has allowed an understanding of the interplay
between ecological and evolutionary processes in
shaping species diversity (Losos & Schluter, 2000;
Schoener, Spiller & Losos, 2001).
The primary feature of the Caribbean region that
makes it particularly useful for examining the interaction between colonization and diversification is its
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
long and complex geological history. The Caribbean
Basin began forming nearly 140 Mya. Islands in the
basin consist of four different types: (1) land-bridge
islands which were connected to each other or to the
mainland at times of lower sea level; (2) continental
islands which broke off from the mainland through
tectonic displacement; (3) uplifted limestone islands;
and (4) volcanic islands (MacPhee & Iturralde-Vinent,
2005; Robertson, 2009). Despite their limited isolation, the age and geologic complexity of the area have
provided ‘well-defined paths of entry by which immigrants may reach’ the islands (Munroe, 1948). Moreover, the islands have served as the setting for
adaptive radiation among lineages with limited dispersal ability, in particular lizards of the genus Anolis
(Losos, 1992, 1994, 2009), frogs of the genus Eleutherodactylus (Hedges, 1989; Heinicke, Duellman &
Hedges, 2007), some lineages of insects [e.g. beetles
(Liebherr, 1988b), flies (Wilder & Hollocher, 2003)]
and plants [e.g. lineages within the Melastomaceae
(Michelangeli et al., 2008) and Asteraceae (FranciscoOrtega et al., 2008)]. Although studies to date have
provided insights into how the individual lineages
have colonized and subsequently diversified within
the island system, notable controversies remain,
including the source of colonists and the means by
which they colonized the islands, biogeographic patterns within lineages and whether these patterns
might be expected to be shared across multiple lineages (Guyer & Savage, 1986; Williams, 1989;
Hedges, Hass & Maxon, 1992; Crother & Guyer, 1996;
Hedges, 1996a,b).
A particular focus of debate has been the role of
vicariance vs. dispersal in shaping the Caribbean
biota. Hedges and colleagues (Hedges et al., 1992;
Hedges, 1996a,b; Hedges & Heinicke, 2007; Heinicke
et al., 2007), working with herpetofauna, have suggested that the absence of lineages older than the
break-up of the proto-Antilles (a contiguous land
mass between North and South America) precludes a
vicariant origin and they argue for the initial colonization of most taxa via over-water dispersal on
flotsam. A similarly dominant role for dispersal has
been suggested for multiple lineages of plants, such
as Miconieae (Michelangeli et al., 2008). In contrast,
other studies have suggested that vicariance has
played a larger role than dispersal in the initial
colonization of the Caribbean; for example, in lizards
(Crother & Guyer, 1996; Iturralde-Vinent & MacPhee,
1999; MacPhee & Iturralde-Vinent, 2005) and some
plants [e.g. Euphorbiaceae (van Ee et al., 2008)].
A related controversy focuses on the hypothesis of
GAARlandia (Greater Antilles + Aves Ridge), first
proposed by Iturralde-Vinent & MacPhee (1999), who
used geological data and fossil evidence to demonstrate the likely existence of a land span connecting
289
South America to the Greater Antilles during the
Eocene–Oligocene transition 35–33 Mya. The land
span, although probably short-lived, may have provided an avenue for terrestrial organisms to colonize
the Greater Antilles from South America. Among
mammals, molecular phylogenies of primates and
hystricognath rodents are consistent with the model,
while sloths and insectivorans are not (Dávalos,
2004). The pattern in plants is similarly mixed.
Molecular phylogenetic data from the genera Croton
(Euphorbiaceae) (van Ee et al., 2008) and Styrax (Styracaceae) (Fritsch, 2003) show that the timing of
divergence of lineages is consistent with the GAARlandia hypothesis. However, similar data from
endemic legume radiations in the Greater Antilles,
although initially thought to indicate ancient splitting
between lineages consistent with the GAARlandia
hypothesis (Lavin et al., 2001), show more recent
diversification (Lavin & Beyra-Matos, 2008), which is
likely to hold also for lineages of Asteraceae
(Francisco-Ortega et al., 2008).
Clearly, the timing and frequency of dispersal and
vicariance, and the interplay between the two, varies
across biotic assemblages. The challenge, then, is to
understand the circumstances dictating the relative
roles of each and how they interact. Arthropods,
because they can provide a fine-scale resolution of
biogeographic patterns (Ferrier et al, 2004), are ideal
candidates for elucidating the nature of these relationships. Although the biogeography of terrestrial
invertebrates in the Caribbean has been examined in
some detail (see Liebherr, 1988a and chapters
therein), few recent studies have been attempted,
with little molecular information on the timing and
nature of the interplay between colonization and
diversification. However, there are some notable
exceptions (Davies & Bermingham, 2002; Wilder &
Hollocher, 2003; Brisson, Wilder & Hollocher, 2006).
In particular, recent studies on spiders (Sicariidae:
Loxosceles) support the GAARlandia hypothesis in the
colonization of the lineage of North from South
America (Binford et al., 2008), while crickets show a
more mixed pattern of both vicariance and dispersal,
coupled with intra-island diversification (Oneal,
2009).
In this study, we combine molecular and morphological methods to examine the phylogenetic relationships and biogeographic history of the cursorial and
dispersal-limited spider genus Selenops (Araneae:
Selenopidae) in the Caribbean. These primarily tropical and subtropical spiders (Muma, 1953; Corronca,
1998; Alayón, 2005) are distinctive in that they are
extremely dorsoventrally flattened and exceedingly
fast. They are found in a variety of habitats and
microhabitats (Crews, Wienskoski & Gillespie, 2008).
Although the genera and species groups have
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
290
S. C. CREWS and R. G. GILLESPIE
Figure 1. Map of the study area. The Americas; the boxed region shows the primary study area.
undergone several revisions (Muma, 1953; Corronca,
1998; Alayón, 2005), there is no phylogenetic framework for the family or for any of the component
genera. They were chosen for the current study
because of their high diversity and abundance in the
Caribbean, where they occur in both the Greater and
Lesser Antilles, as well as on the adjacent mainland
(southern North America and throughout South
America) (Muma, 1953; Crews, 2005; Crews et al.,
2008, 2009;) (Figs 1, 2). Accordingly, they provide the
potential to reveal fine-scale biogeographic patterns
across the islands of the Caribbean. The current
study uses the genus to infer the relative importance
of the following two processes in dictating the biogeographic history of the lineage in the Caribbean: (1)
the frequency of colonization to the Caribbean region
from a mainland source and between islands within
the Caribbean; and (2) whether within-island diversification has occurred through a single radiation or
through dispersal and multiple radiations. We also
test the hypotheses of dispersal and vicariance in the
framework of the GAARlandia hypothesis and likelihood biogeographic analysis.
MATERIAL AND METHODS
TAXON SAMPLING
A comprehensive geographic sample of the genus was
obtained from the Caribbean region, including most
islands and several sites throughout Mexico, Central
America and the South American mainland (see also
Supporting Information, Figs S1, S2). Political reasons
prohibited us from obtaining permits to collect several
endemic species from Cuba and the single species from
Navassa Island. The implications for these omissions
are discussed at the end of this paper. Outgroups
included other genera in the family Selenopidae from
all major geographic locations where the family is
found, in particular the type of the genus (Selenops
radiatus Latreille) from Africa, Selenops bursarius
Karsch from Japan, Selenops montigenus Simon from
Nepal/India and representatives of the three other
genera described from Africa (six species of Anyphops,
one species of Hovops and one species of Garcorops),
as well as an undescribed Australian genus. Chosen
representatives outside of the family include a broad
sample of eight genera from two families, the
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
291
Figure 2. Map of the study area showing the number of localities per region (the first number), the number of total
specimens per region (the second number) and the number of species collected out of possible known species per region
(the third and fourth numbers, respectively). For more detailed collection information, see the Appendix and Supporting
Information (Figs S1, S2).
Table 1. Genetic loci and primer pairs used for PCR amplification
CO1 LCO1_1490
C1N_2568
5′-GGTCAACAAATCATAAAGATATTG-3′ – Folmer et al. 1994
5′-GCTACAACAATAATAAGTATCATG-3′ – Hedin & Maddison, 2001
16S–ND1
12350mod
13398
Histone
H3aF
H3aR
5′-TTDGNTACCAAGCAGACVGC-3′ – this study
5′-CGCCTGTTTAACAAAAACAT-3′ – Simon et al. 1994
5′-ATGGCTCGTACCAAGCAGACVGC-3′ – Colgan et al. 1998
5′-ATATCCTTRGGCATRATRGTGAC-3′ – Colgan et al. 1998
CO1, cytochrome oxidase I; 16S, ribosomal DNA; ND1, NADH dehydrogenase I.
Sparassidae and the Ctenidae, and were based on
unpublished data (M. Ramirez, pers. comm.) (Table 4).
The genus Selenops has also been found in Dominican amber and one of these specimens is an adult
male, described by Schawaller (1984) as Selenops
beynai. The specimen was scanned using X-ray computed tomography, as in Penney et al. (2007);
however, the poor preservation of the genitalia prohibited even tentative incorporation into the phylogenetic framework.
MOLECULAR
METHODS
Four gene fragments were amplified – three mitochondrial [cytochrome oxidase I (CO1), 16S ribosomal DNA
(16S) and the intervening leucine tRNA and NADH
dehydrogenase I (ND1)] and one nuclear [histone 3a
(H3)] (see Table 1). The respective lengths of the
amplification products were ~850, ~800 and ~330 base
pairs (bp). DNA sequences can be found on GenBank
(GU109549–GU110746, HM575429–HM576623, and
HM576658). These markers were chosen as they have
become a standard in spider molecular phylogenetics,
with several primers available for each gene (Hedin &
Maddison, 2001; Arnedo et al., 2004; Crews & Hedin,
2006). Also, the chosen genes evolve at different rates
and contain both protein and non-protein coding
regions. DNA was extracted from a portion of a leg
using a Qiagen DNeasy Tissue Kit following the manufacturer’s protocol. Each new specimen used in this
study was given an individual number (e.g. sel_001)
and has been deposited in the Essig Museum of
Entomology at the University of California, Berkeley
and the California Academy of Sciences. Remaining
genomic DNA is stored at -80 °C in the Gillespie
and Roderick Laboratories, University of California,
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292
S. C. CREWS and R. G. GILLESPIE
Berkeley. Primer pairs used are given in Table 1. In
some cases, primarily with outgroup taxa, amplification was difficult and, in such instances, the Epicentre
FailSafe PCR kit was used. In the majority of cases,
sequence data were obtained for all gene fragments for
multiple representatives of each species. In one situation with the species Selenops insularis Keyserling,
there was evidence for multiple copies of H3a in some
specimens, thus these sequences were not analysed for
these individuals.
PHYLOGENETIC
METHODS
Alignments of the protein-coding loci CO1, ND1 and
H3a were performed manually using Mesquite ver.
2.5 (Maddison & Maddison, 2008), with the aminoacid translations used as a guide. The 16S data
were aligned using secondary structure based on the
model from Masta (2000). While there were some
length differences between taxa, alignment was
straightforward.
Data were partitioned by codon position for protein
coding genes, by stems and loops for ribosomal DNA
and by gene for both the maximum likelihood and
Bayesian analyses to improve the fit of the substitution model to the data (Nylander et al., 2004; Brandley, Schmitz & Reeder, 2005). The doublet model of
nucleotide substitution was used for the stem regions
of 16S and the tRNA (Schöniger & von Haeseler,
1994; Kjer, 2004). Maximum likelihood analyses were
performed with RAxML ver. 7.0.4 (Stamatakis, 2006)
and Bayesian analyses were performed using
MrBayes ver. 3.1.2 (Huelsenbeck & Ronquist, 2001;
Ronquist & Huelsenbeck, 2003; Altekar et al., 2004).
RAxML is able to analyse partitioned data, but only
under the generalised time reversible (GTR) model,
thus, while the same partitioning regime was used in
both Bayesian and likelihood analyses, this was the
model that was used in the maximum likelihood
analyses. To determine the models for each partition
in the Bayesian analysis, MrModeltest ver. 2.3
(Nylander, 2004) was used. Models were chosen using
the Akaike information criterion (AIC; Akaike, 1973;
see Posada & Buckley, 2004) and are listed in Table 2.
LIKELIHOOD
ANALYSES
RAxML maximum likelihood analyses were conducted in a variety of ways following the suggestions
of the author (Stamatakis, 2006). First, one analysis
was conducted which included 893 terminals after
identical haplotypes were removed. However, to ease
the computational strain for more intensive analyses,
terminals that were ⱕ 0.3% different were removed
from the analysis (sensu McGuire et al., 2007). This
truncated data set contained 306 terminals.
Table 2. Partitions used in likelihood and Bayesian
analyses and selected models for each partition used in
Bayesian analyses
Partition
Selected model
16S stems
16S half stems
16S loops
Leucine tRNA stems
Leucine tRNA loops
ND1 postion 1
ND1 position 2
ND1 position 3
CO1 position 1
CO1 position 2
CO1 position 3
H3a position 1
H3a position 2
H3a position 3
GTR + G + doublet
GTR + G
GTR + I + G
HKY + G + doublet
HKY + G
GTR + I + G
GTR + I + G
GTR + G
GTR + I + G
GTR + I + G
GTR + G
GTR + I
JC + I
K80 + G
CO1, cytochrome oxidase I; 16S, ribosomal DNA; ND1,
NADH dehydrogenase I.
The RAxML manual suggests two ways to analyse
data – the ‘fast and easy way’ and the ‘hard and slow
way’ (Stamatakis, 2006). The fast and easy way was
used to analyse the full data set because of its large
size (~900 terminals and ~2000 bps). The hard and
slow way was used to analyse the smaller data set
and allows the program to find ‘good’ settings particular to an individual data set. The user’s manual was
followed exactly for the analysis of the truncated data
set using the ‘hard and slow’ method. First, five
randomized maximum parsimony trees were generated and then each tree was inferred using a fixed
setting of ten for the initial rearrangement. Next, this
setting was automatically determined for the same
five starting trees and whichever settings yielded the
best likelihood scores were used for subsequent analyses. The second part of the ‘hard and slow’ method
involves the number of rate categories. For this, the
number of rate categories is increased by 15, from 10
to 55 for each of the five starting trees, using whichever setting worked best from the initial rearrangement analyses. Finally, ten analyses were run using
the best settings from the above experiments and
bootstraps from 500 iterations were then added to the
tree with the best likelihood. For all RAxML analyses,
the rapid bootstrap algorithm was used (Stamatakis,
Hoover & Rougemong, 2008).
BAYESIAN
ANALYSES
Several analyses were run using MrBayes-mpi on the
cluster at the Museum of Vertebrate Zoology, University of California, Berkeley, as well as on the CIPRES
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
cluster at the San Diego Supercomputer Center.
Despite using the truncated data set and running the
jobs in parallel, analyses required months to near
completion. Programs were run using the default
settings for 40 million generations and, if convergence
was not met, the generations were increased in increments up to 100 million, saving every 1000th tree.
Convergence was assessed using Are We There Yet?
(AWTY) (Wilgenbusch, Warren & Swofford, 2004;
Nylander et al., 2008).
LAGRANGE
ANALYSES
The program Lagrange (Ree et al., 2005; Ree &
Smith, 2008) was used to test hypotheses of vicariance and dispersal. Lagrange uses likelihood models
to test geographic range evolution and allows changes
in dispersal and extinction parameters at different
times in the past, allowing the incorporation of external information such as geological data and dispersal
capabilities. For example, if a land mass did not exist
at a particular time period, because it had not yet
emerged or was inundated, the rate of dispersal to the
land mass would be 0 during this time and could
increase during the time period(s) the land mass was
available for colonization. In an area as geologically
complex as the Caribbean, there are nearly endless
ways to parameterize the models, but simplicity was
maintained throughout each analysis.
Lagrange requires a tree and a matrix of range
data for the included taxa. We analysed a truncated
data set, selecting one specimen from each species,
along with the outgroups, using a partitioned RAxML
search for the best tree. We then pruned the outgroups before conducting the Lagrange analyses to
make the computational load smaller, and because
the focus of the questions concerns only the ingroup.
We ensured the tree had the same basic structure as
trees from the more complete analyses and that all
relationships supported in those analyses also
appeared in this tree.
We divided the range of the Selenopids in North
and Central America into five areas: C (Central
America and Mexico), S (South America), G (Greater
Antilles), N (Northern Lesser Antilles), A (Southern
Lesser Antilles). Although certain parts of these
regions were not available for colonization throughout
particular time periods (i.e. some of the Greater Antilles have been emergent longer than others, etc.), we
simply used the maximum times from their first
appearance. We set the age of the root node of the tree
to 130 Myr, as it is assumed a split between the
ingroup, i.e. American selenopids, and the outgroup,
i.e. African selenopids, was caused by the separation
of Africa from South America. Dating vicariance
events by the initiation of mid-ocean ridge spreading
293
Table 3. Correspondence between time slices and geographical ranges defined for models used in Lagrange
analyses
Time slice
Land availability
3.0
5.0
Closing of the Isthmus of Panamá
Most recent appearance of Northern
Lesser Antilles
Most recent appearance of Southern
Lesser Antilles
Disappearance of GAARlandia
Appearance of GAARlandia
From 55–50 Mya, a part of Jamaica was
connected to Central America via the
Nicaraguan Rise
Time after which land was available in
the Greater Antilles region
Age of root node, corresponds to
separation of Africa and South
America
12.0
33.0
35.0
50.0
55.0
130.0
is problematic, in that in some cases this upper bound
is too old as a result of chance transoceanic interchange after actual separation. However, the existence of distinct clades of Selenops on the different
continents would argue for little genetic exchange
between the continental land masses (Smith & Peterson, 2002) and would therefore indicate that it is
indeed appropriate to use the separation of Africa and
America to date Selenops. Throughout all analyses,
we focused on six time periods which correspond to
the availability of land for colonization (Table 3).
The following three analyses consisted of two
models each, one representing each of three scenarios
with, and without, GAARlandia. The three scenarios
were: (1) a dispersal-based scenario where distance
between land masses determines the probability of
colonization; (2) a dispersal-based scenario in which
the ability to colonize an available land mass is not
dependent on distance, thus the colonization of any
one land mass from another is equiprobable; (3) a
vicariance-based scenario, with little to no over-water
dispersal. This means that colonization of one area
from another could occur only through connections of
one land mass to another. In some cases, certain
areas were never connected to other land masses,
such as the Lesser Antilles. In this case, the probability of dispersal is not set to zero, but rather a very low
probability, as the presence of the spiders indicates
colonization at some point in the past.
It is possible to set different dispersal probabilities
for each direction, so that the probability of moving
from one region to another can be lower or higher
than in the opposite direction. However, to maintain
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
294
S. C. CREWS and R. G. GILLESPIE
simplicity, bidirectional probabilities were set as
equal. Within each model, the only parameter
changed between the two analyses was the probability of colonization with and without the presence of
GAARlandia. The maximum range size was set to two
areas and the areas G, N and A were excluded from
the root (> 130 Myr) as they were not available for
colonization at this time (Table 3).
RESULTS
SAMPLING
We obtained over 1000 specimens from over 200
localities within the area of primary focus for this
study. In total, we have 29 out of 41 Caribbean island
species, half of the known Mexican and Central
American species and one fifth of the described South
American species (Appendix).
PHYLOGENETIC
ANALYSES
Likelihood analysis
The tree from the analysis of the full data set is
shown in Figure 3 and has a likelihood score of
-61 544.60. Nodes with bootstrap (BS) values ⱖ 70%
are considered to be supported. There is no support
(BS < 70%) for many basal nodes. Further discussion
of the results from this tree is given below where
compared with trees from the other analyses.
In the analysis of the truncated data set, the best
likelihood score came from the trial with a fixed
setting of 10 for the initial rearrangement, rather
than the automatic setting (Table 5). The best likelihood from the experiment to determine a good setting
for the number of rate categories occurred when this
setting was at 25 (Table 6). Thus, the initial rearrangement setting was fixed at 10 (-i 10) and the
number of rate categories was set to 25 (-c 25). The
best overall likelihood with these settings from
the MultTrees analysis came from the second run
(Table 7) and the results are shown in Figure 4. The
overall structure is similar to the tree obtained from
the analysis of the full data set, in which many basal
nodes are not supported, while nodes above these are.
This tree is discussed in more detail below.
Bayesian analysis
The analyses were run for 64 million generations (the
maximum possible given limits of storage space for
our output files). According to the cumulative plot
from AWTY (Wilgenbusch et al., 2004; Nylander et al.,
2008), the run reached convergence near 55 million
generations. Because convergence was only reached
very late in the analysis, the first 90% of trees were
eliminated as burn-in, leaving ~12 000 trees from
which to compute a consensus. This tree is shown in
Figure 5 and nodes with posterior probability values
ⱖ 0.95 are considered to be supported. The branch
lengths are longer than in the likelihood analyses
and, while a few more basal nodes are supported than
in the likelihood analyses, the overall pattern is the
same. The similarities and differences among all
three trees are discussed below.
Comparison of trees
All three trees are very similar with many of the
minor differences not supported. The remainder of
the basal nodes occurring below the Selenopids of
North and Central America is only supported as
monophyletic in the Bayesian analysis. The focal taxa
of the study, the Selenopids of North and Central
America, are monophyletic and further subdivided
into a well-supported strictly Caribbean clade (Fig. 6,
clade A) and the remaining taxa, supported as a clade
in the Bayesian tree only; (Fig. 6, clade B), including
taxa from the south-western USA, Mexico, Central
America, the Southern Lesser Antilles (SLA) and
South America. Within this clade B, although basal
relationships are not supported, all analyses support
a southern Caribbean basin clade (Fig. 6, clade C)
consisting of taxa from Aruba, Bonaire, Curaçao,
Trinidad and Tobago. Selenops n. sp. 5 from Aruba is
always sister to Selenops curazao from Bonaire and
Curaçao and, this clade (Fig. 6, clade D), is always
sister to Selenops willinki from northern South
America and Tobago + S. geraldinae from Trinidad
(Fig. 6, clade E). Also within clade B, another wellsupported clade in all analyses consists of Selenops
banksi, found in Panama and South America, and
Selenops micropalpus, found in the Southern Lesser
Antilles from Dominica to St Vincent and the Grenadines (Fig. 6, clade F).
There is support for a sister group relationship
between the South American taxa + the Central and
North American taxa in the Bayesian tree only. Both
the Bayesian analysis and the likelihood analysis
of the truncated data set support a Central
American + North American clade (Fig. 6, clade G), as
well as one between the widespread Selenops mexicanus, Selenops gracilis and a new species found only in
Mexico (Fig. 6, clade H). In the Bayesian tree, S.
mexicanus is paraphyletic. There is little support for
any other relationships in clade B, other than the
species from the Selenops debilis group of the southwestern USA and Northern Mexico (Fig. 6, clade I).
The Caribbean clade (Fig. 6, clade A) consists only
of taxa from Caribbean islands and is strongly supported in all analyses, but, again, with little support
for basal nodes, the exception being the widespread
Selenops lindborgi and its sister species, S. n. sp. 3,
which are supported as sister to the rest of the
Caribbean taxa (Fig. 6, clade J).
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SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
295
Figure 3. Likelihood tree resulting from the RAxML analysis of the full data set. The map above the tree depicts the
Caribbean islands and the colours correspond to branches in the tree and indicate on which island the species is found.
Multiple colours along a branch indicate that the species is found on multiple islands. A branch outlined in black indicates
the species is found in Cuba. Selenops radiatus (highlighted in blue) is the type of the genus.
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
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S. C. CREWS and R. G. GILLESPIE
Table 4. Outgroup taxa used to root trees, collection localities, voucher numbers and location of vouchers
Family
Genus and species
Locality
Voucher number and locality
Ctenidae
Ctenidae
Ctenidae
Madagascar, Ranomafana
French Guiana, Tresor Nature Reserve
French Guiana, Emerald Jungle Village
CASENT9024024 – CAS
CASENT9021738 – CAS
CASENT9021735 – CAS
Ctenidae
Sparassidae
Sparassidae
Vulsor sp.
Phoneutria fera
Cupiennius ca.
granadensis
Acanthoctenus sp.
Olios sp. 1
Olios sp. 2
ARAMR000556 – MACN
In author’s personal collection
In author’s personal collection
Sparassidae
Sparassidae
Sparassidae
Sparassidae
Sparassidae
Polybetes pythagoricus
Heteropoda sp.
Heteropoda sp.
Damastes sp. 1
Damastes sp. 2
Argentina, Parque Nacional Cope, límte NE
USA, California, Esparto
México, Baja California, north of Guerrero
Negro
Argentina, Buenos Aires Prov., José Mármol
Nepal, near Sauraha
Tanzania
Madagascar, Toliara
Madagascar, Ambohitantely
Ar 10876 – MACN
In author’s personal collection
In author’s personal collection
sel_554 – CAS
CASENT9015896 – CAS
CAS, California Academy of Sciences; MACN, Museo Argentino de Ciencias Naturales.
Table 5. Likelihoods from the ‘hard and slow’ RAxML
analyses to determine the best initial rearrangement
setting for the data
-ln(L) for initial
rearrangement setting
fixed at 10
-ln(L) for automatic
initial rearrangement
setting
49 876.238442*
49 881.090042
49 880.995332
49 881.483263
49 879.897717
49 880.146295
49 877.349169
49 893.148152
49 879.924317
49 897.163460
*The best score is denoted.
A relationship consisting of the four Jamaican
species, four species endemic to Hispaniola + S. insularis, from throughout the Greater Antilles, is represented in all three analyses, although not supported
in the full data set (Fig. 6, clade K). The Jamaican
species are monophyletic, with well-supported interrelationships in all analyses (Fig. 6, clade L). The
sister clade, consisting primarily of Hispaniolan
endemics (three of which are undescribed), is also
well supported (Fig. 6, clade M). However, Hispaniola
has several species outside this clade.
Nodes on the branches subtending other major Caribbean lineages (clades N, O, P and Q in Fig. 6) are
unsupported, although many sister group relationships and one small subclade consisting of three
undescribed species from Hispaniola and one from the
Turks and Caicos Islands (Fig. 6, clade N) are supported in all analyses.
Lagrange analyses
The results of the Lagrange analyses are given in
Table 8. Shown in Figures 7–9 are the maximum like-
lihood reconstructions of range evolution under each
of the six models. The best likelihood score overall
(-121.90) was from model 3B, the vicariance-based
model that includes GAARlandia (Fig. 9B). In models
2A–3B, the best likelihood scores were produced from
those that included GAARlandia. The first two analyses (using models 1A and 1B), which take distance
between islands into account, produced very similar
likelihood scores and maximum likelihood reconstructions, although the model without GAARlandia had
an insignificantly greater likelihood score. However,
in the other two analyses, the differences in likelihood
scores were significant and, in the analyses modelled
with no GAARlandia, there was much more uncertainty in the reconstructions (Figs 7–9 – grey
branches indicate that alternative reconstructions fall
within two log-likelihood units of the scenario that is
depicted). Likelihood ratio tests were used to compare
nested models and, when scenarios were not nested
(e.g. – scenario 2A and scenario 3A), the highest
likelihood score is taken as the best.
DISCUSSION
Unique and shared biogeographic patterns are summarized in Table 9.
SOUTHERN CARIBBEAN
BASIN
Members of the well-supported Southern Caribbean
Basin clade (Aruba, Bonaire, Curaçao, Trinidad and
Tobago, Fig. 6, clade C) are never found within the
larger well-supported Caribbean clade (Fig. 6, clade
A). Geological data often suggest a relationship
between these southern islands, known as the Aruba–
Tobago Belt (Iturralde-Vinent & MacPhee, 1999) and
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
297
Table 6. Likelihoods from the ‘hard and slow’ RAxML analyses to determine the best setting for the number of rate
categories for the data
Rate categories = 10
Rate categories = 25
Rate categories = 40
Rate categories = 55
Starting tree
-ln(L)
-ln(L)
-ln(L)
-ln(L)
1
2
3
4
5
49 876.825172
49 909.100516
49 884.108152
49 878.378649
49 885.284097
49 880.146295
49 877.349169*
49 893.148152
49 879.924317
49 897.163460
49 878.266732
49 877.528777
49 882.377547
49 884.766631
49 877.756447
49 881.349421
49 880.668732
49 885.827844
49 884.610743
49 891.633712
*The best score is denoted.
Table 7. Likelihoods from the MultTrees analyses with
the initial rearrangement setting at 10 and the number of
rate categories set to 25
Tree
-ln(L)
1
2
3
4
5
6
7
8
9
10
49 890.352743
49 875.885667*
49 877.963666
49 876.018067
49 884.591963
49 881.261966
49 879.547932
49 883.400698
49 897.114073
49 879.548262
*The best score is denoted.
indeed the affinities are not surprising given the
proximity of the islands to each other and to the
South American continent. The amphibian and reptile
assemblages on each of these islands are largely continental and also distinct from the primary Caribbean
elements (Hedges, 2006).
In Selenops, this Southern Caribbean clade is
apparently not closely related to other Caribbean
taxa, a pattern found in many other groups, including
mammals (Dávalos, 2004) and plants [orchids (TrejoTorres & Ackerman, 2001)]. However, a contrasting
pattern has been found in Anolis lizards in which the
Southern Caribbean Basin taxa show stronger affinities with the Antilles (Jackman et al., 1999; Creer
et al., 2001); these affinities are hypothesized to have
arisen as a result of the Lesser Antilles being much
further west, and thus closer to Bonaire, in the past
(Creer et al., 2001).
NORTHERN
VS.
SOUTHERN LESSER ANTILLES
A pattern that the Selenops spiders share with
several insects (Wilder & Hollocher, 2003), Anolis
lizards (Gorman & Atkins, 1969; Jackman et al.,
1999; 2002; Creer et al., 2001; Schneider, Losos & de
Queiroz, 2001) and Eleutherodactylus frogs (Kaiser,
Sharbel & Green, 1994), is that species in the Northern Lesser Antilles are only distantly related to
species in the Southern Lesser Antilles. The species
S. n. sp. 7 is found in the Northern Lesser Antilles
from Les Saintes northward to Montserrat and
Antigua, while the species occurring in the Southern
Lesser Antilles, from Dominica south to St Vincent
and the Grenadines (at least to Mayreau) is S. micropalpus. The northern species is nested well within the
strictly Caribbean clade, while S. micropalpus shares
a relationship with S. banksi found from Panamá to
Peru to Guyana. The precise location where northern
and southern lineages are separated is variable, being
slightly to the south in other lineages. For example, in
Anolis, it is between Dominica and Martinique (Losos
& Thorpe, 2004); among Lygaeid bugs (Slater, 1988),
carabid beetles (Liebherr, 1988b), butterflies (Davies
& Bermingham, 2002), Eleutherodactylus frogs
(Kaiser et al., 1994) and populations of the bananaquit (Seutin et al., 1994), it is between St Vincent
and St Lucia. Differences in the location of the boundary between northern and southern lineages may
occur as a result of the timing of colonization of the
different groups, which is likely related to the timing
of emergence of the individual islands. Interestingly,
anoles from the Southern Lesser Antilles, like the
spiders, show affinities with Central and South
American anoles (Jackman et al., 1999; Creer et al.,
2001).
ORIGIN
OF TAXA
The basal taxa for the larger Caribbean clade (Fig. 6,
clade A) are the widely distributed S. lindborgi
(Puerto Rico, Culebra, Vieques, all of the Virgin
Islands, St Kitts, Nevis, eastern Hispaniola and Great
Inagua in the Bahamas, see also Supporting Information, Fig. S1E–G) and the very narrowly distributed
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
298
S. C. CREWS and R. G. GILLESPIE
Figure 4. Likelihood tree resulting from the RAxML analysis of the truncated data set. The map above the tree in
Figure 3 depicts the Caribbean islands and the colours correspond to branches in the tree and indicate on which island
the species is found. Multiple colours along a branch indicate that the species is found on multiple islands. A branch
outlined in black indicates the species is found in Cuba. Selenops radiatus (highlighted in blue) is the type of the genus.
(Isla Mona and Puerto Rico, see also Supporting
Information, Fig. S1F) S. n. sp. 3 (Fig. 6 clade J). A
similar pattern is found among Anolis, with Puerto
Rico endemic Anolis occultus also basal (Jackman
et al., 1999). Likewise, the most basal iguana of the
genus Cyclura is also located on the Puerto Rican
bank (Malone et al., 2000). This pattern, which indicates a common origin of Caribbean diversity for
these groups, is in contrast to data from geckos, frogs,
colubrid snakes and butterflies, which suggest Hispaniola as a centre of diversity (Liebherr, 1988a, and
references therein).
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
299
Figure 5. Tree resulting from the Bayesian analysis of the truncated data set. The map above the tree in Figure 3 depicts
the Caribbean islands and the colours correspond to branches in the tree and indicate on which island the species is found.
Multiple colours along a branch indicate that the species is found on multiple islands. A branch outlined in black indicates
the species is found in Cuba. Selenops radiatus (highlighted in blue) is the type of the genus.
UNIQUE
BIOGEOGRAPHY OF
JAMAICA
Jamaica is one of the oldest islands of the Greater
Antilles, with areas that may have had some parts
continuously above sea level for many millions of
years longer than other islands (Iturralde-Vinent &
MacPhee, 1999; Iturralde-Vinent & Gahagan, 2002).
It is also more isolated than other islands as its last
probable connection with a land mass was likely with
Central America through the Nicaraguan Rise
55 Mya. Our data reflect this isolated history, as
Jamaican species of Selenops form a monophyletic
group of endemics (Fig. 6, clade L). Monophyly of
Jamaican taxa is also present in anoles (Jackman
et al., 1999; Nicholson et al., 2005) and Eleutherodactylus frogs (Hedges, 1996a,b). However, affinities of
the Jamaican clade differ between spider and vertebrate groups: The Jamaican clade of Selenops is supported in the Bayesian and truncated likelihood
analyses as being sister to a clade of primarily Hispaniolan species (Fig. 6, clade K). In contrast, the
Jamaican clade of Eleutherodactylus frogs is most
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300
S. C. CREWS and R. G. GILLESPIE
Figure 6. Bayesian tree with species symbols and asterisks indicative of support removed for clarity. The outgroup taxa
have also been removed. Letters on the nodes indicate clades discussed in the text and in Table 9.
closely related to species from Cuba, while the
Jamaican lineages of Anolis lizards (Nicholson et al.,
2005) and short-faced bats (Dávalos, 2007) are sister
to clades from the mainland. Overall, Jamaica’s
history has been quite different from that of the other
Greater Antillean islands and its fauna may have
accumulated via dispersal and in situ speciation
rather than vicariance (Buskirk, 1985; Crother &
Guyer, 1996).
BIOGEOGRAPHICALLY
DERIVED POSITION OF
CUBA
Cuba has often been depicted as a basal locality in
area cladograms (Buskirk, 1985; Crother & Guyer,
1996). In contrast, although not always supported,
Selenops species from Cuba appear not to be basal, at
least based on morphology and our limited molecular
sampling. Only one species (Selenops aissus – collected from the Bahamas, but that also occurs in
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SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
301
Table 8. Results of the Lagrange analyses for each of the six proposed models
Model
-ln(L)
Dispersal
Extinction
1A
1B
2A
2B
3A
3B
135.5
135.9
144.3
128.1*
260.8
121.9*
0.01634
0.01637
0.02604
0.02137
0.4647
0.1189
0.0082
0.008387
0.008411
0.008094
0.006629
0.008262
–
–
–
–
–
–
dispersal where distance is important; without GAARlandia
dispersal where distance is important; with GAARlandia
dispersal where distance is not important; without GAARlandia
dispersal where distance is not important; with GAARlandia
little to no over-water dispersal; without GAARlandia
little to no over-water dispersal; with GAARlandia
Model 1 is a dispersal-based model in which the distance between land masses is considered.
Model 2 is a dispersal-based model in which the distance between land masses is not taken into account.
Model 3 is a vicariance-based model in which dispersal probabilities are very low if dispersal must occur over water.
The ‘A’ portion of each model was run without GAARlandia, while the ‘B’ portion was run with GAARlandia.
The dispersal and extinction values are the maximum likelihoods estimates for the rate of each process and represent the
mean number of events per unit of branch length.
*Hypotheses that were statistically different from the null hypothesis of ‘no GAARlandia’ are marked.
Figure 7. Maximum likelihood reconstruction of geographic range evolution under a dispersal-based model where
distance between land masses is taken into account. Single-area ancestral ranges are shown at nodes. Grey branches
indicate that alternative reconstructions fall within two log-likelihood units of the scenario that is depicted. Range
transitions along branches show sequences of dispersal and extinction events. C, Central America; S, South America; G,
Greater Antilles; N, Northern Lesser Antilles; A, Southern Lesser Antilles. (A) without GAARlandia; (B) with GAARlandia.
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
302
S. C. CREWS and R. G. GILLESPIE
Figure 8. Maximum likelihood reconstruction of geographic range evolution under a dispersal-based model where
distance between land masses is ignored. Single-area ancestral ranges are shown at nodes. Grey branches indicate that
alternative reconstructions fall within two log-likelihood units of the scenario that is depicted. Range transitions along
branches show sequences of dispersal and extinction events. C, Central America; S, South America; G, Greater Antilles;
N, Northern Lesser Antilles; A, Southern Lesser Antilles. (A) without GAARlandia; (B) with GAARlandia.
Cuba) occurs at the base of an internal clade, while all
other sampled species which occur in Cuba (although
all but one – Selenops submaculosus – were collected
from other islands) are nested high within the trees
(S. submaculosus, Selenops simius, Selenops inuslaris) and it is inferred based on morphology (S. C.
Crews & R. G. Gillespie, unpubl. data) that most of
the Cuban endemics are closely related to S. simius
and S. submaculosus (Fig. 6, clade O).
GREATER ANTILLES
and St Maarten and Anguilla are nested well within
a clade of Hispaniolan animals (Fig. 6, clade O). This
suggests that S. n. sp. 7 and S. n. sp. 8 colonized the
Northern Lesser Antilles region from Hispaniola and
thus support the Greater Antilles as a centre of
species diversity via dispersal events in Anolis (Glor,
Losos & Larson, 2005). Also, many species of Selenops
in Hispaniola have very small ranges that mirror
those of many endemic anoles from the Anolis cybotes
group (Glor et al., 2003), indicating similar patterns
of speciation between the two groups.
AS A CENTRE OF
SPECIES DIVERSITY
There are two additional patterns that appear to be
shared between Selenops and Anolis. In Selenops
spiders, the species from the Northern Lesser Antilles
SPECIES–AREA
RELATIONSHIPS
In many taxa as diverse as fungi (Lodge, Baroni &
Cantrell, 2002), vertebrates (Ricklefs & Lovette,
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SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
303
Figure 9. Maximum likelihood reconstruction of geographic range evolution under a vicariance-based model where there
are very low probabilities of over-water dispersal. Single-area ancestral ranges are shown at nodes. Grey branches
indicate that alternative reconstructions fall within two log-likelihood units of the scenario that is depicted. Range
transitions along branches show sequences of dispersal and extinction events. C, Central America; S, South America; G,
Greater Antilles; N, Northern Lesser Antilles; A, Southern Lesser Antilles. (A) without GAARlandia; (B) with GAARlandia.
1999) and invertebrates (Nichols, 1988), the Greater
Antilles harbour more species than the Lesser Antilles. This can be attributed largely to island area
(MacArthur & Wilson, 1963) and associated habitat
diversity and age (Losos, 1996; Ricklefs & Bermingham, 2002, 2008). In Selenops, the same pattern is
found, with larger, older islands (Greater Antilles)
having more species than smaller, younger, less
habitat-diverse islands (Lesser Antilles). In the
Greater Antilles there is often a pattern of number
of species in Cuba > Hispaniola > Jamaica > Puerto
Rico, based on island size. This pattern also prevails
in Selenops. In this genus, 17 species occur in Cuba
with 12 endemics (Alayón, 2005), while in Hispaniola
there are at least 16 species with 11 endemics and, in
Jamaica, at least five species are known, with four
endemics. However, the Bahamas have no known
endemic species of Selenops.
HYPOTHESIS
TESTING
In the maximum likelihood analyses of range expansion, likelihood ratio tests of scores for the scenarios
that include the existence of the GAARlandia land
span are either equally probable or more favourable
than those that do not. This does not mean that
over-water dispersal has not occurred, but rather that
land bridges hold a stronger signature on the phylogeny. These results contrast to those for mammals in
which there was little to no support for a land span
between the Greater Antilles and northern South
America (Dávalos, 2004). Likewise, Hedges and
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
304
S. C. CREWS and R. G. GILLESPIE
Table 9. Biogeographic patterns in Caribbean Selenops species
Shared with
other taxa?
Pattern
1. Distinct, distantly related SLA and
NLA clades
Yes
2. Monophyly of Jamaican taxa
3. More than one colonization and
diversification in Hispaniola
4. Patterns of endemism throughout the
Caribbean (GA harbour more
endemics than LA)
5. Endemic species in Hispaniola with
distributions that overlap endemics of
other taxa
6. Southern Netherlands Antilles form a
clade with Trinidad and Tobago
exlcusive of other Caribbean taxa
7. Location of the split between the NLA
and SLA is between Dominica and
Les Saintes, Guadeloupe
8. Cuban species are not basal in the
Caribbean clade
Yes
Yes
Clades that show
particular patterns
Which taxa?
Lygaeid bugs, Carabid beetles,
fruitflies, butterflies, Anolis,
Eleutherodactylus, bananaquit
Anolis, Eleutherodactylus
Anolis
F and O
L
A, J, M, N
Yes
Fungi, Anolis, birds, carabid
beetles
LA – F,O
GA – J,K,L,M,N,P,Q
Yes
Anolis
N, P, others in clade A
No
–
C
No
–
SLA – F
NLA – O
No
–
Q, S. aissus
If a pattern is shared with other taxa, the taxa are noted. Clades referenced are those that display the patterns mentioned
here and are depicted in Figure 6.
GA, Greater Antilles; LA, Lesser Antilles; NLA, Northern Lesser Antilles, SLA, Southern Lesser Antilles.
others (Hedges, Hass & Maxon, 1992; Hedges,
1996a,b; Hedges & Heinicke, 2007; Heinicke et al.,
2007) found that molecular clock estimates of divergence times precluded a major role of land bridges in
the origin of Caribbean herpetofauna. The suitability
Dispersal–Extinction–Cladogenesis model of geographic range evolution used here, in which dispersal
events cause range expansion, local extinction events
cause range contraction and the probability of each
kind of event is proportional to the branch length, has
been questioned for island fauna, as terminal taxa
may be restricted to single islands (Ree & Smith,
2008). However, in our models, islands were either
grouped together or several species were spread
across multiple islands and thus the model is reasonable in this particular case.
CONCLUSIONS
The current study provides a basis for biogeographic
comparison across different lineages in the Caribbean. It is one of the most extensive data sets for
Caribbean fauna and the most comprehensive
molecular data set of any spider group. While inclusion of taxa currently missing from our analyses, and
possibly the use of other markers, may help resolve
basal relationships, it should also be noted that deep,
short branches, such as those found here, may be very
difficult if not impossible to resolve (Degnan & Salter,
2005; Kubatko & Degnan, 2007). The results reveal
several patterns common to other disparate taxa, as
well as many unique patterns which warrant further
study. Moreover, the data set provides the groundwork for behavioural, ecological and population-level
studies similar to lineages such as Anolis lizards
(Losos, 2009) and passerine birds (Ricklefs & Bermingham, 2007).
ACKNOWLEDGEMENTS
We would like to acknowledge members of S.C.C.’s
dissertation committee for their guidance: George
Roderick, Jim McGuire and Charles Griswold. We
would like to thank the following museums, curators
and collection managers for specimen loans: American
Museum of Natural History – Norman I. Platnick and
Louis Sorkin; Museum of Comparative Zoology –
Laura Leibensperger; California Academy of Sciences
– Charles Griswold; National Museum of Natural
History – Jonathan Coddington; British Museum of
Natural History – Janet Beccaloni; Peabody Musem
at Yale – Raymond Pupedis; Essig Museum of Ento-
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
mology – Cheryl Barr; Museo Nacional de Histora
Natural, Santo Domingo – Sardis Medrano Cabral,
Carmelo Nuñez. We would also like to thank Jim
McGuire for use of the MVZ cluster and Mark Miller
and Lucie Chan for use of the SGE cluster and the
CIPRES portal and the San Diego Supercomputer
Center. We are extremely grateful to Richard Ree for
quickly responding and helping us to understand and
implement Lagrange. We would also like to thank
Matt Brandley for phylogenetic methodology discussions. We are grateful to all of the many people that
aided us in obtaining permits and collecting: Kelvin
Guerrero, Denia Veloz, Eladio Fernandez, Alberto
Puente-Rolón, Beverly Mae Nisbeth, Adriel Thibou,
Germain George, Brian Riggs, Brian Manco, Margaret Jones, Renata Platenberg, Chris Niebuhr, Abel
Pérez-González, G. B. Edwards, Oscar Francke, Alejandro Mondragon, Mark da Silva, Facundo Franken,
Roy Croes, Gijs Van Hoorn, Adolphe O. Debrot, Mark
Vermeij, Fred the Abaco Caveman, Raveen Gibson,
Daniel Palmer, Jim Starrett, Marshal Hedin, Nicole
VanderSal, Sean Schoville, Luke Mahler, Uri García,
Beto Mendoza, Adrian Nieto Montes de Oca, Rebecca
Duncan, Pierre Paquin, Matthew Cottam, Jan den
Dulk, Joey Slowik, Nicole Esteban, Arturo Herrera,
Nancy Bottomley, Inilek Wilmot, Lauren Esposito,
Stephen Touissant, Arlington James, Ferdinand
Tripoli, Daniel Memia Zolo, Nouree-Yvon, Martín
Ramírez, Mark Harvey, Volker Framenau, Jeremy
Miller, Hannah Wood, Yuri Marusik, Caroline
Chaboo, Cheryl Barr, Bill Shepard, Akio Tanikawa, C.
J. Hayden, Aaron Abdel, Dan Warren. Finally, we
would like to acknowledge Matjaz Kunter and an
anonymous reviewer for their straightforward and
constructive reviews of this paper. Funding was provided by the Schlinger Foundation, with additional
support from the Margaret C. Walker Fund.
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SUPPORTING INFORMATION
Additional Supporting Information may be found in the online version of this article:
Figure S1. Expansion of the boxed area in Figure 1 of the main text, divided into regions depicted in the
Figure S2A–I, showing the detailed locality data.
Figure S2. Collecting localities from the Caribbean region, including most islands and several sites throughout
Mexico, Central America and the South American mainland.
Please note: Wiley-Blackwell are not responsible for the content or functionality of any supporting materials
supplied by the authors. Any queries (other than missing material) should be directed to the corresponding
author for the article.
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
309
APPENDIX
Collecting localities and voucher numbers of all animals used in this study. Locality numbers refer to numbers
in the Supporting Information (Figs S1 and S2).
Locality
number
Species
Collection information
Voucher numbers
1
Selenops n. sp. 5
sel_069, sel_070
2
Selenops n. sp. 5
3
Selenops n. sp. 5
4
Selenops curazao
5
Selenops curazao
6
Selenops curazao
7
Selenops curazao
8
Selenops curazao
9
Selenops willinki
Aruba: Bringamosa, house of Roy Croes,
12°39.547′N, 69°58.077′W, 14.X.2004,
SCC04_041
Aruba: Luela, shooting range, 12°29.023′N,
69°57.778′W, 16.X.2004, SCC04_044
Aruba:N. sp. 5 National Park, near hill Gran
Tonel and Valley Rooi Coashati, 12°29.356′N,
69°55.461′W, 16.X.2004, SCC04_045
Curaçao: Carmabi Institute, 12°07.351′N,
68°58.132′W, 7.X.2004, SCC04_026
Curaçao: Girouette Plantation, E of Schottegat
Harbour, house of A. DeBrot, 12°09′03.20″N,
68°54′56.35″W, 9.X.2004, SCC04_032
Bonaire: Sabadaco, near cave, 12°11.587′N,
68°17.765′W, 11.X.2004, SCC04_033
Bonaire: Altamira Ungu, 12°13.949′N,
68°20.703′W, 12.X.2004, SCC04_038
Bonaire: Nort di Saliña, Kaya Otomac,
12°10′55.92″N, 68°16′28.39″W, 11.X.2004,
SCC04_039
Trinidad and Tobago: Little Tobago,
11°18′03.7″N, 60°30′11.1″W, 16.VII.2005,
SCC05_041
10
Selenops
geraldinae
11
Selenops
geraldinae
12
Selenops
geraldinae
13
Selenops
geraldinae
14
Selenops
geraldinae
15
Selenops
hebraicus
Selenops occultus
16
Trinidad and Tobago: St George Co., Point
Gourde Road near Trinidad Military Base,
near Chaguaramas off Chaguaramas Main
Road, 10°40′47.1″N, 61°37′30.9″W,
11.VII.2005, SCC05_035
Trinidad and Tobago: Gaspar Grande Island
south of Chaguaramas, trails around island,
10°39′46.5″N, 61°38′58.0″W, 12.VII.2005,
SCC05_037
Trinidad and Tobago: Monos Island, South Sea,
10°40′54.2″N, 61°41′21.6″W, 13.VII.2005,
SCC05_038
Trinidad and Tobago: Huevos Island,
10°41′28.3″N, 61°42′55.0″W, 13.VII.2005,
SCC05_039
Trinidad and Tobago: Chacachacare Island,
10°41′24.2″N, 61°44′53.7″W, 13.VII.2005,
SCC05_040
Argentina: Parque Nacional Chaco, Sendero
peatonal. 25.II.2004.
Brazil: São Paulo, Universidade São Paulo,
23°34′27.96″S, 46°40′21.29″W
sel_072
sel_068
sel_047, sel_058, sel_217
sel_048, sel_049, sel_050, sel_051,
sel_052
sel_053
sel_059, sel_060, sel_082
sel_054, sel_056, sel_057, sel_061,
sel_062, sel_063, sel_064, sel_065
sel_230, sel_231, sel_232, sel_233,
sel_234, sel_235, sel_236,
sel_238, sel_240, sel_242,
sel_243, sel_245, sel_251,
sel_252, sel_253, sel_255,
sel_258, sel_259, sel_261,
sel_262, sel_263
sel, 218, sel_219, sel_220, sel_221,
sel_222, sel_223
sel_224, sel_225, sel_226, sel_227,
sel_228, sel_229, sel_237,
sel_241, sel_257
sel_244, sel_247, sel_248, sel_249,
sel_250
sel_239
sel_246
MACN-Ar#12782
sel_995
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
310
S. C. CREWS and R. G. GILLESPIE
APPENDIX Continued
Locality
number
Species
Collection information
Voucher numbers
17
Selenops occultus
sel_284
18
Selenops occultus
18
Selenops
melanurus
Selenops
micropalpus
Brazil: Rio de Janeiro, Sierra da Carioca,
22°57′52.22″S, 43°16′31.34″W
Brazil: Rio de Janeiro, Pao de Açucar,
22°55′22.05″S, 43°09′33.11″W
Brazil: Rio de Janeiro, Pao de Açucar,
22°55′22.05″S, 43°09′33.11″W
St Vincent and the Grenadines: Young Island,
South of Villa Beach, 13°07.895′N,
61°12.142′W, 24.X.2004, SCC04_053
St Vincent and the Grenadines: King’s Hill
Forest Reserve, 13°08.825′N, 61°10.021′W,
27.X.2004, SCC04_055
St Lucia: Vieux Fort, on hill above airport,
13°44′20.2″N, 60°56′40.8″W, 13.III.2007
St Lucia: Anse la Ray, Ti-Kaye, 13°55′29.6″N,
61°02′41.3″W, 13.III.2007, SCC07_046
St Lucia: Dennery: south of Dennery on east
coast road, eastern nature trail, heritage
tourism site, 13°53′50.8″N, 60°52′51.2″W,
13.III.2007, SCC07_047
St Lucia: Gros Islet, Pigeon Island, on top of
hill, 14°05′31.3″N, 60°57′03.8″W, 12.III.2007,
SCC07_044
St Lucia: Gros Islet, Beausejour, past cricket
sponsor’s office, 14°04′43.1″N, 60°56′31.1″W,
12.III.2007, SCC07_045
Martinique: Le Diamant, Grand Anse du
Diamant, off of road D37, 14°28′32.9″N,
61°02′13.4″W, 10.III.2007, SCC07_043
Martinique: La Caravelle Reserve Naturelle,
trail to Pointe Caricoli, 14°46′09.3″N,
60°53′24.7″W, 8.III.2007, SCC07_041
Martinique: Anse Ceran, off of road D-10,
14°50′01.5″N, 61°13′24.7″W, 9.III.2007,
SCC07_042
Dominica: Roseau: Botanical Park, top of hill
near shrine, 15°17.998′N, 61°22.754′W,
1.XI.2004, SCC04_059
Dominica: Jimmit-Warner, on top of hill with
cellular phone antenna, 15°22.690′N,
061°24.003′W, 2.XI.2004, SCC04_061
Dominica: Cabrits National Park, near trail to
fort, 15°35.049′N, 61°28.371′W, 2.XI.2004,
SCC04_060
Guadeloupe: Les Saintes, top of Le Chameau,
15°51′28.1″N, 61°35′39.8″W, 6.III.2007,
SCC07_039
Guadeloupe: Basse-Terre, Parc Archélogique
des Roches Gravées, near Trois-Rivieres,
15°58.394′N, 61°38.347′W, 10.XI.2004,
SCC04_064
19
20
Selenops
micropalpus
21
Selenops
micropalpus
Selenops
micropalpus
Selenops
micropalpus
22
23
24
Selenops
micropalpus
25
Selenops
micropalpus
26
Selenops
micropalpus
27
Selenops
micropalpus
28
Selenops
micropalpus
29
Selenops
micropalpus
30
Selenops
micropalpus
31
Selenops
micropalpus
32
Selenops n. sp. 7
33
Selenops n. sp. 7
sel_283
sel_277, sel_278, sel_279, sel_280,
sel_281, sel_282
sel_091, sel_113
sel_088, sel_090, sel_092, sel_093,
sel_094, sel_095, sel_112
sel_832, sel_833, sel_834
sel_820, sel_821, sel_822, sel_823
sel_825, sel_826, sel_827, sel_828,
sel_829, sel_830, sel_831
sel_811, sel_812, sel_813, sel_814
sel_815, sel_816, sel_817, sel_818,
sel_819
sel_804, sel_805, sel_806, sel_807,
sel_808, sel_809, sel_810
sel_791, sel_792, sel_793, sel_794,
sel_795, sel_797, sel_798, sel_799
sel_800, sel_801, sel_802, sel_803
sel_097, sel_098, sel_099, sel_100,
sel_101
sel_102, sel_103, sel_104, sel_105
sel_106, sel_107, sel_108, sel_110,
sel_111, sel_135
sel_783, sel_784, sel_785
sel_114
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
311
APPENDIX Continued
Locality
number
Species
Collection information
Voucher numbers
34
Selenops n. sp. 7
sel_778, sel_779, sel_780, sel_781
35
Selenops n. sp. 7
36
Selenops n. sp. 7
37
Selenops n. sp. 7
Guadeloupe: Basse-Terre, Trois Rivieres, end of
Trail Sentier de l′Acomat off Rue Nelson
Mandela, 15°58′03.0″N, 61°37′50.1″W,
5.III.2007, SCC04_038
Guadeloupe: Basse-Terre, near Vieux Fort on
the D6, along road at Forêt Domaniale du
Litoral, 15°57.943′N, 061°42.517′W,
11.XI.2004, SCC04_065
Guadeloupe: Gran-Terre, Pointe du Chateaux,
16°14′51.6″N, 61°11′02.6″W, 7.III.2007,
SCC07_040
Montserrat: Jack Boy Hill, 17°46′02.1″N,
62°10′17.0″W, 2.III.2007, SCC07_035
38
Selenops n. sp. 7
39
Selenops n. sp. 7
40
Selenops n. sp. 7
41
Selenops n. sp. 7
42
Selenops
lindborgi
43
Selenops
lindborgi
44
Selenops
lindborgi
45
Selenops
lindborgi
46
Selenops n. sp. 8
47
Selenops n. sp. 8
48
Selenops n. sp. 8
49
Selenops n. sp. 8
Montserrat: Sweet Water Ghaut, 16°47′07.2″N,
62°10′59.8″W, 2.III.2007, SCC07_036
Montserrat: Silver Hills, north side of Silver
Hill, 16°48′41.3″N, 62°11′28.7″W, 3.III.2007,
SCC07_037
Antigua: Nelson’s Dockyard National Park,
Shirley’s Heights lookout, 17°00′06.7″N,
61°44′57.6″W, 27.II.2007, SCC07_034
Antigua: Indian Town, east of Veranda Resort,
17°05′50.2″N, 61°40′53.0″W, 27.II.2007,
SCC07_033
St Kitts and Nevis: Nevis, Tamarind Bay,
Galliput Restaurant, 17°09′48.53″N,
62°37′50.02″W, 23.II.2007, SCC07_030
St Kitts and Nevis: Nevis, Round Hill entrance
to Mt Nevis, 17°11′13.83″N, 62°36′00.96″W,
23.II.2007, SCC07_029
St Kitts and Nevis: St Kitts, Major’s Bay,
17°13′37.9″N, 62°38′49.3″W, 24.II.2007,
SCC07_032
St Kitts and Nevis: St Kitts, Sand Bank Bay,
17°14′59.1″N, 62°38′40.8″W, 24.II.2007,
SCC07_031
Saba: Giles Quarter Trail, 17°36′54.36″N,
63°14′35.52″W, 12.III.2008
St Maarten: Mullet Bay, abandoned Mullet
Bay Resort, 18°02′48.0″N, 63°027′29.7″W,
21.III.2007, SCC07_027
St Maarten: Emilio Wilson Estate and Park,
18°02′32.7″N, 63°03′53.1″W, 20.II.2007,
SCC07_026
St Maarten: Upper Princess Quarter,
18°01′48.0″N, 63°02′08.0″W, 21.III.2007,
SCC07_028
sel_115
sel_786, sel_787, sel_788, sel_789,
sel_790
sel_758, sel_759, sel_760, sel_761,
sel_762, sel_763, sel_764,
sel_765, sel_766, sel_767, sel_768
sel_769
sel_771, sel_772, sel_773, sel_774,
sel_775, sel_776, sel_777
sel_754, sel_755, sel_756, sel_757
sel_753
sel_740, sel_741, sel_742
sel_735, sel_736, sel_737, sel_738,
sel_739
sel_750, sel_751, sel_752
sel_743, sel_745, sel_746, sel_747,
sel_748, sel_749
sel_1021, sel_1022
sel_712, sel_713, sel_718, sel_719,
sel_720, sel_721, sel_722,
sel_723, sel_724, sel_725,
sel_726, sel_727, sel_728,
sel_729, sel_730
sel_711, sel_714, sel_715, sel_716,
sel_717
sel_731, sel_732
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
312
S. C. CREWS and R. G. GILLESPIE
APPENDIX Continued
Locality
number
Species
Collection information
Voucher numbers
50
Selenops n. sp. 8
sel_116, sel_119, sel_120, sel_121,
sel_708, sel_709, sel_710
51
Selenops
mexicanus
52
Selenops n. sp. 8
53
Selenops n. sp. 8
54
Selenops n. sp. 8
55
Selenops n. sp. 8
56
Selenops n. sp. 8
57
Selenops
lindborgi
58
Selenops
lindborgi
59
Selenops
lindborgi
60
Selenops
lindborgi
61
Selenops
lindborgi
62
Selenops
lindborgi
Selenops
insularis
Selenops
lindborgi
Selenops
insularis
Selenops
lindborgi
Selenops
insularis
Selenops
lindborgi
St Maarten: south-east side of island, trail
from Back Bay to Geneve Bay, 18°00.929′N,
63°01′840W, 12–13.XI.2004 and 20.II.2007,
SCC04_066, 068, SCC07_025
St Maarten: Philipsburg, Front Street, near
entrance to cruise ship dock, 18°00.906′N,
63°02.587′W, 12.XI.2004, SCC04_067
Anguilla: Shoal Bay West, 18°09′52.8″N,
63°09′21.3″W, 13.II.2007, SCC07_016
Anguilla: The Cove, 18°10′14.1″N,
63°07′52.6″W, 13.II.2007, SCC07_017
Anguilla: Long Bay, Long Bay Beach,
18°11′29.3″N, 63°07′49.7″W, 13.II.2007,
SCC07_015
Anguilla: Blowing Point, 18°10′18.0″N,
63°05′28.7″W, 13.II.2007, SCC07_018
Anguilla: Windward Point, 18°16′18.2″N,
62°58′05.3″W, 12.II.2007, SCC07_014
USVI: St Croix: Fredericksted, Sprat Hall
Beach, Rte.63, 17°44′09.8″N, 64°53′24.0″W,
14.VI.2006, SCC06_056
USVI: St Croix: Sprat Hall Hill off West Shore
Road, first right after subtracking station,
17°44′38.4″N, 64°53′22.3″W,14.VI.2006
SCC06_055
USVI: St Croix: Butler Bay, West Shore Road,
17°45′49.7″N, 64°52′58.8″W, 14.VI.2006,
SCC06_054
USVI: St Croix: intersecting road from Creque
Dam to Mahogany Road (intersects Mount
Victory Camp) 17°44′27.1″N, 64°51′25.4″W,
14.VI.2006, SCC06_057
USVI: St Croix: Radio Telescope Station, east
island, 17°45.398′N, 64°35.045′W, 18.XI.2004,
SCC04_072
Puerto Rico: Vieques: Laguna Kiani,
18°07′02.2″N, 65°33′41.4″W, 19.VI.2006,
SCC06_065
63
64
65
sel_117, sel_118
sel_697
sel_703, sel_704
sel_698, sel_699, sel_700, sel_701,
sel_702
sel_705, sel_706, sel_707
sel_696
sel_497, sel_498, sel_499, sel_500
sel_494, sel_495, sel_496
sel_490, sel_491, sel_492, sel_493
sel_501
sel_527
sel_522, sel_523, sel_524, sel_525,
sel_526
Puerto Rico: Vieques: Ruinas Central Playa
Grande, 18°05′43.2″N, 65°31′13.2″W,
19.VI.2006, SCC06_064
sel_516, sel_517, sel_518, sel_519,
sel_520, sel_521
Puerto Rico: Vieques: Cano Hondo, Cerca de
Puerto Mosquito, 18°06′11.0″N, 65°27′05.5″W,
19.VI.2006, SCC06_061
sel_504, sel_505, sel_506, sel_507,
sel_508, sel_509, sel_510,
sel_511, sel_512
Puerto Rico: Vieques: Refugio Nacional de Vida
Silvestre, road to Playa Caracas, Laguna
Puerto Ferro, 18°06′24″N, 65°25′25.8″W,
19.VI.2006, SCC06_063
sel_513, sel_514, sel_515
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
313
APPENDIX Continued
Locality
number
Species
Collection information
Voucher numbers
66
Selenops
lindborgi
sel_480, sel_481, sel_482, sel_483,
sel_484, sel_485
67
Selenops
lindborgi
68
Selenops
lindborgi
69
Selenops
lindborgi
70
Selenops
lindborgi
71
Selenops
lindborgi
72
Selenops
lindborgi
73
Selenops
lindborgi
74
Selenops
lindborgi
75
Selenops
lindborgi
76
Selenops
lindborgi
77
Selenops
lindborgi
78
Selenops
lindborgi
79
Selenops
lindborgi
80
81
Selenops
submaculosus
Selenops simius
Puerto Rico: Culebra: Monte Resaca,
18°19′30.7″N, 65°18′10.5″W, 12.VI.2006,
SCC06_052
Puerto Rico: Culebra: Brava Beach Trail,
18°19′38.9″N, 65°16′54.1″W, 12.VI.2006,
SCC06_053
USVI: St Thomas, Estate Perserverance,
Perserverance Bay Trail, 18°21.463′N,
64°59.753′W, 22.X.2004, SCC04_050
USVI: St Thomas, St Peter, house of R.
Platenberg, 18°21′22.17″N, 64°56′49.53″W,
23.X.2004, SCC04_075
USVI: St Thomas, Magen’s Bay Trail,
18°21.350′N, 64°55.231′W, 22.X.2004 and
9.XI.2004, SCC04_052 and SCC04_073
USVI: St Thomas, Benner Hill, above armory,
18°19.533′N, 64°51.703′W, 19.XI.2004,
SCC04_074
USVI: St Thomas, East End, Estate Nazareth,
Dolphin House, 18°19.128′N, 64°51.567′W,
19.XI.2004, SCC04_076
USVI: St John, Bordeaux Ridge Road,
18°20.125′N, 64°43.672′W, 17.XI.2004,
SCC04_071
USVI: St John, Cinnamon Bay Loop Trail,
18°21.226′N, 64°45.259′W, 16-17.XI.2004,
SCC04_070
USVI: St John, Leinster Bay Trail,
18°21.825′N, 64°43.743′W, 16.XI.2004,
SCC04_069
BVI: Tortola: vic. Sage Mountain,
18°24′46.44″N, 64°39′18.43″W, 20.X.2004,
SCC04_049
BVI: Guana Island: north side near beach
house, 18°28.793′N, 64°34.473′W, 18.X.2004,
SCC04_047
BVI: Guana Island: south side near salt pond,
18°28.619′N, 64°34.475′W, 18.X.2004,
SCC04_046
BVI: Virgin Gorda, lower trail up Gorda Peak,
18°28.774′N, 64°24.210′W, 19.X.2004,
SCC04_048
Cuba: Sierra de Mesa, Pinar del Rio
82
Selenops simius
Cayman Islands: Grand Cayman, Queen
Elizabeth II Botanic gardens, storage facility
south of main building, 19°19.055′N,
81°09.527′W, 30.IX.2004, SCC04_021
Cayman Islands: Grand Cayman, Queen
Elizabeth II Botanic gardens, tree trail
behind iguanas, 19°19.042′N, 81°10.081′W,
2.X.2004, SCC04_022
sel_486, sel_487, sel_488, sel_489
sel_076
sel_131
sel_124, sel_125, sel_128, sel_129,
sel_130, sel_164
sel_123, sel_132, sel_133, sel_134
sel_143, sel_144, sel_145, sel_146,
sel_147
sel_140, sel_141, sel_142, sel_157
sel_126, sel_127
sel_136, sel_137, sel_138, sel_139
sel_078
sel_071, sel_081, sel_085, sel_089
sel_079, sel_084, sel_086, sel_087
sel_073, sel_074, sel_075, sel_083,
sel_089, sel_096, sel_109
sel_276
sel_046, sel_066, sel_067
sel_080
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
314
S. C. CREWS and R. G. GILLESPIE
APPENDIX Continued
Locality
number
Species
Collection information
Voucher numbers
83
Selenops simius
sel_022
84
Selenops simius
85
Selenops
candidus
86
Selenops
candidus
87
Selenops
candidus
88
Selenops n. sp.
17
89
Selenops
candidus
90
Selenops
candidus
91
Selenops
candidus
92
Selenops
petrunkevitchi
Cayman Islands: Little Cayman, on road
across street from Pirate’s Point Resort,
19°39.754′N, 80°06.032′W, 3.X.2004,
SCC04_023
Cayman Islands: Cayman Brac, National Trust
House off West End Road, 19°42.019′N,
79°52.084′W, 3.X.2004, SCC04_025
Jamaica: Westmoreland Paris, near New Hope
on road toward Savanna-la-Mar,
18°14′55.4″N, 78°14′41.0″W, 29.V.2006,
SCC06_027
Jamaica: St Ann Parish, North Coast Highway,
between Discovery Bay and Rio Bueno,
18°28′31.3″N, 77°25′49.0″W, 28.V.2006,
SCC06_024
Jamaica: Clarendon Parish, off road to
Lluidasvale, 18°07′50.8″N, 77°10′05.0″W, 31
May 2006, SCC06_028
Jamaica: St Catherine Parish, Hellshire Hills,
A2 Depression, 17°51′59.3″N, 76°57′54.0″W,
3.VI.2006, SCC06_031
Jamaica: St Mary Parish, near Mango Valley,
1.6 km off North Coast Highway,
18°24′23.4″N, 77°02′37.6″W, 28.V.2006,
SCC06_023
Jamaica: St Andrew Parish, Castleton Botanic
Gardens, 18°10′20.3″N, 76°49′27.6″W,
27.V.2006, SCC06_022
Jamaica: St Andrew Parish, Hermitage Dam
Road, 2–6 km from junction with Stony Hill,
18°04′25.4″N, 76°47′01.3″W, 5.VI.2006,
SCC06_033
Jamaica: St Thomas Parish, Blue Mountains
National Park, Whitfield Hall, 18°02′54.8″N,
76°37′03.7″W, 1.VI.2006, SCC06_029
93
Selenops n. sp.
16
94
Selenops
insularis
Selenops
pensilis
Selenops n. sp.
13
95
96
Selenops
marcanoi
Jamaica: St Thomas Parish, near 12 mile Bull
Bay, on left side of road heading east,
17°55′32.5″N, 76°38′31.0″W, 5.VI.2006,
SCC06_034
Hispaniola: Haiti: Jacmel, St Cyr 72°31′41.2″N,
18°14′16.6″W, 23.X.2006, SCC06_078
Hispaniola: Dominican Republic: Prov.
Pedernales: road to Aguacate from Rio
Mulito, 18°13.895′N, 71°45.190′W,
25.XI.2004, SCC04_082
Hispaniola: Dominican Republic: Prov.
Pedernales, Rio Mulito (El Banano),
18°09.165′N, 071°45.388′W, 25.XI.2004,
SCC04_081
sel_023, sel_024, sel_025, sel_026,
sel_027, sel_028
sel_362
sel_357, sel_358, sel_359, sel_360
sel_363, sel_364
sel_376, sel_377, sel_378, sel_379,
sel_380, sel_381, sel_382,
sel_383, sel_384
sel_352, sel_353, sel_354, sel_355,
sel_356
sel_350, sel_351
sel_385
sel_365, sel_366, sel_367, sel_368,
sel_369, sel_370, sel_371,
sel_372, sel_373, sel_374,
sel_375, sel_376
sel_386, sel_387, sel_388, sel_389,
sel_390, sel_391
sel_655, sel_656, sel_657, sel_658,
sel_659, sel_660, sel_661,
sel_662, sel_663, sel_664, sel_665
sel_156
sel_150, sel_151, sel_152, sel_153,
sel_154, sel_155
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
315
APPENDIX Continued
Locality
number
Species
Collection information
Voucher numbers
97
Selenops phaselus
sel_158, sel_160, sel_632, sel_633,
sel_634
98
Selenops phaselus
99
Selenops n. sp.
14
Hispaniola: Dominican Republic: Prov.
Pedernales, Parque Nacional Sierra de
Baoruco, Las Abejas, 18°08.804′N,
71°37.164′W, 24-Nov-04 and 10-Oct-06,
SCC04_077 and SCC06_072
Hispaniola: Dominican Republic: Prov.
Pedernales, 26 km north of Cabo Rojo, Sierra
de Baoruco, 18°06.490′N, 71°37.316′W,
24.XI.2004, SCC04_078
Hispaniola: Dominican Republic: Prov.
Pedernales, 13–14 km north of Cabo Rojo on
Carretera ALCOA, 18°01.962′N, 71°38.748′W,
24.XI.2004, SCC04_079
Hispaniola: Dominican Republic: Prov.
Pedernales, Parque Jaragua, VII.2006
Hispaniola: Dominican Republic: Prov.
Pedernales, Boca de la Cañada,
Pedernales-N. sp. 15 Road, 9.X.2006,
SCC06_071
Hispaniola: Dominican Republic: Prov.
Pedernales, N. sp. 15, Fondo de Mama
Cocoño, 25.X.2003, SCC03_021
Hispaniola: Dominican Republic, Prov.
Pedernales, Laguna N. sp. 15, El Cajuil,
9.X.2006, SCC06_070
Hispaniola: Dominican Republic: Prov.
Barahona, Carretera Higuero-Polo,
26.XI.2004, SCC04_083
Hispaniola: Dominican Republic: Prov.
Barahona, Polo Coffee Plantation, July 2006
Hispaniola: Dominican Republic: Prov.
Barahona, Barahona, CoralSol Resort, San
Rafael Beach, July 2006
sel_537, sel_538, sel_539, sel_540,
sel_541, sel_542, sel_543,
sel_544, sel_545, sel_561,
sel_563, sel_564, sel_567,
sel_573, sel_574, sel_575,
sel_576, sel_587, sel_588
sel_175
100
101
102
Selenops
insularis
Selenops n. sp.
14
Selenops n. sp.
15
Selenops
insularis
103
Selenops n. sp.
15
104
Selenops
insularis s. n.
sp. 1
Selenops phaselus
Selenops n. sp.
13
Selenops
insularis
105
106
107
Selenops
insularis
108
Selenops
insularis
109
Selenops
insularis
110
Selenops
lindborgi
Hispaniola: Dominican Republic: Prov. Peravia,
Bani, road from Bani to Manaclar, past La
Laguna, 18°21.343′N, 70°21.077′W,
27.XI.2004, SCC04_086
Hispaniola: Dominican Republic: Prov. Peravia,
Bani, Rio Nizao, 18°16.915′N, 70°12.101′W,
27.XI.2004, SCC04_087
Hispaniola: Dominican Republic: Prov. San
Cristóbal, Engombe, farm of autonomous
university, 18°27.360′N, 70°00.306′W,
27.X.2003, SCC03_023
Hispaniola: Dominican Republic: Prov. La
Altagracia, Parque del Este, Guaraguao,
18°19.968′N, 68°48.709′W, 30.XI.2004,
SCC04_090
sel_148, sel_159, sel_215
sel_161, sel_162, sel_163, sel_165,
sel_166, sel_167
sel_586, sel_589
sel_630, sel_631, sel_635, sel_636,
sel_637
sel_018
sel_621, sel_622, sel_633, sel_634,
sel_635, sel_636, sel_637, sel_638
sel_169, sel_170, sel_171, sel_172,
sel_173
sel_562, sel_565, sel_566, sel_568,
sel_585
sel_176, sel_177, sel_178, sel_179
sel_010, sel_011, sel_012, sel_013,
sel_014, sel_015, sel_016, sel_017
sel_200, sel_201, sel_202, sel_203,
sel_204, sel_205, sel_206, sel_207
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
316
S. C. CREWS and R. G. GILLESPIE
APPENDIX Continued
Locality
number
111
112
113
114
115
116
117
118
Species
Collection information
Voucher numbers
Selenops
lindborgi
Selenops
insularis
Selenops
lindborgi
Selenops
insularis
Selenops n. sp.
9
Selenops
insularis
Hispaniola: Dominican Republic: Prov. La
Altagracia, Parque del Este, Boca de Yuma,
18°21.875′N, 68°37.080′W, 29-30.XI.2004,
SCC04_089
Hispaniola: Domnican Republic: Prov. La
Altagracia, Punta Cana Resort,
18°30′55.53″N, 68°22′28.73″W, 5-9.VII.2006,
SCC04_066
sel_191, sel_192, sel_193, sel_194,
sel_195, sel_196, sel_197,
sel_199, sel_216
Hispaniola: Dominican Republic: Prov: Hato
Major, Los Haitises, 1 km south of El Valley
near Hato Mayor, 18°58′24.01″N,
69°22′34.58″W, VI.2006
Hispaniola: Dominican Republic: Prov.
Samaná, Las Terrenas, 19°19′28.88″N,
69°32′50.77″W, VI.2006
sel_555
Hispaniola: Dominican Republic: Prov. La
Vega, Constanza, Alto Cerro Hotel,
18°54′14.16″N, 70°44′41.14″W, VI.2006
Hispaniola: Dominican Republic: Prov. Puerto
Plata, Sosua, VI.2006
Hispaniola: Dominican Republic: Prov. Puerto
Plata, entrance to Loma de Isabel Torres,
19°46′41.9″N, 70°42′01.1″W, 7.X.2006,
SCC06_068
Hispaniola: Dominican Republic, Prov.
Santiago, Mata Grande, 19°11′43.0″N,
70°59′42.0″W, 14–15.X.2006, SCC06_075
Hispaniola: Dominican Republic, Prov.
Santiago, Armando Bermudéz Park, ranger
station at trail to Loma del Oro,
19°12′05.2″N, 71°00′04.8″W, 13.X.2006,
SCC06_074
Hispaniola: Dominican Republic, Prov. Monti
Cristi, Monte Cristi, El Morro,
19°53′42.44″N, 71°39′14.48″W, 8.X.2006,
SCC06_069
sel_546
Selenops
lindborgi
Selenops
insularis
Selenops n. sp. 2
Selenops
insularis
Selenops
insularis
Selenops n. sp.
12
Selenops n. sp.
13
119
Selenops n. sp.
13
120
Selenops
insularis
Selenops n. sp.
10
121
122
Selenops
insularis
Selenops
pensilis
Selenops bani
Selenops phaselus
123
Selenops n. sp. 3
124
Selenops
insularis
Hispaniola: Dominican Republic: Prov.
Independencia, La Descubierta, El Azufrada,
north side of Lago Enriquillo, 18°33.751′N,
71°41.853′W, 26.XI.2004, SCC04_084
Hispaniola: Haiti: Kenskoff, Belot-Montcel,
18°27′11.3″N, 72°21′06.4″W, 20-21.X.2006,
SCC06_076
Puerto Rico: Isla Mona: Bajura de Empalme,
18°06′25.07″N, 67°53′10.41″W
Puerto Rico: Isla Mona: Sardinera,
18°05′46.76″N, 67°56′12.35″W
sel_528, sel_529, sel_530, sel_531,
sel_532, sel_533, sel_534,
sel_535, sel_536
sel_577, sel_590, sel_591, sel_592,
sel_593, sel_594
sel_578
sel_604, sel_605, sel_606, sel_607,
sel_608, sel_609
sel_640, sel_641, sel_642, sel_643,
sel_644, sel_645, sel_646
sel_639
sel_579, sel_580, sel_581, sel_582,
sel_583, sel_584, sel_610,
sel_611, sel_612, sel_613,
sel_614, sel_615, sel_616,
sel_617, sel_618
sel_180, sel_181, sel_182, sel_183,
sel_184, sel_185, sel_186,
sel_187, sel_188, sel_189,
sel_190, sel_569, sel_570,
sel_571, sel_572, sel_599
sel_647, sel_648, sel_649, sel_650,
sel_651, sel_652, sel_653, sel_654
sel_846, sel_847
sel_838, sel_839, sel_840, sel_841,
sel _842, sel_843, sel_844,
sel_845
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
317
APPENDIX Continued
Locality
number
Species
Collection information
Voucher numbers
125
Selenops n. sp. 3
sel_502
126
Selenops n. sp. 3
Selenops
insularis
Selenops
lindborgi
Selenops
lindborgi
Selenops
insularis
Selenops
lindborgi
Selenops
insularis
Selenops
lindborgi
Selenops
insularis
Selenops
lindborgi
Selenops
insularis
Selenops
insularis
Puerto Rico: Isla Mona: Camino de los Cobros,
18°04′02.68″N, 67°52′45.75″W, VII.2006
Puerto Rico: Maricao, Bosque Estatl de
Maricao, 18°08′51.2″N, 66°59′35.0″W,
10.VI.2006, SCC06_045
sel_434, sel_435, sel_436, sel_437,
sel_438, sel_439, sel_440,
sel_441, sel_442
Puerto Rico: Susua State Forest, Sabana
Grande, 18°04′15.0″N, 66°54′31.6″W,
10.VI.2006, SCC06_046
sel_443, sel_444, sel_445, sel_446,
sel_447, sel_448, sel_449,
sel_450, sel_451, sel_452, sel_453
Puerto Rico: Quebradillas, Merendero de
Guajataca, 18°29′23.7″N, 66°56′59.4″W,
9.VI.2006, SCC06_043
sel_423, sel_424, sel_425, sel_426,
sel_427, sel_428, sel_429,
sel_430, sel_431, sel_432
Puerto Rico: Arecibo, Arenalejos, Carretera
657, km 1.9, 18°25′15.9″N, 66°40′35.2″W,
7.VI.2006, SCC06_035
sel_392, sel_393, sel_394, sel_395,
sel_396, sel_397, sel_398
Puerto Rico: between Barceloneta and Arecibo,
Bosque Cambalacheo, 18°27′07.0″N,
66°35′49.9″W, 9.VI.2006, SCC06_041
sel_419, sel_420, sel_421, sel_422
Puerto Rico: Ciales, Bosque Fronton, Carretera
146, km 16.3 Interior Camino Maximo
Nuñez, Sector Los Gonzalez, 18°18′33.8″N,
66°32′42.8″W, 15.VI.2006, SCC06_060
Puerto Rico: Coamo, Baños de Coamo,
18°02′19.4″N, 66°22′27.0″W, 11.VI.2006,
SCC06_048
Puerto Rico: Salina, Reserva Jobos, Parque
Jagueys, 17°57′13.9″N, 66°15′03.5″W,
11.VI.2006, SCC06_049
Puerto Rico: Toa Baja, Bosque Media Luna,
PR-2, km 21–6, 18°24′38.30″N,
66°14′44.36″W, 14.VI.2006, SCC06_058
Puerto Rico: Manuabo, Mariani Creek,
18°00′29.7″N, 65°52′17.0″W, 11.VI.2006,
SCC06_050
sel_503
127
128
129
130
131
132
Selenops
insularis
133
Selenops
lindborgi
134
Selenops
insularus
135
Selenops
lindborgi
Selenops
insularis
Selenops
insularis
136
137
Selenops
lindborgi
138
Selenops
lindborgi
Selenops
insularis
Puerto Rico: Humacao, Barrio Collores,
18°09′44.82″N, 65°49′06.50″W, 11.VI.2006,
SCC06_051
Puerto Rico: Loiza, Punta Vacia, Talega,
18°27′03.8″N, 65°54′16.7″W, 8.VI.2006,
SCC06_038
Puerto Rico: Ceiva, Los Corchos, 18°12′13.8″N,
65°40′06.5″W, 8.VI.2006, SCC06_040
sel_462, sel_463
sel_464
sel_461
sel_465, sel_466, sel_467, sel_468
sel_469, sel_470, sel_471, sel_472,
sel_473, sel_474, sel_475,
sel_476, sel_477, sel_478, sel_479
sel_399, sel_400, sel_401, sel_401,
sel_402, sel_403
sel_416, sel_417, sel_418
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
318
S. C. CREWS and R. G. GILLESPIE
APPENDIX Continued
Locality
number
139
140
Species
Collection information
Voucher numbers
Selenops
lindborgi
Selenops
insularis
Selenops
submaculosus
Puerto Rico: Fajardo, Seven Seas Public Beach,
18°22′03.7″N, 65°38′04.9″W, 8.VI.2006,
SCC06_039
sel_405, sel_406, sel_407, sel_408,
sel_409, sel_410, sel_411,
sel_412, sel_413, sel_414, sel_415
Bahamas: Andros Island: Owens Town,
24°52′30.1″N, 78°02′03.6″W, 13.V.2006,
SCC06_002
Bahamas: Andros Island: Morgan’s Cave at
Morgan’s Bluff, 25°10′30.1″N, 78°01′26.2″W,
13.V.2006, SCC06_003
Bahamas: Andros Island: International Field
Station, 24°53′51.1″N, 77°55′50.1″W,
12.V.2006, SCC06_001
sel_302, sel_303, sel_304, sel_305,
sel_306, sel_307, sel_308
141
Selenops
submaculosus
142
Selenops
submaculosus
143
Selenops
submaculosus
144
145
Selenops
submaculosus
Selenops aissus
146
Selenops aissus
147
Selenops
submaculosus
148
Selenops aissus
149
Selenops aissus
150
Selenops aissus
151
Selenops
lindborgi
152
Selenops
lindborgi
153
Selenops n. sp. 11
Bahamas: Pigeon Cay, near IFS on Andros
Island, 24°52′54.4″N, 77°53′53.5″W,
13.V.2006, SCC06_004
Bahamas: Andros Island: Cargill Creek,
24°30′00.37″N, 77°43′15.61″W, 13.V.2006
Bahamas: Abaco: Ralph’s Chimney off Queen’s
(Abaco) Highway, 26°14′58.2N, 77°11′25.4″W,
14.V.2006, SCC06_006
Bahamas: Abaco: Abaco National Park,
26°03′44.0″N, 77°12′46.2″W, 14.V.2006,
SCC06_005
Bahamas: Great Exuma: Bahamas Sound
Subdivision near old airport, 23°27′56.0″N,
75°46′24.8″W, 18.V.2006, SCC06_011
Bahamas: Great Exuma: Regatta Point,
23°30′24.7″N, 75°45′58.0″W, 18.V.2006,
SCC06_009
Bahamas: Stocking Island, near Great Exuma,
23°32′08.9″N, 75°46′29.6″W, 18.V.2006,
SCC06_010
Bahamas: San Salvador: Gerace Field Station,
trails behind field station, 24°06.9′N,
74°27.8′W, 19.V.2006, SCC06_012
Bahamas: Great Inagua: Man o’War Bay,
21°04′30.2″N, 73°38′36.7″W, 16.V.2006,
SCC06_007
Bahamas: Great Inagua: Old Aerostat Base,
21°06.06.7″N, 73°39′01.9″W, 16.V.2006,
SCC06_228
Turks and Caicos: Providenciales, North-West
Point Pond Nature Reserve, 21°50′32.1″N,
72°19′43.7″W, 8.II.2007, SCC07_010 and
SCC07_011
sel_309, sel_310, sel_311, sel_666
sel_286, sel_287, sel_288, sel_289,
sel_290, sel_291, sel_292,
sel_293, sel_294, sel_295,
sel_296, sel_297, sel_298,
sel_299, sel_300, sel_301,
sel_556, sel_557, sel_558,
sel_595, sel_596, sel_667,
sel_669, sel_671, sel_835, sel_836
sel_312
sel_668
sel_315
sel_313
sel_332
sel_319, sel_320, sel_321, sel_322,
sel_323, sel_324
sel_325, sel_326, sel_327, sel_328,
sel_329, sel_330, sel_331
sel_333, sel_334, sel_335, sel_336,
sel_337, sel_338, sel_339,
sel_340, sel_341, sel_342, sel_343
sel_316
sel_317, sel_318
sel_689, sel_690, sel_691, sel_692
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
319
APPENDIX Continued
Locality
number
Species
Collection information
Voucher numbers
154
Selenops n. sp. 11
sel_346, sel_347, sel_348, sel_349,
sel_694, sel_695
155
Selenops n. sp. 11
156
Selenops n. sp. 11
157
Selenops n. sp. 11
158
Selenops n. sp. 11
159
Selenops debils
gp. species 1
160
Selenops debils
gp. species 1
161
Selenops debils
gp. species 1
162
Selenops debils
gp. species 1
163
Selenops debils
gp. species 1
164
Selenops debils
gp. species 1
Selenops debils
gp. species 1
Turks and Caicos: Providenciales, Turtle Cove,
Third Turtle Drive, 21°47′01.1″N,
72°13′45.4″W, 22.V.2006 and 10.II.2007,
SCC06_017 and SCC07_013
Turks and Caicos: Providenciales, The Bight,
21°47′00.6″N, 72°13′06.4″W, 22.V.2006,
SCC06_015
Turks and Caicos: North Caicos, Wade’s Green
Plantation, 21°55′13.36″N, 72°01′12.45″W,
2–3.II.2007, SCC07_001 and SCC07_005
Turks and Caicos: Middle Caicos, Garden Pond
Field Road, 21°48′24.76″N, 71°45′42.78″W,
3.II.2007, SCC07_004
Turks and Caicos: Providenciales, South View
Drive off of Leeward Highway, 21°46′45.7″N,
72°13′45.4″W, 10.II.2007, SCC07_012
USA: California, San Diego Co., Upper Otay
River Valley, 32°59′50.33″N, 116°19′09.27″W,
19.V.2003
USA: California, San Diego Co., Jamul, Lyons
Valley, north of Lyons Peak, 32°44′03.93″N,
116°53′51.23″W, 29.VII.2007
USA: California, San Diego Co., Anza Borrego
Desert State Park, Carrizo Palm Grove,
32°44′31.29″N, 116°12′51.11″W
Mexico: Baja California Sur, 2 km east of
Ballena between San Ignacio and San
Juanico, 26°27′10.60″N, 111°34′53.39″W
Mexico: Baja California Sur, 3 km west of Villa
Insurgentes on road to San Miguel de
Comundu, 25°16′50.46″N, 111°50′03.75″W
Mexico: Baja California Sur, Cuevas Pintas,
26°01′38.63″N, 111°30′24.45’W
USA: Arizona, Coconino Co., Monument Trail,
flat near archaeological site, 36°25.309′N,
112°27.483′W, 13.VIII.2004, SCC04_013
USA: Arizona: Santa Cruz Co., Madera
Canyon, Mt Wrightson Trail, 31°43′06.86’N,
110°52′22.45’W
Mexico: Sonora, Sonoran Highway on road to
Yecora, 28°23′15.01’N, 108°55′41.42’W
USA: Texas: Val Verde Co., Seminole Canyon,
Highway 90, under bridge, 29°42′21.12’N,
101°18′28.48’W
Mexico: Hidalgo, Villa Flores, Ejido ′El Rayo’,
24°14′1.00’N, 99°1′4.50’W, 2.XI.2007
Mexico: Colima, Manzanillo, Municipio
Manzanillo, 1.2 to 1.4 km east La Central,
19°8′56.4’N, 104°25′35.3’W, X.2005
Mexico: Colima, Municipio Ixtlahuacan,
Tamala, 19°5′2.13’N, 103°47′26.21″
165
166
Selenops debils
gp. species 2
167
Selenops debils
gp. species 3
Selenops debils
gp. species 3
168
169
170
Selenops
mexicanus
Selenops abyssus
171
Selenops abyssus
sel_344, sel_345
sel_675, sel_680, sel_681, sel_682,
sel_683, sel_684, sel_685,
sel_686, sel_687, sel_688, sel_689
sel_676, sel_677, sel_678, sel_679
sel_693
sel_002
sel_837
sel_021, sel_214
sel_210, sel_213
sel_212
sel_009
sel_211
sel_264, sel_270, sel_271, sel_272
sel_851
sel_208
sel_1008
sel_1004
sel_1013
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
320
S. C. CREWS and R. G. GILLESPIE
APPENDIX Continued
Locality
number
Species
Collection information
Voucher numbers
172
Selenops abyssus
sel_1006, sel_1012
173
Selenops n. sp. 4
174
Selenops gracilis
175
Selenops
nigromaculatus?
This is an
immature
specimen, but
is found near
the type
locality of S.
nigromaculatus,
hence the ‘?’
Selenops n. sp. 4
Mexico: Michoacan, Municipio Coalcomán,
Coalcomán, 18°24′12.81″N, 103°07′58.82″W,
X.1005
Mexico: Morelos, Cuernavaca, Colonia
Chamilpa, Instituto de Biotecnología,
UNAM, 18°55′51.86″N, 99°14′16.60″W
Mexico: Guerrero, Arcelia, Campo Morado,
17°34′60.00″N, 100°4′60.00″W
Mexico: Guerrero, Omiltemi, Chilpancingo,
Camino al Omiltemi, 17°33′3.76″N,
99°30′21.83″W
Mexico: Puebla, Municipio Zapotitlan de las
Salinas, Cerro el Pajarito, 18°22′48.3″N,
97°30′26.9″W, V.2005
Mexico: Chiapas, Tuxtla-Gutierrez, Cañon
deSumidero, 16°49′41.46″N, 93°6′22.22″W
Mexico: Chiapas, Berriozabal, dirt road from
Efrain A. Gutierrez, approx. 8 km north
Berriozabal, 16°52′27.45″N, 93°17′28.20″W
Mexico: Chiapas, Pueblo Nueva Solistahuacan,
17°11.550′N, 92°54.875′W, 17.IX.2004,
SCC04_018
Mexico: Chiapas, La Reforma, Municipio La
Concordia, 15°54.212′N, 92°40.157′W,
18.IX.2004, SCC04_018b
Mexico: Chiapas, Huixtla, Las Golindrinas,
15°25.747′N, 92°39.270′W, 23.IX.2004,
SCC04_020
sel_1002, sel_1005
176
177
178
Selenops
mexicanus
Selenops
mexicanus
179
Selenops n. sp.
19
180
Selenops
mexicanus
181
Selenops
mexicanus
Selenops n. sp.
19
Selenops
mexicanus
Selenops n. sp.
19
Selenops
mexicanus
182
183
184
185
186
Selenops
mexicanus
Selenops
mexicanus
Selenops n. sp.
18
Selenops
bifurcatus
Mexico: Chiapas, Municipio Motozintla de
Mendoza, Chevolcan, 15°20′52.4″N,
92°19′25.4″W, 21.XI.2004, SCC04_019
Mexico: Chiapas: road to Roberto Barrio,
~4 km south-west Nuevo Sonora,
17°23′41.10″N, 91°54′10.70″W
Mexico: Veracruz, Municipio Tamalin, El
Mamey, 21°31′13.63″N, 97°38′31.75″W
Guatemala: Petén, Sta Elena de la Cruz,
Colonia del Bosque, near Flores, Cueva
Actun Kan, 16°54′10.9″N, 89°53′44.3″W,
1.I.2008, SCC08_001
Guatemala: Zacatán, Las Guacamayas,
Carretera Sta Rosalia Marmol,
Hídroelectrica Pasabíen, 15°01′39.7″N,
89°41′41.2″W, 1.I.2008, SCC08_002
sel_1010
sel_1014, sel_1015
sel_1007
1016, 1017
sel_848
sel_043
sel_044
sel_031, sel_034, sel_035, sel_036,
sel_037, sel_038, sel_039,
sel_040, sel_041, sel_045,
sel_1011
sel_029, sel_030, sel_032, sel_033,
sel_038
sel_849
sel_1018, sel_1019, sel_1020
sel_865, sel_866, sel_867, sel_868
sel_869, sel_870, sel_871, sel_872,
sel_873, sel_874, sel_875,
sel_876, sel_877, sel_878, sel_879
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS
321
APPENDIX Continued
Locality
number
Species
Collection information
Voucher numbers
187
Selenops
bifurcatus
sel_927
188
Selenops
bifurcatus
189
Selenops
bifurcatus
Selenops
mexicanus
Selenops
bifurcatus
El Salvador: Dep. Chaletenango, Chaletenango
off Carretera Troncal del Norte, Hotel Maya,
14°16′29.1″N, 89°08′32.7″W, 5-6.I.2008,
SCC08_009
El Salvador: Dep. Chaletenango, Mun.
Chaletenango, La Cueva del Corridor,
6.I.2008, SCC08_010
El Salvador: Dep. Chaletenango, Chaletenango,
Municipio Tejutto, Rest. Eucalyptos,
14°12′20.4″N, 89°06′43.9″W, 5.I.2008,
SCC08_008
El Salvador: San Salvador, Museo Nacional de
Historia Natural grounds, 13°40′23.4″N,
89°11′53.6″W, 3.I.2008, SCC08_003
El Salvador: near San Vicente, Mun. Tepetitán
vic. Finca El Carmen, 13°37′53.0″N,
88°50′19.5″W, 4.I.2008, SCC08_005
190
191
Selenops
bifurcatus
Selenops
mexicanus
192
Selenops
bifurcatus
193
Selenops
bifurcatus
194
Selenops
bifurcatus
195
Selenops
bifurcatus
196
Selenops
bifurcatus
Selenops
mexicanus
Selenops
bifurcatus
197
198
Selenops
bifurcatus
199
Selenops
mexicanus
200
Selenops
bifurcatus
Selenops
mexicanus
El Salvador: Dep. San Vicente, vic. San
Vicente, road to Zacatecoluca, behind strip
club ‘Dreamed Girl’, 13°37′43.4″N,
88°46′49.6″W, 3.I.2008, SCC08_004
El Salvador: Dept La Union, Mun. El Carmen,
Lotificacion Amaya, 13°21.44.9″N,
87°59′58.2″W, 5.I.2008, SCC08_007
El Salvador: Dept La Union, Conchagua,
Volcán Conchagua near La Union,
13°18′14.1″N, 87°51′19.6″W, 4.I.2008,
SCC08_006
Nicaragua: Dept Nuevo Segovia, Alc. Ocotal,
Barrio Roberto Gomez above Rio Coco,
13°37′05.8″N, 86°27′57.3″W, 11.I.2008,
SCC08_012
Nicaragua: Dept Madríz, Alc. Ocotal,
Totogalpa, 13°33′49.5″N, 86°29′54.6″W,
11.I.2008, SCC08_013
Nicaragua: Dept Leon, Alc. San Jacinto, Mina
El Límon, Rancho Las Brisas, 12°37′03.8″N,
86°44′34.3″W, 14.I.2008, SCC08_016
Nicaragua: Dept Leon, Alc. El Jicaral, Camino
Santa Rosa, Puente La Guayabita,
12°44′31.2″N, 86°22′44.6″W, SCC08_017,
SCC08_017
Nicaragua: Dept Matagalpa, Alc. San Ramon,
Mata Palo, 12°56′16.5″N, 85°51′12.2″W,
14.I.2008, SCC08_018
Nicaragua: Dept Boaco, Aguas calientes, Alc.
Teustepe, Camino La Cuesta, 12°22′57.8″N,
85°47′30.7″W, 15.I.2008, SCC08_020
sel_925, sel_926, sel_927, sel_928,
sel_929
sel_921, sel_922, sel_923, sel_924
sel_881, sel_882, sel_883, sel_884,
sel_885, sel_886, sel_887,
sel_888, sel_889
sel_897, sel_898, sel_899, sel_900,
sel_901, sel_902, sel_903,
sel_904, sel_905, sel_906,
sel_907, sel_908, sel_909,
sel_910, sel_911
sel_890, sel_891, sel_892, sel_893,
sel_894, sel_895, sel_896
sel_916, sel_917, sel_918, sel_919,
sel_920
sel_912, sel_913, sel_914, sel_915
sel_930
sel_931, sel_932, sel_933, sel_934,
sel_935, sel_936, sel_937,
sel_938, sel_939
sel_954, sel_955, sel_956, sel_957,
sel_958, sel_959, sel_960
sel_961, sel_962, sel_963, sel_964,
sel_965, sel_966, sel_967,
sel_968, sel_969, sel_970
sel_971, sel_972, sel_973, sel_974
sel_963, sel_975, sel_976, sel_977,
sel_978, sel_979, sel_980,
sel_981, sel_982, sel_983,
sel_985, sel_986
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322
322
S. C. CREWS and R. G. GILLESPIE
APPENDIX Continued
Locality
number
Species
Collection information
Voucher numbers
201
Selenops
bifurcatus
Selenops
mexicanus
sel_940, sel_941, sel_942, sel_943,
sel_944, sel_945, sel_946,
sel_947, sel_948, sel_949,
sel_950, sel_951, sel_952, sel_953
202
Selenops
mexicanus
203
Selenops
bifurcatus
Selenops
mexicanus
Selenops
mexicanus
Nicaragua: Lago Nicaragua, Isla Ometepe,
Volcán Concepción, Charco Verde, Hotel
Finca Vincenzia and up hill across the
street, 11°28′42.6″N, 35°38′20.6″W and
11°29′31.2″N, 85°38′14.1″W, 12-13.I.2008,
SCC08_014, SCC08_015
Costa Rica: Guanacaste, Palo Verde National
Park, Cueva Las Tigres, 10°21′58.9″N,
85°21′14.2″W, 17.I.2008, SCC08_022
Costa Rica: Guanacaste: Palo Verde Field
Station, hill behind OTES office,
10°20′42.5″N, 85°20′19.1″W, 17.I.2008,
SCC08_021
Costa Rica: Guanacaste: Nicoya Peninsula,
near Loma Bonita, 10°15′04.0″N,
85°17′30.5″W, 18.I.2008, SCC08_023
Panamá: Barro Colorado Island, I.2008
Panamá: STRI, Galeta, Plot F, 2004
sel_1000, sel_1001
sel_265, sel_266, sel_267, sel_268
Japan: Kyoto, Shugakuin, 23.VIII.2007
sel_861, sel_862, sel_863
Namibia: 12.VI.2006
Tanzania: Iringa, Lutheran House Hostel
South Africa: Guateng, Roodeport, Ruimsig
Butterfly Reserve
South Africa: Tsitsikamma National Park,
78 km E Knysna
South Africa: Grahmstown Municipal. Caravan
Park
South Africa: Eastern Cape, Kai Mouth, 58 km
NE East London
South Africa: Table Mountain National Park,
Newland’s Forest
Madagascar: Reserve Nat. Integrale de Lokobe,
3.61 km ESE Hellville
Madagascar: Park National Montagne d’Ambre
montane rainforest
Australia: Western Australia: Ravensthorpe,
Ravensthorpe Ranges South, WAM 10,
33°38′16.03″S, 120°10′46.01″E, 17.V.2007,
under rock, MCLeng, ML Moir
Australia: Western Australia: Two Peoples Bay
Nature Reserve, granite outcrop, site 6,
34°59′18″S, 118°44″E, 14.X.2006, under
granite rock, ML Moir, JM Waldock
Australia: Western Australia: Fiztgerald River
National Park, East Mt Barren, site 7,
33°55′28″S, 120°01′13″E, 25.XI.2006, under
rock ML Moir, KEC Brennan
sel_998, 999
sel_997
sel_547, sel_548
204
205
206
207
208
209
210
211
212
213
214
215
216
Selenops banksi
Selenops
mexicanus
Selenops
bursarius
Selenops radiatus
Selenops radiatus
Anyphops
barnardi
Anyphops
parvulus
Anyphops
tugelanus
Anyphops
stauntoni
Anyphops kraussi
Garcorops
madagascarensis
Hovops sp.
217
New genus sp. 1
Australia
218
New genus sp. 2
Australia
219
New genus sp. 3
Australia
sel_989, sel_990
sel_987, sel_988
sel_991, sel_992, sel_993, sel_994
sel_549
sel_550
sel_551
sel_552
sel_553
sel_275
T80881, T80996
T78485, T78489
T78500, T78498
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322

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