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

1University of California Berkeley, Department of Environmental Sciences Policy and Management, 137 Mulford Hall, Berkeley, CA 94720-3114, USA 2Berkeley 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 2010bij_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 insights into the complex interaction between colonization and diversification. In particular, the Carib- Remote islands form the basis for many biological bean has served as the setting for the establishment studies because of their ability to act as a laboratory, of most of the central tenets in the equilibrium theory with repeated sets of ecological and/or evolutionary of island biogeography (Munroe, 1948), the argu- experiments occurring within a circumscribed time ments being formulated independently by MacArthur frame (Cronk, 1997; Losos et al., 1998; Gillespie & and Wilson (1963, 1967) much later (Lomolino & Roderick, 2002; Gillespie, 2004; Ricklefs & Berming- Brown, 2009). More recent research on the islands ham, 2008). While the Hawaiian Islands have served has allowed an understanding of the interplay as a model system for processes of in situ diversifica- between ecological and evolutionary processes in tion, the long history of studies on the biota of the shaping species diversity (Losos & Schluter, 2000; Caribbean has provided some of the most important Schoener, Spiller & Losos, 2001). The primary feature of the Caribbean region that makes it particularly useful for examining the inter- *Corresponding author. E-mail: [email protected] action between colonization and diversification is its

288 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 289 long and complex geological history. The Caribbean South America to the Greater Antilles during the Basin began forming nearly 140 Mya. Islands in the Eocene–Oligocene transition 35–33 Mya. The land basin consist of four different types: (1) land-bridge span, although probably short-lived, may have pro- islands which were connected to each other or to the vided an avenue for terrestrial organisms to colonize mainland at times of lower sea level; (2) continental the Greater Antilles from South America. Among islands which broke off from the mainland through mammals, molecular phylogenies of primates and tectonic displacement; (3) uplifted limestone islands; hystricognath rodents are consistent with the model, and (4) volcanic islands (MacPhee & Iturralde-Vinent, while sloths and insectivorans are not (Dávalos, 2005; Robertson, 2009). Despite their limited isola- 2004). The pattern in plants is similarly mixed. tion, the age and geologic complexity of the area have Molecular phylogenetic data from the genera Croton provided ‘well-defined paths of entry by which immi- (Euphorbiaceae) (van Ee et al., 2008) and Styrax (Sty- grants may reach’ the islands (Munroe, 1948). More- racaceae) (Fritsch, 2003) show that the timing of over, the islands have served as the setting for divergence of lineages is consistent with the GAAR- adaptive radiation among lineages with limited dis- landia hypothesis. However, similar data from persal ability, in particular lizards of the genus Anolis endemic legume radiations in the Greater Antilles, (Losos, 1992, 1994, 2009), frogs of the genus Eleuth- although initially thought to indicate ancient splitting erodactylus (Hedges, 1989; Heinicke, Duellman & between lineages consistent with the GAARlandia Hedges, 2007), some lineages of insects [e.g. beetles hypothesis (Lavin et al., 2001), show more recent (Liebherr, 1988b), flies (Wilder & Hollocher, 2003)] diversification (Lavin & Beyra-Matos, 2008), which is and plants [e.g. lineages within the Melastomaceae likely to hold also for lineages of Asteraceae (Michelangeli et al., 2008) and Asteraceae (Francisco- (Francisco-Ortega et al., 2008). Ortega et al., 2008)]. Although studies to date have Clearly, the timing and frequency of dispersal and provided insights into how the individual lineages vicariance, and the interplay between the two, varies have colonized and subsequently diversified within across biotic assemblages. The challenge, then, is to the island system, notable controversies remain, understand the circumstances dictating the relative including the source of colonists and the means by roles of each and how they interact. Arthropods, which they colonized the islands, biogeographic pat- because they can provide a fine-scale resolution of terns within lineages and whether these patterns biogeographic patterns (Ferrier et al, 2004), are ideal might be expected to be shared across multiple lin- candidates for elucidating the nature of these rela- eages (Guyer & Savage, 1986; Williams, 1989; tionships. Although the biogeography of terrestrial Hedges, Hass & Maxon, 1992; Crother & Guyer, 1996; invertebrates in the Caribbean has been examined in Hedges, 1996a,b). some detail (see Liebherr, 1988a and chapters A particular focus of debate has been the role of therein), few recent studies have been attempted, vicariance vs. dispersal in shaping the Caribbean with little molecular information on the timing and biota. Hedges and colleagues (Hedges et al., 1992; nature of the interplay between colonization and Hedges, 1996a,b; Hedges & Heinicke, 2007; Heinicke diversification. However, there are some notable et al., 2007), working with herpetofauna, have sug- exceptions (Davies & Bermingham, 2002; Wilder & gested that the absence of lineages older than the Hollocher, 2003; Brisson, Wilder & Hollocher, 2006). break-up of the proto-Antilles (a contiguous land In particular, recent studies on spiders (Sicariidae: mass between North and South America) precludes a Loxosceles) support the GAARlandia hypothesis in the vicariant origin and they argue for the initial coloni- colonization of the lineage of North from South zation of most taxa via over-water dispersal on America (Binford et al., 2008), while crickets show a flotsam. A similarly dominant role for dispersal has more mixed pattern of both vicariance and dispersal, been suggested for multiple lineages of plants, such coupled with intra-island diversification (Oneal, as Miconieae (Michelangeli et al., 2008). In contrast, 2009). other studies have suggested that vicariance has In this study, we combine molecular and morpho- played a larger role than dispersal in the initial logical methods to examine the phylogenetic relation- colonization of the Caribbean; for example, in lizards ships and biogeographic history of the cursorial and (Crother & Guyer, 1996; Iturralde-Vinent & MacPhee, dispersal-limited spider genus Selenops (Araneae: 1999; MacPhee & Iturralde-Vinent, 2005) and some Selenopidae) in the Caribbean. These primarily tropi- plants [e.g. Euphorbiaceae (van Ee et al., 2008)]. cal and subtropical spiders (Muma, 1953; Corronca, A related controversy focuses on the hypothesis of 1998; Alayón, 2005) are distinctive in that they are GAARlandia (Greater Antilles + Aves Ridge), first extremely dorsoventrally flattened and exceedingly proposed by Iturralde-Vinent & MacPhee (1999), who fast. They are found in a variety of habitats and used geological data and fossil evidence to demon- microhabitats (Crews, Wienskoski & Gillespie, 2008). strate the likely existence of a land span connecting Although the genera and species groups have

Figure 1. Map of the study area. The Americas; the boxed region shows the primary study area. undergone several revisions (Muma, 1953; Corronca, MATERIAL AND METHODS 1998; Alayón, 2005), there is no phylogenetic frame- TAXON SAMPLING work for the family or for any of the component genera. They were chosen for the current study A comprehensive geographic sample of the genus was because of their high diversity and abundance in the obtained from the Caribbean region, including most Caribbean, where they occur in both the Greater and islands and several sites throughout Mexico, Central Lesser Antilles, as well as on the adjacent mainland America and the South American mainland (see also (southern North America and throughout South Supporting Information, Figs S1, S2). Political reasons America) (Muma, 1953; Crews, 2005; Crews et al., prohibited us from obtaining permits to collect several 2008, 2009;) (Figs 1, 2). Accordingly, they provide the endemic species from Cuba and the single species from potential to reveal ﬁne-scale biogeographic patterns Navassa Island. The implications for these omissions across the islands of the Caribbean. The current are discussed at the end of this paper. Outgroups study uses the genus to infer the relative importance included other genera in the family Selenopidae from of the following two processes in dictating the biogeo- all major geographic locations where the family is graphic history of the lineage in the Caribbean: (1) found, in particular the type of the genus (Selenops the frequency of colonization to the Caribbean region radiatus Latreille) from Africa, Selenops bursarius from a mainland source and between islands within Karsch from Japan, Selenops montigenus Simon from the Caribbean; and (2) whether within-island diver- Nepal/India and representatives of the three other siﬁcation has occurred through a single radiation or genera described from Africa (six species of Anyphops, through dispersal and multiple radiations. We also one species of Hovops and one species of Garcorops), test the hypotheses of dispersal and vicariance in the as well as an undescribed Australian genus. Chosen framework of the GAARlandia hypothesis and likeli- representatives outside of the family include a broad hood biogeographic analysis. sample of eight genera from two families, the

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 5′-TTDGNTACCAAGCAGACVGC-3′ – this study 13398 5′-CGCCTGTTTAACAAAAACAT-3′ – Simon et al. 1994 Histone H3aF 5′-ATGGCTCGTACCAAGCAGACVGC-3′ – Colgan et al. 1998 H3aR 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 ampliﬁcation products were ~850, ~800 and ~330 base unpublished data (M. Ramirez, pers. comm.) (Table 4). pairs (bp). DNA sequences can be found on GenBank The genus Selenops has also been found in Domini- (GU109549–GU110746, HM575429–HM576623, and can amber and one of these specimens is an adult HM576658). These markers were chosen as they have male, described by Schawaller (1984) as Selenops become a standard in spider molecular phylogenetics, beynai. The specimen was scanned using X-ray com- with several primers available for each gene (Hedin & puted tomography, as in Penney et al. (2007); Maddison, 2001; Arnedo et al., 2004; Crews & Hedin, however, the poor preservation of the genitalia pro- 2006). Also, the chosen genes evolve at different rates hibited even tentative incorporation into the phyloge- and contain both protein and non-protein coding netic framework. regions. DNA was extracted from a portion of a leg using a Qiagen DNeasy Tissue Kit following the manu- facturer’s protocol. Each new specimen used in this MOLECULAR METHODS study was given an individual number (e.g. sel_001) Four gene fragments were ampliﬁed – three mitochon- and has been deposited in the Essig Museum of drial [cytochrome oxidase I (CO1), 16S ribosomal DNA Entomology at the University of California, Berkeley (16S) and the intervening leucine tRNA and NADH and the California Academy of Sciences. Remaining dehydrogenase I (ND1)] and one nuclear [histone 3a genomic DNA is stored at -80 °C in the Gillespie (H3)] (see Table 1). The respective lengths of the and Roderick Laboratories, University of California,

Berkeley. Primer pairs used are given in Table 1. In Table 2. Partitions used in likelihood and Bayesian some cases, primarily with outgroup taxa, amplifica- analyses and selected models for each partition used in tion was difficult and, in such instances, the Epicentre Bayesian analyses FailSafe PCR kit was used. In the majority of cases, sequence data were obtained for all gene fragments for Partition Selected model multiple representatives of each species. In one situa- tion with the species Selenops insularis Keyserling, 16S stems GTR +G+doublet there was evidence for multiple copies of H3a in some 16S half stems GTR +G specimens, thus these sequences were not analysed for 16S loops GTR + I +G Leucine tRNA stems HKY doublet these individuals. +G+ Leucine tRNA loops HKY +G ND1 postion 1 GTR + I +G PHYLOGENETIC METHODS ND1 position 2 GTR + I +G ND1 position 3 GTR +G Alignments of the protein-coding loci CO1, ND1 and CO1 position 1 GTR + I +G H3a were performed manually using Mesquite ver. CO1 position 2 GTR + I +G 2.5 (Maddison & Maddison, 2008), with the amino- CO1 position 3 GTR +G acid translations used as a guide. The 16S data H3a position 1 GTR + I were aligned using secondary structure based on the H3a position 2 JC + I model from Masta (2000). While there were some H3a position 3 K80 +G length differences between taxa, alignment was straightforward. CO1, cytochrome oxidase I; 16S, ribosomal DNA; ND1, Data were partitioned by codon position for protein NADH dehydrogenase I. coding genes, by stems and loops for ribosomal DNA and by gene for both the maximum likelihood and The RAxML manual suggests two ways to analyse Bayesian analyses to improve the fit of the substitu- data – the ‘fast and easy way’ and the ‘hard and slow tion model to the data (Nylander et al., 2004; Brand- way’ (Stamatakis, 2006). The fast and easy way was ley, Schmitz & Reeder, 2005). The doublet model of used to analyse the full data set because of its large nucleotide substitution was used for the stem regions size (~900 terminals and ~2000 bps). The hard and of 16S and the tRNA (Schöniger & von Haeseler, slow way was used to analyse the smaller data set 1994; Kjer, 2004). Maximum likelihood analyses were and allows the program to find ‘good’ settings particu- performed with RAxML ver. 7.0.4 (Stamatakis, 2006) lar to an individual data set. The user’s manual was and Bayesian analyses were performed using followed exactly for the analysis of the truncated data MrBayes ver. 3.1.2 (Huelsenbeck & Ronquist, 2001; set using the ‘hard and slow’ method. First, five Ronquist & Huelsenbeck, 2003; Altekar et al., 2004). randomized maximum parsimony trees were gener- RAxML is able to analyse partitioned data, but only ated and then each tree was inferred using a fixed under the generalised time reversible (GTR) model, setting of ten for the initial rearrangement. Next, this thus, while the same partitioning regime was used in setting was automatically determined for the same both Bayesian and likelihood analyses, this was the five starting trees and whichever settings yielded the model that was used in the maximum likelihood best likelihood scores were used for subsequent analy- analyses. To determine the models for each partition ses. The second part of the ‘hard and slow’ method in the Bayesian analysis, MrModeltest ver. 2.3 involves the number of rate categories. For this, the (Nylander, 2004) was used. Models were chosen using number of rate categories is increased by 15, from 10 the Akaike information criterion (AIC; Akaike, 1973; to 55 for each of the five starting trees, using which- see Posada & Buckley, 2004) and are listed in Table 2. ever setting worked best from the initial rearrangement analyses. Finally, ten analyses were run using the best settings from the above experiments and LIKELIHOOD ANALYSES bootstraps from 500 iterations were then added to the RAxML maximum likelihood analyses were con- tree with the best likelihood. For all RAxML analyses, ducted in a variety of ways following the suggestions the rapid bootstrap algorithm was used (Stamatakis, of the author (Stamatakis, 2006). First, one analysis Hoover & Rougemong, 2008). was conducted which included 893 terminals after identical haplotypes were removed. However, to ease the computational strain for more intensive analyses, BAYESIAN ANALYSES terminals that were Յ 0.3% different were removed Several analyses were run using MrBayes-mpi on the from the analysis (sensu McGuire et al., 2007). This cluster at the Museum of Vertebrate Zoology, Univer- truncated data set contained 306 terminals. sity 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 293 cluster at the San Diego Supercomputer Center. Table 3. Correspondence between time slices and geo- Despite using the truncated data set and running the graphical ranges defined for models used in Lagrange jobs in parallel, analyses required months to near analyses completion. Programs were run using the default settings for 40 million generations and, if convergence Time slice Land availability was not met, the generations were increased in incre- ments up to 100 million, saving every 1000th tree. 3.0 Closing of the Isthmus of Panamá Convergence was assessed using Are We There Yet? 5.0 Most recent appearance of Northern (AWTY) (Wilgenbusch, Warren & Swofford, 2004; Lesser Antilles 12.0 Most recent appearance of Southern Nylander et al., 2008). Lesser Antilles 33.0 Disappearance of GAARlandia LAGRANGE ANALYSES 35.0 Appearance of GAARlandia 50.0 From 55–50 Mya, a part of Jamaica was The program Lagrange (Ree et al., 2005; Ree & connected to Central America via the Smith, 2008) was used to test hypotheses of vicari- Nicaraguan Rise ance and dispersal. Lagrange uses likelihood models 55.0 Time after which land was available in to test geographic range evolution and allows changes the Greater Antilles region in dispersal and extinction parameters at different 130.0 Age of root node, corresponds to times in the past, allowing the incorporation of exter- separation of Africa and South nal information such as geological data and dispersal America 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 is problematic, in that in some cases this upper bound increase during the time period(s) the land mass was is too old as a result of chance transoceanic inter- available for colonization. In an area as geologically change after actual separation. However, the exist- complex as the Caribbean, there are nearly endless ence of distinct clades of Selenops on the different ways to parameterize the models, but simplicity was continents would argue for little genetic exchange maintained throughout each analysis. between the continental land masses (Smith & Peter- Lagrange requires a tree and a matrix of range son, 2002) and would therefore indicate that it is data for the included taxa. We analysed a truncated indeed appropriate to use the separation of Africa and data set, selecting one specimen from each species, America to date Selenops. Throughout all analyses, along with the outgroups, using a partitioned RAxML we focused on six time periods which correspond to search for the best tree. We then pruned the out- the availability of land for colonization (Table 3). groups before conducting the Lagrange analyses to The following three analyses consisted of two make the computational load smaller, and because models each, one representing each of three scenarios the focus of the questions concerns only the ingroup. with, and without, GAARlandia. The three scenarios We ensured the tree had the same basic structure as were: (1) a dispersal-based scenario where distance trees from the more complete analyses and that all between land masses determines the probability of relationships supported in those analyses also colonization; (2) a dispersal-based scenario in which appeared in this tree. the ability to colonize an available land mass is not We divided the range of the Selenopids in North dependent on distance, thus the colonization of any and Central America into five areas: C (Central one land mass from another is equiprobable; (3) a America and Mexico), S (South America), G (Greater vicariance-based scenario, with little to no over-water Antilles), N (Northern Lesser Antilles), A (Southern dispersal. This means that colonization of one area Lesser Antilles). Although certain parts of these from another could occur only through connections of regions were not available for colonization throughout one land mass to another. In some cases, certain particular time periods (i.e. some of the Greater Anti- areas were never connected to other land masses, lles have been emergent longer than others, etc.), we such as the Lesser Antilles. In this case, the probabil- simply used the maximum times from their first ity of dispersal is not set to zero, but rather a very low appearance. We set the age of the root node of the tree probability, as the presence of the spiders indicates to 130 Myr, as it is assumed a split between the colonization at some point in the past. ingroup, i.e. American selenopids, and the outgroup, It is possible to set different dispersal probabilities i.e. African selenopids, was caused by the separation for each direction, so that the probability of moving of Africa from South America. Dating vicariance from one region to another can be lower or higher events by the initiation of mid-ocean ridge spreading 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 Figure 5 and nodes with posterior probability values equal. Within each model, the only parameter Ն 0.95 are considered to be supported. The branch changed between the two analyses was the probabil- lengths are longer than in the likelihood analyses ity of colonization with and without the presence of and, while a few more basal nodes are supported than GAARlandia. The maximum range size was set to two in the likelihood analyses, the overall pattern is the areas and the areas G, N and A were excluded from same. The similarities and differences among all the root (> 130 Myr) as they were not available for three trees are discussed below. colonization at this time (Table 3). Comparison of trees All three trees are very similar with many of the RESULTS minor differences not supported. The remainder of SAMPLING the basal nodes occurring below the Selenopids of We obtained over 1000 specimens from over 200 North and Central America is only supported as localities within the area of primary focus for this monophyletic in the Bayesian analysis. The focal taxa study. In total, we have 29 out of 41 Caribbean island of the study, the Selenopids of North and Central species, half of the known Mexican and Central America, are monophyletic and further subdivided American species and one fifth of the described South into a well-supported strictly Caribbean clade (Fig. 6, American species (Appendix). 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 PHYLOGENETIC ANALYSES America, the Southern Lesser Antilles (SLA) and Likelihood analysis South America. Within this clade B, although basal The tree from the analysis of the full data set is relationships are not supported, all analyses support shown in Figure 3 and has a likelihood score of a southern Caribbean basin clade (Fig. 6, clade C) -61 544.60. Nodes with bootstrap (BS) values Ն 70% consisting of taxa from Aruba, Bonaire, Curaçao, are considered to be supported. There is no support Trinidad and Tobago. Selenops n. sp. 5 from Aruba is (BS < 70%) for many basal nodes. Further discussion always sister to Selenops curazao from Bonaire and of the results from this tree is given below where Curaçao and, this clade (Fig. 6, clade D), is always compared with trees from the other analyses. sister to Selenops willinki from northern South In the analysis of the truncated data set, the best America and Tobago + S. geraldinae from Trinidad likelihood score came from the trial with a fixed (Fig. 6, clade E). Also within clade B, another well- setting of 10 for the initial rearrangement, rather supported clade in all analyses consists of Selenops than the automatic setting (Table 5). The best likeli- banksi, found in Panama and South America, and hood from the experiment to determine a good setting Selenops micropalpus, found in the Southern Lesser for the number of rate categories occurred when this Antilles from Dominica to St Vincent and the Grena- setting was at 25 (Table 6). Thus, the initial rear- dines (Fig. 6, clade F). rangement setting was fixed at 10 (-i 10) and the There is support for a sister group relationship number of rate categories was set to 25 (-c 25). The between the South American taxa + the Central and best overall likelihood with these settings from North American taxa in the Bayesian tree only. Both the MultTrees analysis came from the second run the Bayesian analysis and the likelihood analysis (Table 7) and the results are shown in Figure 4. The of the truncated data set support a Central overall structure is similar to the tree obtained from American + North American clade (Fig. 6, clade G), as the analysis of the full data set, in which many basal well as one between the widespread Selenops mexica- nodes are not supported, while nodes above these are. nus, Selenops gracilis and a new species found only in This tree is discussed in more detail below. Mexico (Fig. 6, clade H). In the Bayesian tree, S. mexicanus is paraphyletic. There is little support for Bayesian analysis any other relationships in clade B, other than the The analyses were run for 64 million generations (the species from the Selenops debilis group of the south- maximum possible given limits of storage space for western USA and Northern Mexico (Fig. 6, clade I). our output files). According to the cumulative plot The Caribbean clade (Fig. 6, clade A) consists only from AWTY (Wilgenbusch et al., 2004; Nylander et al., of taxa from Caribbean islands and is strongly sup- 2008), the run reached convergence near 55 million ported in all analyses, but, again, with little support generations. Because convergence was only reached for basal nodes, the exception being the widespread very late in the analysis, the first 90% of trees were Selenops lindborgi and its sister species, S. n. sp. 3, eliminated as burn-in, leaving ~12 000 trees from which are supported as sister to the rest of the which to compute a consensus. This tree is shown in Caribbean taxa (Fig. 6, clade J).

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.

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 Vulsor sp. Madagascar, Ranomafana CASENT9024024 – CAS Ctenidae Phoneutria fera French Guiana, Tresor Nature Reserve CASENT9021738 – CAS Ctenidae Cupiennius ca. French Guiana, Emerald Jungle Village CASENT9021735 – CAS granadensis Ctenidae Acanthoctenus sp. Argentina, Parque Nacional Cope, límte NE ARAMR000556 – MACN Sparassidae Olios sp. 1 USA, California, Esparto In author’s personal collection Sparassidae Olios sp. 2 México, Baja California, north of Guerrero In author’s personal collection Negro Sparassidae Polybetes pythagoricus Argentina, Buenos Aires Prov., José Mármol Ar 10876 – MACN Sparassidae Heteropoda sp. Nepal, near Sauraha In author’s personal collection Sparassidae Heteropoda sp. Tanzania In author’s personal collection Sparassidae Damastes sp. 1 Madagascar, Toliara sel_554 – CAS Sparassidae Damastes sp. 2 Madagascar, Ambohitantely CASENT9015896 – CAS

CAS, California Academy of Sciences; MACN, Museo Argentino de Ciencias Naturales.

Table 5. Likelihoods from the ‘hard and slow’ RAxML lihood reconstructions of range evolution under each analyses to determine the best initial rearrangement of the six models. The best likelihood score overall setting for the data (-121.90) was from model 3B, the vicariance-based model that includes GAARlandia (Fig. 9B). In models -ln(L) for initial -ln(L) for automatic 2A–3B, the best likelihood scores were produced from rearrangement setting initial rearrangement those that included GAARlandia. The first two analy- fixed at 10 setting ses (using models 1A and 1B), which take distance between islands into account, produced very similar 49 876.238442* 49 880.146295 likelihood scores and maximum likelihood reconstruc- 49 881.090042 49 877.349169 tions, although the model without GAARlandia had 49 880.995332 49 893.148152 an insignificantly greater likelihood score. However, 49 881.483263 49 879.924317 in the other two analyses, the differences in likelihood 49 879.897717 49 897.163460 scores were significant and, in the analyses modelled *The best score is denoted. with no GAARlandia, there was much more uncer- tainty in the reconstructions (Figs 7–9 – grey A relationship consisting of the four Jamaican branches indicate that alternative reconstructions fall species, four species endemic to Hispaniola + S. insu- within two log-likelihood units of the scenario that is laris, from throughout the Greater Antilles, is repre- depicted). Likelihood ratio tests were used to compare sented in all three analyses, although not supported nested models and, when scenarios were not nested in the full data set (Fig. 6, clade K). The Jamaican (e.g. – scenario 2A and scenario 3A), the highest species are monophyletic, with well-supported inter- likelihood score is taken as the best. relationships in all analyses (Fig. 6, clade L). The sister clade, consisting primarily of Hispaniolan endemics (three of which are undescribed), is also DISCUSSION well supported (Fig. 6, clade M). However, Hispaniola Unique and shared biogeographic patterns are sum- has several species outside this clade. marized in Table 9. Nodes on the branches subtending other major Car- ibbean lineages (clades N, O, P and Q in Fig. 6) are unsupported, although many sister group relation- SOUTHERN CARIBBEAN BASIN ships and one small subclade consisting of three undescribed species from Hispaniola and one from the Members of the well-supported Southern Caribbean Turks and Caicos Islands (Fig. 6, clade N) are sup- Basin clade (Aruba, Bonaire, Curaçao, Trinidad and ported in all analyses. Tobago, Fig. 6, clade C) are never found within the larger well-supported Caribbean clade (Fig. 6, clade Lagrange analyses A). Geological data often suggest a relationship The results of the Lagrange analyses are given in between these southern islands, known as the Aruba– Table 8. Shown in Figures 7–9 are the maximum like- Tobago Belt (Iturralde-Vinent & MacPhee, 1999) and

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 49 876.825172 49 880.146295 49 878.266732 49 881.349421 2 49 909.100516 49 877.349169* 49 877.528777 49 880.668732 3 49 884.108152 49 893.148152 49 882.377547 49 885.827844 4 49 878.378649 49 879.924317 49 884.766631 49 884.610743 5 49 885.284097 49 897.163460 49 877.756447 49 891.633712

*The best score is denoted.

Table 7. Likelihoods from the MultTrees analyses with lizards (Gorman & Atkins, 1969; Jackman et al., the initial rearrangement setting at 10 and the number of 1999; 2002; Creer et al., 2001; Schneider, Losos & de rate categories set to 25 Queiroz, 2001) and Eleutherodactylus frogs (Kaiser, Sharbel & Green, 1994), is that species in the North- Tree -ln(L) ern Lesser Antilles are only distantly related to species in the Southern Lesser Antilles. The species 1 49 890.352743 S. n. sp. 7 is found in the Northern Lesser Antilles 2 49 875.885667* from Les Saintes northward to Montserrat and 3 49 877.963666 Antigua, while the species occurring in the Southern 4 49 876.018067 Lesser Antilles, from Dominica south to St Vincent 5 49 884.591963 and the Grenadines (at least to Mayreau) is S. micro- 6 49 881.261966 palpus. The northern species is nested well within the 7 49 879.547932 strictly Caribbean clade, while S. micropalpus shares 8 49 883.400698 9 49 897.114073 a relationship with S. banksi found from Panamá to 10 49 879.548262 Peru to Guyana. The precise location where northern and southern lineages are separated is variable, being *The best score is denoted. 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), indeed the affinities are not surprising given the carabid beetles (Liebherr, 1988b), butterﬂies (Davies proximity of the islands to each other and to the & Bermingham, 2002), Eleutherodactylus frogs South American continent. The amphibian and reptile (Kaiser et al., 1994) and populations of the banan- assemblages on each of these islands are largely con- aquit (Seutin et al., 1994), it is between St Vincent tinental and also distinct from the primary Caribbean and St Lucia. Differences in the location of the bound- elements (Hedges, 2006). ary between northern and southern lineages may In Selenops, this Southern Caribbean clade is occur as a result of the timing of colonization of the apparently not closely related to other Caribbean different groups, which is likely related to the timing taxa, a pattern found in many other groups, including of emergence of the individual islands. Interestingly, mammals (Dávalos, 2004) and plants [orchids (Trejo- anoles from the Southern Lesser Antilles, like the Torres & Ackerman, 2001)]. However, a contrasting spiders, show affinities with Central and South pattern has been found in Anolis lizards in which the American anoles (Jackman et al., 1999; Creer et al., Southern Caribbean Basin taxa show stronger affini- 2001). ties 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 ORIGIN OF TAXA further west, and thus closer to Bonaire, in the past (Creer et al., 2001). 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 NORTHERN VS.SOUTHERN LESSER ANTILLES Islands, St Kitts, Nevis, eastern Hispaniola and Great A pattern that the Selenops spiders share with Inagua in the Bahamas, see also Supporting Informa- several insects (Wilder & Hollocher, 2003), Anolis tion, Fig. S1E–G) and the very narrowly distributed

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 bank (Malone et al., 2000). This pattern, which indi- Information, Fig. S1F) S. n. sp. 3 (Fig. 6 clade J). A cates a common origin of Caribbean diversity for similar pattern is found among Anolis, with Puerto these groups, is in contrast to data from geckos, frogs, Rico endemic Anolis occultus also basal (Jackman colubrid snakes and butterﬂies, which suggest His- et al., 1999). Likewise, the most basal iguana of the paniola as a centre of diversity (Liebherr, 1988a, and genus Cyclura is also located on the Puerto Rican references therein).

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 Jamaican species of Selenops form a monophyletic group of endemics (Fig. 6, clade L). Monophyly of Jamaica is one of the oldest islands of the Greater Jamaican taxa is also present in anoles (Jackman Antilles, with areas that may have had some parts et al., 1999; Nicholson et al., 2005) and Eleutherodac- continuously above sea level for many millions of tylus frogs (Hedges, 1996a,b). However, affinities of years longer than other islands (Iturralde-Vinent & the Jamaican clade differ between spider and verte- MacPhee, 1999; Iturralde-Vinent & Gahagan, 2002). brate groups: The Jamaican clade of Selenops is sup- It is also more isolated than other islands as its last ported in the Bayesian and truncated likelihood probable connection with a land mass was likely with analyses as being sister to a clade of primarily His- Central America through the Nicaraguan Rise paniolan species (Fig. 6, clade K). In contrast, the 55 Mya. Our data reﬂect this isolated history, as Jamaican clade of Eleutherodactylus frogs is most

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 BIOGEOGRAPHICALLY DERIVED POSITION OF CUBA Jamaican lineages of Anolis lizards (Nicholson et al., 2005) and short-faced bats (Dávalos, 2007) are sister Cuba has often been depicted as a basal locality in to clades from the mainland. Overall, Jamaica’s area cladograms (Buskirk, 1985; Crother & Guyer, history has been quite different from that of the other 1996). In contrast, although not always supported, Greater Antillean islands and its fauna may have Selenops species from Cuba appear not to be basal, at accumulated via dispersal and in situ speciation least based on morphology and our limited molecular rather than vicariance (Buskirk, 1985; Crother & sampling. Only one species (Selenops aissus – col- Guyer, 1996). lected from the Bahamas, but that also occurs in

Table 8. Results of the Lagrange analyses for each of the six proposed models

Model -ln(L) Dispersal Extinction

1A – dispersal where distance is important; without GAARlandia 135.5 0.01634 0.0082 1B – dispersal where distance is important; with GAARlandia 135.9 0.01637 0.008387 2A – dispersal where distance is not important; without GAARlandia 144.3 0.02604 0.008411 2B – dispersal where distance is not important; with GAARlandia 128.1* 0.02137 0.008094 3A – little to no over-water dispersal; without GAARlandia 260.8 0.4647 0.006629 3B – little to no over-water dispersal; with GAARlandia 121.9* 0.1189 0.008262

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 GAAR- landia.

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 and St Maarten and Anguilla are nested well within other sampled species which occur in Cuba (although a clade of Hispaniolan animals (Fig. 6, clade O). This all but one – Selenops submaculosus – were collected suggests that S. n. sp. 7 and S. n. sp. 8 colonized the from other islands) are nested high within the trees Northern Lesser Antilles region from Hispaniola and (S. submaculosus, Selenops simius, Selenops inus- thus support the Greater Antilles as a centre of laris) and it is inferred based on morphology (S. C. species diversity via dispersal events in Anolis (Glor, Crews & R. G. Gillespie, unpubl. data) that most of Losos & Larson, 2005). Also, many species of Selenops the Cuban endemics are closely related to S. simius in Hispaniola have very small ranges that mirror and S. submaculosus (Fig. 6, clade O). those of many endemic anoles from the Anolis cybotes group (Glor et al., 2003), indicating similar patterns of speciation between the two groups. GREATER ANTILLES AS A CENTRE OF SPECIES DIVERSITY There are two additional patterns that appear to be SPECIES–AREA RELATIONSHIPS shared between Selenops and Anolis.InSelenops In many taxa as diverse as fungi (Lodge, Baroni & spiders, the species from the Northern Lesser Antilles Cantrell, 2002), vertebrates (Ricklefs & Lovette,

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 GAAR- landia.

1999) and invertebrates (Nichols, 1988), the Greater endemics. However, the Bahamas have no known Antilles harbour more species than the Lesser Anti- endemic species of Selenops. lles. This can be attributed largely to island area (MacArthur & Wilson, 1963) and associated habitat diversity and age (Losos, 1996; Ricklefs & Berming- HYPOTHESIS TESTING ham, 2002, 2008). In Selenops, the same pattern is In the maximum likelihood analyses of range expan- found, with larger, older islands (Greater Antilles) sion, likelihood ratio tests of scores for the scenarios having more species than smaller, younger, less that include the existence of the GAARlandia land habitat-diverse islands (Lesser Antilles). In the span are either equally probable or more favourable Greater Antilles there is often a pattern of number than those that do not. This does not mean that of species in Cuba > Hispaniola > Jamaica > Puerto over-water dispersal has not occurred, but rather that Rico, based on island size. This pattern also prevails land bridges hold a stronger signature on the phylog- in Selenops. In this genus, 17 species occur in Cuba eny. These results contrast to those for mammals in with 12 endemics (Alayón, 2005), while in Hispaniola which there was little to no support for a land span there are at least 16 species with 11 endemics and, in between the Greater Antilles and northern South Jamaica, at least ﬁve species are known, with four America (Dávalos, 2004). Likewise, Hedges and

Table 9. Biogeographic patterns in Caribbean Selenops species

Shared with Clades that show Pattern other taxa? Which taxa? particular patterns

1. Distinct, distantly related SLA and Yes Lygaeid bugs, Carabid beetles, F and O NLA clades fruitflies, butterflies, Anolis, Eleutherodactylus, bananaquit 2. Monophyly of Jamaican taxa Yes Anolis, Eleutherodactylus L 3. More than one colonization and Yes Anolis A, J, M, N diversification in Hispaniola 4. Patterns of endemism throughout the Yes Fungi, Anolis, birds, carabid LA–F,O Caribbean (GA harbour more beetles GA – J,K,L,M,N,P,Q endemics than LA) 5. Endemic species in Hispaniola with Yes Anolis N, P, others in clade A distributions that overlap endemics of other taxa 6. Southern Netherlands Antilles form a No – C clade with Trinidad and Tobago exlcusive of other Caribbean taxa 7. Location of the split between the NLA No – SLA – F and SLA is between Dominica and NLA–O Les Saintes, Guadeloupe 8. Cuban species are not basal in the No – Q, S. aissus Caribbean clade

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, basal relationships, it should also be noted that deep, 1996a,b; Hedges & Heinicke, 2007; Heinicke et al., short branches, such as those found here, may be very 2007) found that molecular clock estimates of diver- difficult if not impossible to resolve (Degnan & Salter, gence times precluded a major role of land bridges in 2005; Kubatko & Degnan, 2007). The results reveal the origin of Caribbean herpetofauna. The suitability several patterns common to other disparate taxa, as Dispersal–Extinction–Cladogenesis model of geo- well as many unique patterns which warrant further graphic range evolution used here, in which dispersal study. Moreover, the data set provides the ground- events cause range expansion, local extinction events work for behavioural, ecological and population-level cause range contraction and the probability of each studies similar to lineages such as Anolis lizards kind of event is proportional to the branch length, has (Losos, 2009) and passerine birds (Ricklefs & Ber- been questioned for island fauna, as terminal taxa mingham, 2007). may be restricted to single islands (Ree & Smith, 2008). However, in our models, islands were either grouped together or several species were spread ACKNOWLEDGEMENTS across multiple islands and thus the model is reason- We would like to acknowledge members of S.C.C.’s able in this particular case. dissertation committee for their guidance: George Roderick, Jim McGuire and Charles Griswold. We would like to thank the following museums, curators CONCLUSIONS and collection managers for specimen loans: American The current study provides a basis for biogeographic Museum of Natural History – Norman I. Platnick and comparison across different lineages in the Carib- Louis Sorkin; Museum of Comparative Zoology – bean. It is one of the most extensive data sets for Laura Leibensperger; California Academy of Sciences Caribbean fauna and the most comprehensive – Charles Griswold; National Museum of Natural molecular data set of any spider group. While inclu- History – Jonathan Coddington; British Museum of sion of taxa currently missing from our analyses, and Natural History – Janet Beccaloni; Peabody Musem possibly the use of other markers, may help resolve 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 305 mology – Cheryl Barr; Museo Nacional de Histora Binford GJ, Callahan MS, Bodner MR, Rynerson MR, Natural, Santo Domingo – Sardis Medrano Cabral, Berea Nuñez P, Ellison CE, Duncan RP. 2008. Phylo- Carmelo Nuñez. We would also like to thank Jim genetic relationships of Loxosceles and Sicarius are consis- McGuire for use of the MVZ cluster and Mark Miller tent with Western Gondwanan vicariance. Molecular and Lucie Chan for use of the SGE cluster and the Phylogenetics and Evolution 49: 538–553. CIPRES portal and the San Diego Supercomputer Brandley MC, Schmitz A, Reeder TW. 2005. Partitioned Center. We are extremely grateful to Richard Ree for Bayesian analyses, partition choice, and the phylogenetic quickly responding and helping us to understand and relationships of scincid lizards. Systematic Biology 54: 373– 390. implement Lagrange. We would also like to thank Brisson JA, Wilder J, Hollocher H. 2006. Phylogenetic Matt Brandley for phylogenetic methodology discus- analysis of the cardini group of Drosophila with respect to sions. We are grateful to all of the many people that changes in pigmentation. Evolution 60: 1228–1241. aided us in obtaining permits and collecting: Kelvin Buskirk RE. 1985. Zoogeographic patterns and tectonic Guerrero, Denia Veloz, Eladio Fernandez, Alberto history of Jamaica and the Northern Caribbean. Journal of Puente-Rolón, Beverly Mae Nisbeth, Adriel Thibou, Biogeography 12: 445–461. Germain George, Brian Riggs, Brian Manco, Marga- Colgan DJ, McLauchlan A, Wilson GDF, Livingston SP, ret Jones, Renata Platenberg, Chris Niebuhr, Abel Edgecombe GD, Macaranas J, Cassis G, Gray MR. Pérez-González, G. B. Edwards, Oscar Francke, Ale- 1998. Histone H3 and U2 snRNA DNA sequences and jandro Mondragon, Mark da Silva, Facundo Franken, arthropod molecular evolution. Australian Journal of Roy Croes, Gijs Van Hoorn, Adolphe O. Debrot, Mark Zoology 46: 419–437. Vermeij, Fred the Abaco Caveman, Raveen Gibson, Corronca JA. 1998. A taxonomic revision of the afrotropical Daniel Palmer, Jim Starrett, Marshal Hedin, Nicole species of Selenops Latreille, 1819 (Araneae, Selenopidae). VanderSal, Sean Schoville, Luke Mahler, Uri García, Zootaxa 107: 1–35. Beto Mendoza, Adrian Nieto Montes de Oca, Rebecca Creer DA, de Queiroz K, Jackman TR, Losos JB, Larson Duncan, Pierre Paquin, Matthew Cottam, Jan den A. 2001. Systematics of the Anolis roquet series of the Dulk, Joey Slowik, Nicole Esteban, Arturo Herrera, Southern Lesser Antilles. Journal of Herpetology. 35: 428– Nancy Bottomley, Inilek Wilmot, Lauren Esposito, 441. Stephen Touissant, Arlington James, Ferdinand Crews SC. 2005. Selenopidae. In: Ubick D, Paquin P, Tripoli, Daniel Memia Zolo, Nouree-Yvon, Martín Cushing PE, Roth V, eds. Spiders of North America: an Ramírez, Mark Harvey, Volker Framenau, Jeremy identiﬁcation manual. Keene, NH: American Arachnological Miller, Hannah Wood, Yuri Marusik, Caroline Society, 221. Chaboo, Cheryl Barr, Bill Shepard, Akio Tanikawa, C. Crews SC, Hedin MC. 2006. Studies of morphological and molecular phylogenetic divergence in spiders (Araneae: J. Hayden, Aaron Abdel, Dan Warren. Finally, we Homalonychus) from the American southwest, including would like to acknowledge Matjaz Kunter and an divergence along the Baja California Peninsula. Molecular anonymous reviewer for their straightforward and Phylogenetics and Evolution 38: 470–487. constructive reviews of this paper. Funding was pro- Crews SC, Puente-Rolón AR, Rutstein E, Gillespie RG. vided by the Schlinger Foundation, with additional 2009. A comparison of populations of island and adjacent support from the Margaret C. Walker Fund. mainland species of Caribbean Selenops (Araneae: Selenopi- dae) spiders. Molecular Phylogenetics and Evolution 54: 970–983. REFERENCES Crews SC, Wienskoski E, Gillespie RG. 2008. Life history of the spider Selenops occultus Mello-Leitão (Araeae, Sele- Akaike H. 1973. Information theory and an extension of the nopidae) from Brazil with notes on the natural history of the maximum likelihood principle. In: Petrov BN, Caski F, eds. genus. Journal of Natural History 42-43: 2747–2761. Second international symposium on information theory. Cronk QCB. 1997. Islands: stability, diversity, conservation. Budapest, Hungary: Akademiai Kiado, 267–281. Biodiversity and Conservation 6: 447–493. Alayón G. 2005. La familia Selenopidae (Arachnidae: Crother BI, Guyer C. 1996. Caribbean historical biogeogra- Araneae) en Cuba. Solenodon 5: 10–52. phy: was the dispersal vicariance debate eliminated by an Altekar G, Dwarkadas S, Huelsenbeck JP, Ronquist F. extraterrestrial bolide? Herpetologica 52: 440–465. 2004. Parallel Metropolis coupled Markov chain Monte Dávalos LM. 2004. Phylogeny and biogeography of Carib- Carlo for Bayesian phylogenetic inference. Bioinformatics bean mammals. Biological Journal of the Linnean Society 20: 407–415. 81: 373–394. Arnedo MA, Coddington JA, Agnarsson I, Gillespie RG. Dávalos LM. 2007. Short-faced bats (Phyllostomidae: Steno- 2004. From a comb to a tree: phylogenetic relationships of dermatina): a Caribbean radiation of strict frugivores. the comb-footed spiders (Araneae, Theridiidae) inferred Journal of Biogeography 34: 364–375. from nuclear and mitochondrial genes. Molecular Phyloge- Davies N, Bermingham E. 2002. The historical biogeogra- netics and Evolution 31: 225–245. phy of two Caribbean butterﬂies (Lepidoptera: Heliconiidae)

as inferred from genetic variation at multiple loci. Evolution Academy of Sciences of the United States of America 89: 56: 573–589. 1909–1913. Degnan JH, Salter LA. 2005. Gene tree distributions under Hedges SB, Heinicke MP. 2007. Molecular phylogeny and the coalescent process. Evolution 59: 24–37. biogeography of West Indian frogs of the genus Leptodacty- Ferrier S, Powell GVN, Richardson KS, Manion G, lus. Molecular Phylogenetics and Evolution 44: 308–314. Overton JM, Allnutt TF, Cameron SE, Mantle K, Hedin MC, Maddison WP. 2001. A combined molecular Burgess ND, Faith DP, Lamoreux JF, Kier G, Hijmans approach to phylogeny of the jumping spider subfamily RJ, Funk VA, Cassis GA, Fisher BL, Flemons P, Lees Dendryphantinae (Araneae: Salticidae. Molecular Phyloge- D, Lovett JC, Van Rompaey RSAR. 2004. Mapping more netics and Evolution 18: 386–403. of terrestrial biodiversity for global conservation assess- Heinicke MP, Duellman WE, Hedges SB. 2007. Major ment. BioScience 54: 1101–1109. Caribbean and Central American frog faunas originated by Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R. 1994. ancient oceanic dispersal. Proceedings of the National DNA primers for amplification of mitochondrial cytochrome Academy of Sciences of the United Sates of America 104: C oxidase subunit I from diverse metazoan invertebrates. 10092–10097. Molecular Marine Biology and Biotechnology 3: 294–299. Huelsenbeck JP, Ronquist FR. 2001. MrBayes: Bayesian Francisco-Ortega J, Ventosa I, Oviedo R, Jiménez F, inference of phylogeny. Bioinformatics 17: 754–755. Herrera P, Maunder M, Panero JL. 2008. Caribbean Iturralde-Vinent MA, Gahagan L. 2002. Late Eocene to island Asteraceae: systematics, molecules, and conservation Middle Miocene tectonic evolution of theCaribbean: Some on a biodiversity hotspot. The Botanical Review 74: 112–131. principles and their implications for plate tectonic modeling. Fritsch PW. 2003. Multiple geographic origins of Antillean In: Jackson TA, ed. Caribbean geology into the third mil- Styrax. Systematic Botany 28: 421–430. lennium: transactions of the Fifteenth Caribbean Geological Gillespie RG. 2004. Community assembly through adaptive Conference. Mona, Jamaica: University of the West Indies radiation in Hawaiian spiders. Science 16: 356–359. Press, 47–62. Gillespie RG, Roderick GK. 2002. Arthropods on islands: Iturralde-Vinent MA, MacPhee RDE. 1999. Paleogeogra- colonization, speciation, and conservation. Annual Review of phy of the Caribbean region: implications for Cenzoic bio- Entomology 47: 595–632. geography. Bulletin of the American Museum of Natural Glor RE, Kolbe JK, Powell R, Larson A, Losos JB. 2003. History 238: 1–95. Phylogenetic analysis of ecological and morphological diver- Jackman TR, Irschick DI, de Queiroz K, Losos JB, sification in Hispaniolan trunk-ground anoles (Anolis Larson A. 2002. Molecular phylogenetic perspective on cybotes group). Evolution 58: 2383–2397. evolution of lizards of the Anolis grahami series. Journal of Glor RE, Losos JB, Larson A. 2005. Out of Cuba: overwater Experimental Zoology Part B 294: 1–16. dispersal and speciation among lizards in the Anolis caroli- Jackman TR, Larson A, de Queiroz K, Losos JB. 1999. inensis subgroup. Moleular Ecology 14: 2419–2432. Phylogenetic relationships and tempo of early diversifica- Gorman GC, Atkins L. 1969. The zoogeography of Lesser tion in Anolis lizards. Systematic Biology 48: 254–285. Antillean Anolis lizards – an analysis based upon chromo- Kaiser H, Sharbel TF, Green DM. 1994. Systematics and somes and lactic dehydrogenases. Bulletin of the Museum of biogeography of eastern Caribbean Eleutherodactylus Comparative Zoology 138: 554–579. (Anura: Leptodactylidae): evidence from allozymes. Guyer C, Savage JM. 1986. Cladistic relationships among Amphibia. Reptilia 15: 375–394. anoles (Sauria:Iguanidae). Systematic Zoology 35: 509–531. Kjer KM. 2004. Aligned 18S and insect phylogeny. Systematic Hedges SB. 1989. Evolution and biogeography of West Indian Biology 53: 506–514. frogs of the genus Eleutherodactylus: slow-evolving loci and Kubatko LS, Degnan JH. 2007. Inconsistency of phyloge- the major groups. In: Woods CA, ed. Biogeography of the netic estimates from concatenated data under coalescence. West Indies. Past, present and future. Gainesville, FL: San- Systematic Biology 56: 17–24. dhill Crane Press, 305–370. Lavin M, Beyra-Matos A. 2008. The impact of ecology and Hedges SB. 1996a. The origin of West Indian amphibians biogeography on legume diversity and phylogeny in the and reptiles. In: Powell R, Henderson RW, eds. Contribu- Caribbean region: a new direction in historical biogeogra- tions to West Indian herpetology: a tribute to Albert phy. The Botanical Review 74: 178–196. Schwartz. Ithaca, NY: Society For The Study of Amphibians Lavin M, Wojciechowski MF, Richman A, Sanderson M. and Reptiles, 95–128. 2001. Identifying Tertiary radiations of Fabaceae in Hedges SB. 1996b. Historical biogeography of West Indian the Greater Antilles: alternatives to cladistic vicariance vertebrates. Annual Review of Ecology and Systematics 27: analysis. International Journal of Plant Science 162: S53– 163–196. S76. Hedges SB. 2006. Paleogeography of the Antilles and the Liebherr JK. 1988a. Zoogeography of Caribbean insects. origin of West Indian amphibians and reptiles. Applied Ithaca, NY: Cornell University Press. 285p. Herpetology 3: 281–292. Liebherr JK. 1988b. Biogeographic patterns of West Indian Hedges SB, Hass CA, Maxon LR. 1992. Caribbean bioge- Platynus carabid beetles (Coleoptera). In: Liebherr JK, ed. ography: molecular evidence for dispersal in West Indian Zoogeography of Caribbean insects. Ithaca, NY: Cornell Uni- terrestrial vertebrates. Proceedings of the National versity Press, 121–152.

Lodge DJ, Baroni TJ, Cantrell SA. 2002. Basidiomycetes of Michelangeli FA, Judd WS, Penneys DS, Skean Jr. JD, the Greater Antilles Project. In: Watling R, Frankland JC, Bécquer-Granados ER, Goldenberg R, Martin CV. Ainsworth AM, Isaac S, Robinson CH, eds. Tropical mycol- 2008. Multiple events of dispersal and radiation of the tribe ogy V. 1 macromycetes. Engham UK: CABI Publishing, Miconieae (Melastomataceae) in the Caribbean. Botanical 45–60. Review 74: 53–77. Lomolino MV, Brown JH. 2009. The reticulating phylogeny Muma MH. 1953. A study of the spider family Selenopidae in of island biogeography theory. Quarterly Review of Biology North and Central America and the West Indies. American 84: 357–390. Museum Novitates 1619: 1–55. Losos JB. 1992. The evolution of convergent community Munroe EG. 1948. The geographical distribution of butter- structure in Caribbean Anolis communities. Systematic flies in the West Indies. PhD Dissertation. Ithaca, NY: Biology. 41: 403–420. Cornell University. Losos JB. 1994. Historical contingency and lizard community Nichols SW. 1988. Kaleidoscopic biogeography of West ecology. In: Vitt LJ, Pianka ER, eds. Lizard ecology: histori- Indian Scaritinae (Coleoptera: Carabidae). In: Liebherr JK, cal and experimental perspectives. Princeton, NJ: Princeton ed. Zoogeography of Caribbean insects. Ithaca, NY: Cornell University Press, 319–333. University Press, 71–121. Losos JB. 1996. Ecological and evolutionary determinants of Nicholson KE, Glor RE, Kolbe JJ, Larson A, Hedges SB, the species-area relation in Caribbean anoline lizards. Philo- Losos JB. 2005. Mainland colonization by island lizards. sophical Transactions of the Royal Society B 351: 847–854. Journal of Biogeography 32: 929–938. Losos JB. 2009. Lizards in an evolutionary tree: ecology and Nylander JAA. 2004. MrModeltest v2. Program distributed adaptive radiation of anoles. Berkeley, CA: University of by the author. Evolutionary Biology Centre, Uppsala Uni- California Press. versity. Losos JB, Jackman TR, Larson A, de Queiroz K, Nylander JAA, Ronquist F, Huelsenbeck JP, Nieves- Rodríguez-Schettino L. 1998. Historical contingency and Aldrey JL. 2004. Bayesian phylogenetic analysis of com- determinism in replicated adaptive radiations of island bined data. Systematic Biology 53: 47–67. lizards. Science 279: 2115–2118. Nylander JAA, Wilgenbusch JC, Warren DL, Swofford Losos JB, Schluter D. 2000. Analysis of an evolutionary DL. 2008. AWTY (are we there yet?): a system for graphical species-area relationship. Nature 408: 847–850. exploration of MCMC convergence in Bayesian phylogenet- Losos JB, Thorpe RS. 2004. Evolutionary diversification of ics. Bioinformatics 24: 581–583. Anolis lizards: introduction. In: Dieckmann U, Metz HAJ, Oneal E. 2009. Biogeographic and evolutionary mechanisms Doebeli M, Tautz D, eds. Adaptive speciation. Cambridge: driving diversification in Caribbean ground crickets (genus Cambridge University Press, 343–344. Amphiacusta). PhD dissertation. Ann Arbor, MI: University MacArthur RH, Wilson EO. 1963. An equilibrium theory of of Michigan. insular zoogeography. Evolution 17: 373–387. Penney D, Dierick M, Cnudde V, Masschaele B, Vlas- MacArthur RH, Wilson EO. 1967. The theory of island senbroeck J, Van Hoorebeke L, Jacobs P. 2007. The biogeography. Princeton, NJ: Princeton University Press, first Micropholcommatidae (Araneae), imaged in Eocene 224p. Paris amber using X-ray computed tomography. Zootaxa MacPhee RDE, Iturralde-Vinent MA. 2005. The interpre- 1623: 47–53. tation of Caribbean paleogeography: reply to Hedges. In: Posada D, Buckley TR. 2004. Advantages of AIC and Baye- Alcover JA, Bover P, eds. Proceedings of the International sian approaches over likelihood ratio tests for model selec- Symposium ‘Insular Vertebrate Evolution: The Palaeonto- tion in phylogenetics. Systematic Biology 53: 793–808. logical Approach’. Monografies de la Societat d’Història Ree RH, Moore BR, Webb CO, Donoghue MJ. 2005. A Natural de les Balears 12. likelihood framework for inferring the evolution of geo- Maddison WP, Maddison DR. 2008. Mesquite: a modular graphic range on phylogenetic trees. Evolution 59: 2229– system for evolutionary analysis. Version 2.5. Available at 2311. http://mesquiteproject.org Ree RH, Smith SA. 2008. Maximum-likelihood inference of Malone CL, Wheeler TC, Davis SK, Taylor JF. 2000. geographic range evolution by dispersal, local extinction, Biogeography and systematic of the Caribbean rock iguana and cladogenesis. Systematic Biology 57: 4–414. (Cyclura): implications for conservation and insights into Ricklefs RE, Bermingham E. 2002. The concept of the the biogeographic history of the West Indies. Molecular taxon cycle in biogeography. Global Ecology and Biogeogra- Phylogenetics and Evolution 17: 269–279. phy 11: 353–361. Masta SE. 2000. Mitochondrial sequence evolution in spiders: Ricklefs RE, Bermingham E. 2007. Evolutionary radiations intraspecific variation in tRNAs lacking the TYC arm. of passerine birds in archipelagoes. The American Natural- Molecular Biology and Evolution 17: 1091–1100. ist 169: 285–297. McGuire JA, Linkem CW, Koo MS, Hutcison DW, Lappin Ricklefs RE, Bermingham E. 2008. The West Indies as a AK, Orange DI, Lemos-Espinal J, Riddle BR, Jaeger laboratory of biogeography and evolution. Philosophical JR. 2007. Mitochondrial introgression and incomplete Transactions of the Royal Society B 363: 2393–2413. lineage sorting through space and time: phylogenetics of Ricklefs RE, Lovette IJ. 1999. The roles of island area per crotophytid lizards. Evolution 61: 2879–2897. se and habitat diversity in the species–area relationships of

four Lesser Antillean faunal groups. Journal of Animal Caribbean insects. Ithaca, NY: Cornell University Press, Ecology 68: 1142–1160. 38–60. Robertson REA. 2009. Antilles geology. In: Gillespie RG, Smith AB, Peterson KJ. 2002. Dating the time of origin of Clague DA, eds. Encyclopedia of islands. Berkeley, CA: major clades: molecular clocks and the fossil record. Annual University of California Press, 29–35. Review of Earth and Planetary Sciences 30: 65–88. Ronquist F, Huelsenbeck JP. 2003. MRBAYES 3: Bayesian Stamatakis A. 2006. RAxML-VI-HPC: maximum likelihood- phylogenetic inference under mixed models. Bioinformatics based phylogenetic analyses with thousands of taxa 19: 1572–1574. and mixed models. Bioinformatics 22: 2688–2690. Schawaller W. 1984. Die families Selenopidae in Dominika- Stamatakis A, Hoover P, Rougemong J. 2008. A rapid nischem Berrnstein (Arachnida, Araneae). Stuttgarter bootstrap algorithm for the RAxML web servers. Systematic Beiträge zur Naturkunde Serie B (Geologie und Paläontolo- Biology 57: 758–771. gie) 103: 1–8. Trejo-Torres JC, Ackerman D. 2001. Biogeography of the Schneider CJ, Losos JB, de Queiroz K. 2001. Evolutionary Antilles based on a parsimony analysis of orchid distribu- relationships of the Anolis bimaculatus group from the tions. Journal of Biogeography 28: 775–794. Northern Lesser Antilles. Journal of Herpetology 35: 1–12. van Ee B, Berry PE, Riina R, Gutiérrez Amaro JE. 2008. Schoener TW, Spiller DA, Losos JB. 2001. Natural resto- Molecular phylogenetics and biogeography of the ration of the species–area relation for a lizard after a Caribbean-centered Croton subgenus Moacroton (Euphorbi- hurricane. Science 294: 1525–1528. aceae s.s.). Botanical Review 74: 132–165. Schöniger M, von Haeseler A. 1994. A stochastic model and Wilder JA, Hollocher H. 2003. Recent radiation of endemic the evolution of autocorrelated DNA sequences. Molecular Caribbean Drosophila of the dunni subgroup inferred from Phylogenetics and Evolution 3: 240–247. multilocus DNA sequence variation. Evolution 57: 2566– Seutin G, Klein NK, Ricklefs RE, Bermingham E. 1994. 2579. Historical biogeography of the bananaquit (Coereba ﬂa- Wilgenbusch JC, Warren DL, Swofford DL. 2004. AWTY: veola) in the Caribbean region: a mitochondrial DNA assess- a system for graphical exploration of MCMC convergence in ment. Evolution 48: 1041–1061. Bayesian phylogenetic inference. Available at http://king2. Simon C, Frati F, Bechenbach A, Crespi B, Liu H, Flook scs.fsu.edu/CEBProjects/awty/awty_start.php P. 1994. Evolution, weighting, and phylogenetic utility of Williams EE. 1989. A critique of Guyer and Savage (1986): mitochondrial gene sequences and a compilation of con- cladistic relationships among anoles (Sauria: Iguanidae): served polymerase chain reaction primers. Annals of the are the data available to reclassify the anoles? In: Woods Entomological Society of America 87: 651–701. CA, ed. Biogeography of the West Indies: past, present, Slater J. 1988. Zoogeography of West Indian Lygaeidae and future. Gainesville, FL: Sandhill Crane Press, 433– (Hemiptera). In: Liebherr JK, ed. Zoogeography of 478.

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.

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 Aruba: Bringamosa, house of Roy Croes, sel_069, sel_070 12°39.547′N, 69°58.077′W, 14.X.2004, SCC04_041 2 Selenops n. sp. 5 Aruba: Luela, shooting range, 12°29.023′N, sel_072 69°57.778′W, 16.X.2004, SCC04_044 3 Selenops n. sp. 5 Aruba:N. sp. 5 National Park, near hill Gran sel_068 Tonel and Valley Rooi Coashati, 12°29.356′N, 69°55.461′W, 16.X.2004, SCC04_045 4 Selenops curazao Curaçao: Carmabi Institute, 12°07.351′N, sel_047, sel_058, sel_217 68°58.132′W, 7.X.2004, SCC04_026 5 Selenops curazao Curaçao: Girouette Plantation, E of Schottegat sel_048, sel_049, sel_050, sel_051, Harbour, house of A. DeBrot, 12°09′03.20″N, sel_052 68°54′56.35″W, 9.X.2004, SCC04_032 6 Selenops curazao Bonaire: Sabadaco, near cave, 12°11.587′N, sel_053 68°17.765′W, 11.X.2004, SCC04_033 7 Selenops curazao Bonaire: Altamira Ungu, 12°13.949′N, sel_059, sel_060, sel_082 68°20.703′W, 12.X.2004, SCC04_038 8 Selenops curazao Bonaire: Nort di Saliña, Kaya Otomac, sel_054, sel_056, sel_057, sel_061, 12°10′55.92″N, 68°16′28.39″W, 11.X.2004, sel_062, sel_063, sel_064, sel_065 SCC04_039 9 Selenops willinki Trinidad and Tobago: Little Tobago, sel_230, sel_231, sel_232, sel_233, 11°18′03.7″N, 60°30′11.1″W, 16.VII.2005, sel_234, sel_235, sel_236, SCC05_041 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 10 Selenops Trinidad and Tobago: St George Co., Point sel, 218, sel_219, sel_220, sel_221, geraldinae Gourde Road near Trinidad Military Base, sel_222, sel_223 near Chaguaramas off Chaguaramas Main Road, 10°40′47.1″N, 61°37′30.9″W, 11.VII.2005, SCC05_035 11 Selenops Trinidad and Tobago: Gaspar Grande Island sel_224, sel_225, sel_226, sel_227, geraldinae south of Chaguaramas, trails around island, sel_228, sel_229, sel_237, 10°39′46.5″N, 61°38′58.0″W, 12.VII.2005, sel_241, sel_257 SCC05_037 12 Selenops Trinidad and Tobago: Monos Island, South Sea, sel_244, sel_247, sel_248, sel_249, geraldinae 10°40′54.2″N, 61°41′21.6″W, 13.VII.2005, sel_250 SCC05_038 13 Selenops Trinidad and Tobago: Huevos Island, sel_239 geraldinae 10°41′28.3″N, 61°42′55.0″W, 13.VII.2005, SCC05_039 14 Selenops Trinidad and Tobago: Chacachacare Island, sel_246 geraldinae 10°41′24.2″N, 61°44′53.7″W, 13.VII.2005, SCC05_040 15 Selenops Argentina: Parque Nacional Chaco, Sendero MACN-Ar#12782 hebraicus peatonal. 25.II.2004. 16 Selenops occultus Brazil: São Paulo, Universidade São Paulo, sel_995 23°34′27.96″S, 46°40′21.29″W

APPENDIX Continued

Locality number Species Collection information Voucher numbers

17 Selenops occultus Brazil: Rio de Janeiro, Sierra da Carioca, sel_284 22°57′52.22″S, 43°16′31.34″W 18 Selenops occultus Brazil: Rio de Janeiro, Pao de Açucar, sel_283 22°55′22.05″S, 43°09′33.11″W 18 Selenops Brazil: Rio de Janeiro, Pao de Açucar, sel_277, sel_278, sel_279, sel_280, melanurus 22°55′22.05″S, 43°09′33.11″W sel_281, sel_282 19 Selenops St Vincent and the Grenadines: Young Island, sel_091, sel_113 micropalpus South of Villa Beach, 13°07.895′N, 61°12.142′W, 24.X.2004, SCC04_053 20 Selenops St Vincent and the Grenadines: King’s Hill sel_088, sel_090, sel_092, sel_093, micropalpus Forest Reserve, 13°08.825′N, 61°10.021′W, sel_094, sel_095, sel_112 27.X.2004, SCC04_055 21 Selenops St Lucia: Vieux Fort, on hill above airport, sel_832, sel_833, sel_834 micropalpus 13°44′20.2″N, 60°56′40.8″W, 13.III.2007 22 Selenops St Lucia: Anse la Ray, Ti-Kaye, 13°55′29.6″N, sel_820, sel_821, sel_822, sel_823 micropalpus 61°02′41.3″W, 13.III.2007, SCC07_046 23 Selenops St Lucia: Dennery: south of Dennery on east sel_825, sel_826, sel_827, sel_828, micropalpus coast road, eastern nature trail, heritage sel_829, sel_830, sel_831 tourism site, 13°53′50.8″N, 60°52′51.2″W, 13.III.2007, SCC07_047 24 Selenops St Lucia: Gros Islet, Pigeon Island, on top of sel_811, sel_812, sel_813, sel_814 micropalpus hill, 14°05′31.3″N, 60°57′03.8″W, 12.III.2007, SCC07_044 25 Selenops St Lucia: Gros Islet, Beausejour, past cricket sel_815, sel_816, sel_817, sel_818, micropalpus sponsor’s office, 14°04′43.1″N, 60°56′31.1″W, sel_819 12.III.2007, SCC07_045 26 Selenops Martinique: Le Diamant, Grand Anse du sel_804, sel_805, sel_806, sel_807, micropalpus Diamant, off of road D37, 14°28′32.9″N, sel_808, sel_809, sel_810 61°02′13.4″W, 10.III.2007, SCC07_043 27 Selenops Martinique: La Caravelle Reserve Naturelle, sel_791, sel_792, sel_793, sel_794, micropalpus trail to Pointe Caricoli, 14°46′09.3″N, sel_795, sel_797, sel_798, sel_799 60°53′24.7″W, 8.III.2007, SCC07_041 28 Selenops Martinique: Anse Ceran, off of road D-10, sel_800, sel_801, sel_802, sel_803 micropalpus 14°50′01.5″N, 61°13′24.7″W, 9.III.2007, SCC07_042 29 Selenops Dominica: Roseau: Botanical Park, top of hill sel_097, sel_098, sel_099, sel_100, micropalpus near shrine, 15°17.998′N, 61°22.754′W, sel_101 1.XI.2004, SCC04_059 30 Selenops Dominica: Jimmit-Warner, on top of hill with sel_102, sel_103, sel_104, sel_105 micropalpus cellular phone antenna, 15°22.690′N, 061°24.003′W, 2.XI.2004, SCC04_061 31 Selenops Dominica: Cabrits National Park, near trail to sel_106, sel_107, sel_108, sel_110, micropalpus fort, 15°35.049′N, 61°28.371′W, 2.XI.2004, sel_111, sel_135 SCC04_060 32 Selenops n. sp. 7 Guadeloupe: Les Saintes, top of Le Chameau, sel_783, sel_784, sel_785 15°51′28.1″N, 61°35′39.8″W, 6.III.2007, SCC07_039 33 Selenops n. sp. 7 Guadeloupe: Basse-Terre, Parc Archélogique sel_114 des Roches Gravées, near Trois-Rivieres, 15°58.394′N, 61°38.347′W, 10.XI.2004, SCC04_064

APPENDIX Continued

Locality number Species Collection information Voucher numbers

34 Selenops n. sp. 7 Guadeloupe: Basse-Terre, Trois Rivieres, end of sel_778, sel_779, sel_780, sel_781 Trail Sentier de l′Acomat off Rue Nelson Mandela, 15°58′03.0″N, 61°37′50.1″W, 5.III.2007, SCC04_038 35 Selenops n. sp. 7 Guadeloupe: Basse-Terre, near Vieux Fort on sel_115 the D6, along road at Forêt Domaniale du Litoral, 15°57.943′N, 061°42.517′W, 11.XI.2004, SCC04_065 36 Selenops n. sp. 7 Guadeloupe: Gran-Terre, Pointe du Chateaux, sel_786, sel_787, sel_788, sel_789, 16°14′51.6″N, 61°11′02.6″W, 7.III.2007, sel_790 SCC07_040 37 Selenops n. sp. 7 Montserrat: Jack Boy Hill, 17°46′02.1″N, sel_758, sel_759, sel_760, sel_761, 62°10′17.0″W, 2.III.2007, SCC07_035 sel_762, sel_763, sel_764, sel_765, sel_766, sel_767, sel_768 38 Selenops n. sp. 7 Montserrat: Sweet Water Ghaut, 16°47′07.2″N, sel_769 62°10′59.8″W, 2.III.2007, SCC07_036 39 Selenops n. sp. 7 Montserrat: Silver Hills, north side of Silver sel_771, sel_772, sel_773, sel_774, Hill, 16°48′41.3″N, 62°11′28.7″W, 3.III.2007, sel_775, sel_776, sel_777 SCC07_037 40 Selenops n. sp. 7 Antigua: Nelson’s Dockyard National Park, sel_754, sel_755, sel_756, sel_757 Shirley’s Heights lookout, 17°00′06.7″N, 61°44′57.6″W, 27.II.2007, SCC07_034 41 Selenops n. sp. 7 Antigua: Indian Town, east of Veranda Resort, sel_753 17°05′50.2″N, 61°40′53.0″W, 27.II.2007, SCC07_033 42 Selenops St Kitts and Nevis: Nevis, Tamarind Bay, sel_740, sel_741, sel_742 lindborgi Galliput Restaurant, 17°09′48.53″N, 62°37′50.02″W, 23.II.2007, SCC07_030 43 Selenops St Kitts and Nevis: Nevis, Round Hill entrance sel_735, sel_736, sel_737, sel_738, lindborgi to Mt Nevis, 17°11′13.83″N, 62°36′00.96″W, sel_739 23.II.2007, SCC07_029 44 Selenops St Kitts and Nevis: St Kitts, Major’s Bay, sel_750, sel_751, sel_752 lindborgi 17°13′37.9″N, 62°38′49.3″W, 24.II.2007, SCC07_032 45 Selenops St Kitts and Nevis: St Kitts, Sand Bank Bay, sel_743, sel_745, sel_746, sel_747, lindborgi 17°14′59.1″N, 62°38′40.8″W, 24.II.2007, sel_748, sel_749 SCC07_031 46 Selenops n. sp. 8 Saba: Giles Quarter Trail, 17°36′54.36″N, sel_1021, sel_1022 63°14′35.52″W, 12.III.2008 47 Selenops n. sp. 8 St Maarten: Mullet Bay, abandoned Mullet sel_712, sel_713, sel_718, sel_719, Bay Resort, 18°02′48.0″N, 63°027′29.7″W, sel_720, sel_721, sel_722, 21.III.2007, SCC07_027 sel_723, sel_724, sel_725, sel_726, sel_727, sel_728, sel_729, sel_730 48 Selenops n. sp. 8 St Maarten: Emilio Wilson Estate and Park, sel_711, sel_714, sel_715, sel_716, 18°02′32.7″N, 63°03′53.1″W, 20.II.2007, sel_717 SCC07_026 49 Selenops n. sp. 8 St Maarten: Upper Princess Quarter, sel_731, sel_732 18°01′48.0″N, 63°02′08.0″W, 21.III.2007, SCC07_028

APPENDIX Continued

Locality number Species Collection information Voucher numbers

50 Selenops n. sp. 8 St Maarten: south-east side of island, trail sel_116, sel_119, sel_120, sel_121, from Back Bay to Geneve Bay, 18°00.929′N, sel_708, sel_709, sel_710 63°01′840W, 12–13.XI.2004 and 20.II.2007, SCC04_066, 068, SCC07_025 51 Selenops St Maarten: Philipsburg, Front Street, near sel_117, sel_118 mexicanus entrance to cruise ship dock, 18°00.906′N, 63°02.587′W, 12.XI.2004, SCC04_067 52 Selenops n. sp. 8 Anguilla: Shoal Bay West, 18°09′52.8″N, sel_697 63°09′21.3″W, 13.II.2007, SCC07_016 53 Selenops n. sp. 8 Anguilla: The Cove, 18°10′14.1″N, sel_703, sel_704 63°07′52.6″W, 13.II.2007, SCC07_017 54 Selenops n. sp. 8 Anguilla: Long Bay, Long Bay Beach, sel_698, sel_699, sel_700, sel_701, 18°11′29.3″N, 63°07′49.7″W, 13.II.2007, sel_702 SCC07_015 55 Selenops n. sp. 8 Anguilla: Blowing Point, 18°10′18.0″N, sel_705, sel_706, sel_707 63°05′28.7″W, 13.II.2007, SCC07_018 56 Selenops n. sp. 8 Anguilla: Windward Point, 18°16′18.2″N, sel_696 62°58′05.3″W, 12.II.2007, SCC07_014 57 Selenops USVI: St Croix: Fredericksted, Sprat Hall sel_497, sel_498, sel_499, sel_500 lindborgi Beach, Rte.63, 17°44′09.8″N, 64°53′24.0″W, 14.VI.2006, SCC06_056 58 Selenops USVI: St Croix: Sprat Hall Hill off West Shore sel_494, sel_495, sel_496 lindborgi Road, ﬁrst right after subtracking station, 17°44′38.4″N, 64°53′22.3″W,14.VI.2006 SCC06_055 59 Selenops USVI: St Croix: Butler Bay, West Shore Road, sel_490, sel_491, sel_492, sel_493 lindborgi 17°45′49.7″N, 64°52′58.8″W, 14.VI.2006, SCC06_054 60 Selenops USVI: St Croix: intersecting road from Creque sel_501 lindborgi Dam to Mahogany Road (intersects Mount Victory Camp) 17°44′27.1″N, 64°51′25.4″W, 14.VI.2006, SCC06_057 61 Selenops USVI: St Croix: Radio Telescope Station, east sel_527 lindborgi island, 17°45.398′N, 64°35.045′W, 18.XI.2004, SCC04_072 62 Selenops Puerto Rico: Vieques: Laguna Kiani, sel_522, sel_523, sel_524, sel_525, lindborgi 18°07′02.2″N, 65°33′41.4″W, 19.VI.2006, sel_526 Selenops SCC06_065 insularis 63 Selenops Puerto Rico: Vieques: Ruinas Central Playa sel_516, sel_517, sel_518, sel_519, lindborgi Grande, 18°05′43.2″N, 65°31′13.2″W, sel_520, sel_521 Selenops 19.VI.2006, SCC06_064 insularis 64 Selenops Puerto Rico: Vieques: Cano Hondo, Cerca de sel_504, sel_505, sel_506, sel_507, lindborgi Puerto Mosquito, 18°06′11.0″N, 65°27′05.5″W, sel_508, sel_509, sel_510, Selenops 19.VI.2006, SCC06_061 sel_511, sel_512 insularis 65 Selenops Puerto Rico: Vieques: Refugio Nacional de Vida sel_513, sel_514, sel_515 lindborgi Silvestre, road to Playa Caracas, Laguna Puerto Ferro, 18°06′24″N, 65°25′25.8″W, 19.VI.2006, SCC06_063

APPENDIX Continued

Locality number Species Collection information Voucher numbers

66 Selenops Puerto Rico: Culebra: Monte Resaca, sel_480, sel_481, sel_482, sel_483, lindborgi 18°19′30.7″N, 65°18′10.5″W, 12.VI.2006, sel_484, sel_485 SCC06_052 67 Selenops Puerto Rico: Culebra: Brava Beach Trail, sel_486, sel_487, sel_488, sel_489 lindborgi 18°19′38.9″N, 65°16′54.1″W, 12.VI.2006, SCC06_053 68 Selenops USVI: St Thomas, Estate Perserverance, sel_076 lindborgi Perserverance Bay Trail, 18°21.463′N, 64°59.753′W, 22.X.2004, SCC04_050 69 Selenops USVI: St Thomas, St Peter, house of R. sel_131 lindborgi Platenberg, 18°21′22.17″N, 64°56′49.53″W, 23.X.2004, SCC04_075 70 Selenops USVI: St Thomas, Magen’s Bay Trail, sel_124, sel_125, sel_128, sel_129, lindborgi 18°21.350′N, 64°55.231′W, 22.X.2004 and sel_130, sel_164 9.XI.2004, SCC04_052 and SCC04_073 71 Selenops USVI: St Thomas, Benner Hill, above armory, sel_123, sel_132, sel_133, sel_134 lindborgi 18°19.533′N, 64°51.703′W, 19.XI.2004, SCC04_074 72 Selenops USVI: St Thomas, East End, Estate Nazareth, sel_143, sel_144, sel_145, sel_146, lindborgi Dolphin House, 18°19.128′N, 64°51.567′W, sel_147 19.XI.2004, SCC04_076 73 Selenops USVI: St John, Bordeaux Ridge Road, sel_140, sel_141, sel_142, sel_157 lindborgi 18°20.125′N, 64°43.672′W, 17.XI.2004, SCC04_071 74 Selenops USVI: St John, Cinnamon Bay Loop Trail, sel_126, sel_127 lindborgi 18°21.226′N, 64°45.259′W, 16-17.XI.2004, SCC04_070 75 Selenops USVI: St John, Leinster Bay Trail, sel_136, sel_137, sel_138, sel_139 lindborgi 18°21.825′N, 64°43.743′W, 16.XI.2004, SCC04_069 76 Selenops BVI: Tortola: vic. Sage Mountain, sel_078 lindborgi 18°24′46.44″N, 64°39′18.43″W, 20.X.2004, SCC04_049 77 Selenops BVI: Guana Island: north side near beach sel_071, sel_081, sel_085, sel_089 lindborgi house, 18°28.793′N, 64°34.473′W, 18.X.2004, SCC04_047 78 Selenops BVI: Guana Island: south side near salt pond, sel_079, sel_084, sel_086, sel_087 lindborgi 18°28.619′N, 64°34.475′W, 18.X.2004, SCC04_046 79 Selenops BVI: Virgin Gorda, lower trail up Gorda Peak, sel_073, sel_074, sel_075, sel_083, lindborgi 18°28.774′N, 64°24.210′W, 19.X.2004, sel_089, sel_096, sel_109 SCC04_048 80 Selenops Cuba: Sierra de Mesa, Pinar del Rio sel_276 submaculosus 81 Selenops simius Cayman Islands: Grand Cayman, Queen sel_046, sel_066, sel_067 Elizabeth II Botanic gardens, storage facility south of main building, 19°19.055′N, 81°09.527′W, 30.IX.2004, SCC04_021 82 Selenops simius Cayman Islands: Grand Cayman, Queen sel_080 Elizabeth II Botanic gardens, tree trail behind iguanas, 19°19.042′N, 81°10.081′W, 2.X.2004, SCC04_022

APPENDIX Continued

Locality number Species Collection information Voucher numbers

83 Selenops simius Cayman Islands: Little Cayman, on road sel_022 across street from Pirate’s Point Resort, 19°39.754′N, 80°06.032′W, 3.X.2004, SCC04_023 84 Selenops simius Cayman Islands: Cayman Brac, National Trust sel_023, sel_024, sel_025, sel_026, House off West End Road, 19°42.019′N, sel_027, sel_028 79°52.084′W, 3.X.2004, SCC04_025 85 Selenops Jamaica: Westmoreland Paris, near New Hope sel_362 candidus on road toward Savanna-la-Mar, 18°14′55.4″N, 78°14′41.0″W, 29.V.2006, SCC06_027 86 Selenops Jamaica: St Ann Parish, North Coast Highway, sel_357, sel_358, sel_359, sel_360 candidus between Discovery Bay and Rio Bueno, 18°28′31.3″N, 77°25′49.0″W, 28.V.2006, SCC06_024 87 Selenops Jamaica: Clarendon Parish, off road to sel_363, sel_364 candidus Lluidasvale, 18°07′50.8″N, 77°10′05.0″W, 31 May 2006, SCC06_028 88 Selenops n. sp. Jamaica: St Catherine Parish, Hellshire Hills, sel_376, sel_377, sel_378, sel_379, 17 A2 Depression, 17°51′59.3″N, 76°57′54.0″W, sel_380, sel_381, sel_382, 3.VI.2006, SCC06_031 sel_383, sel_384 89 Selenops Jamaica: St Mary Parish, near Mango Valley, sel_352, sel_353, sel_354, sel_355, candidus 1.6 km off North Coast Highway, sel_356 18°24′23.4″N, 77°02′37.6″W, 28.V.2006, SCC06_023 90 Selenops Jamaica: St Andrew Parish, Castleton Botanic sel_350, sel_351 candidus Gardens, 18°10′20.3″N, 76°49′27.6″W, 27.V.2006, SCC06_022 91 Selenops Jamaica: St Andrew Parish, Hermitage Dam sel_385 candidus Road, 2–6 km from junction with Stony Hill, 18°04′25.4″N, 76°47′01.3″W, 5.VI.2006, SCC06_033 92 Selenops Jamaica: St Thomas Parish, Blue Mountains sel_365, sel_366, sel_367, sel_368, petrunkevitchi National Park, Whitﬁeld Hall, 18°02′54.8″N, sel_369, sel_370, sel_371, 76°37′03.7″W, 1.VI.2006, SCC06_029 sel_372, sel_373, sel_374, sel_375, sel_376 93 Selenops n. sp. Jamaica: St Thomas Parish, near 12 mile Bull sel_386, sel_387, sel_388, sel_389, 16 Bay, on left side of road heading east, sel_390, sel_391 17°55′32.5″N, 76°38′31.0″W, 5.VI.2006, SCC06_034 94 Selenops Hispaniola: Haiti: Jacmel, St Cyr 72°31′41.2″N, sel_655, sel_656, sel_657, sel_658, insularis 18°14′16.6″W, 23.X.2006, SCC06_078 sel_659, sel_660, sel_661, Selenops sel_662, sel_663, sel_664, sel_665 pensilis 95 Selenops n. sp. Hispaniola: Dominican Republic: Prov. sel_156 13 Pedernales: road to Aguacate from Rio Mulito, 18°13.895′N, 71°45.190′W, 25.XI.2004, SCC04_082 96 Selenops Hispaniola: Dominican Republic: Prov. sel_150, sel_151, sel_152, sel_153, marcanoi Pedernales, Rio Mulito (El Banano), sel_154, sel_155 18°09.165′N, 071°45.388′W, 25.XI.2004, SCC04_081

APPENDIX Continued

Locality number Species Collection information Voucher numbers

97 Selenops phaselus Hispaniola: Dominican Republic: Prov. sel_158, sel_160, sel_632, sel_633, Pedernales, Parque Nacional Sierra de sel_634 Baoruco, Las Abejas, 18°08.804′N, 71°37.164′W, 24-Nov-04 and 10-Oct-06, SCC04_077 and SCC06_072 98 Selenops phaselus Hispaniola: Dominican Republic: Prov. sel_148, sel_159, sel_215 Pedernales, 26 km north of Cabo Rojo, Sierra de Baoruco, 18°06.490′N, 71°37.316′W, 24.XI.2004, SCC04_078 99 Selenops n. sp. Hispaniola: Dominican Republic: Prov. sel_161, sel_162, sel_163, sel_165, 14 Pedernales, 13–14 km north of Cabo Rojo on sel_166, sel_167 Carretera ALCOA, 18°01.962′N, 71°38.748′W, 24.XI.2004, SCC04_079 100 Selenops Hispaniola: Dominican Republic: Prov. sel_586, sel_589 insularis Pedernales, Parque Jaragua, VII.2006 101 Selenops n. sp. Hispaniola: Dominican Republic: Prov. sel_630, sel_631, sel_635, sel_636, 14 Pedernales, Boca de la Cañada, sel_637 Selenops n. sp. Pedernales-N. sp. 15 Road, 9.X.2006, 15 SCC06_071 102 Selenops Hispaniola: Dominican Republic: Prov. sel_018 insularis Pedernales, N. sp. 15, Fondo de Mama Cocoño, 25.X.2003, SCC03_021 103 Selenops n. sp. Hispaniola: Dominican Republic, Prov. sel_621, sel_622, sel_633, sel_634, 15 Pedernales, Laguna N. sp. 15, El Cajuil, sel_635, sel_636, sel_637, sel_638 9.X.2006, SCC06_070 104 Selenops Hispaniola: Dominican Republic: Prov. sel_169, sel_170, sel_171, sel_172, insularis s. n. Barahona, Carretera Higuero-Polo, sel_173 sp. 1 26.XI.2004, SCC04_083 105 Selenops phaselus Hispaniola: Dominican Republic: Prov. sel_562, sel_565, sel_566, sel_568, Selenops n. sp. Barahona, Polo Coffee Plantation, July 2006 sel_585 13 106 Selenops Hispaniola: Dominican Republic: Prov. sel_537, sel_538, sel_539, sel_540, insularis Barahona, Barahona, CoralSol Resort, San sel_541, sel_542, sel_543, Rafael Beach, July 2006 sel_544, sel_545, sel_561, sel_563, sel_564, sel_567, sel_573, sel_574, sel_575, sel_576, sel_587, sel_588 107 Selenops Hispaniola: Dominican Republic: Prov. Peravia, sel_175 insularis Bani, road from Bani to Manaclar, past La Laguna, 18°21.343′N, 70°21.077′W, 27.XI.2004, SCC04_086 108 Selenops Hispaniola: Dominican Republic: Prov. Peravia, sel_176, sel_177, sel_178, sel_179 insularis Bani, Rio Nizao, 18°16.915′N, 70°12.101′W, 27.XI.2004, SCC04_087 109 Selenops Hispaniola: Dominican Republic: Prov. San sel_010, sel_011, sel_012, sel_013, insularis Cristóbal, Engombe, farm of autonomous sel_014, sel_015, sel_016, sel_017 university, 18°27.360′N, 70°00.306′W, 27.X.2003, SCC03_023 110 Selenops Hispaniola: Dominican Republic: Prov. La sel_200, sel_201, sel_202, sel_203, lindborgi Altagracia, Parque del Este, Guaraguao, sel_204, sel_205, sel_206, sel_207 18°19.968′N, 68°48.709′W, 30.XI.2004, SCC04_090

APPENDIX Continued

Locality number Species Collection information Voucher numbers

111 Selenops Hispaniola: Dominican Republic: Prov. La sel_191, sel_192, sel_193, sel_194, lindborgi Altagracia, Parque del Este, Boca de Yuma, sel_195, sel_196, sel_197, Selenops 18°21.875′N, 68°37.080′W, 29-30.XI.2004, sel_199, sel_216 insularis SCC04_089 112 Selenops Hispaniola: Domnican Republic: Prov. La sel_528, sel_529, sel_530, sel_531, lindborgi Altagracia, Punta Cana Resort, sel_532, sel_533, sel_534, Selenops 18°30′55.53″N, 68°22′28.73″W, 5-9.VII.2006, sel_535, sel_536 insularis SCC04_066 Selenops n. sp. 9 113 Selenops Hispaniola: Dominican Republic: Prov: Hato sel_555 insularis Major, Los Haitises, 1 km south of El Valley near Hato Mayor, 18°58′24.01″N, 69°22′34.58″W, VI.2006 114 Selenops Hispaniola: Dominican Republic: Prov. sel_577, sel_590, sel_591, sel_592, lindborgi Samaná, Las Terrenas, 19°19′28.88″N, sel_593, sel_594 Selenops 69°32′50.77″W, VI.2006 insularis 115 Selenops n. sp. 2 Hispaniola: Dominican Republic: Prov. La sel_546 Vega, Constanza, Alto Cerro Hotel, 18°54′14.16″N, 70°44′41.14″W, VI.2006 116 Selenops Hispaniola: Dominican Republic: Prov. Puerto sel_578 insularis Plata, Sosua, VI.2006 117 Selenops Hispaniola: Dominican Republic: Prov. Puerto sel_604, sel_605, sel_606, sel_607, insularis Plata, entrance to Loma de Isabel Torres, sel_608, sel_609 Selenops n. sp. 19°46′41.9″N, 70°42′01.1″W, 7.X.2006, 12 SCC06_068 118 Selenops n. sp. Hispaniola: Dominican Republic, Prov. sel_640, sel_641, sel_642, sel_643, 13 Santiago, Mata Grande, 19°11′43.0″N, sel_644, sel_645, sel_646 70°59′42.0″W, 14–15.X.2006, SCC06_075 119 Selenops n. sp. Hispaniola: Dominican Republic, Prov. sel_639 13 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_074 120 Selenops Hispaniola: Dominican Republic, Prov. Monti sel_579, sel_580, sel_581, sel_582, insularis Cristi, Monte Cristi, El Morro, sel_583, sel_584, sel_610, Selenops n. sp. 19°53′42.44″N, 71°39′14.48″W, 8.X.2006, sel_611, sel_612, sel_613, 10 SCC06_069 sel_614, sel_615, sel_616, sel_617, sel_618 121 Selenops Hispaniola: Dominican Republic: Prov. sel_180, sel_181, sel_182, sel_183, insularis Independencia, La Descubierta, El Azufrada, sel_184, sel_185, sel_186, Selenops north side of Lago Enriquillo, 18°33.751′N, sel_187, sel_188, sel_189, pensilis 71°41.853′W, 26.XI.2004, SCC04_084 sel_190, sel_569, sel_570, Selenops bani sel_571, sel_572, sel_599 122 Selenops phaselus Hispaniola: Haiti: Kenskoff, Belot-Montcel, sel_647, sel_648, sel_649, sel_650, 18°27′11.3″N, 72°21′06.4″W, 20-21.X.2006, sel_651, sel_652, sel_653, sel_654 SCC06_076 123 Selenops n. sp. 3 Puerto Rico: Isla Mona: Bajura de Empalme, sel_846, sel_847 18°06′25.07″N, 67°53′10.41″W 124 Selenops Puerto Rico: Isla Mona: Sardinera, sel_838, sel_839, sel_840, sel_841, insularis 18°05′46.76″N, 67°56′12.35″W sel _842, sel_843, sel_844, sel_845

APPENDIX Continued

Locality number Species Collection information Voucher numbers

125 Selenops n. sp. 3 Puerto Rico: Isla Mona: Camino de los Cobros, sel_502 18°04′02.68″N, 67°52′45.75″W, VII.2006 126 Selenops n. sp. 3 Puerto Rico: Maricao, Bosque Estatl de sel_434, sel_435, sel_436, sel_437, Selenops Maricao, 18°08′51.2″N, 66°59′35.0″W, sel_438, sel_439, sel_440, insularis 10.VI.2006, SCC06_045 sel_441, sel_442 Selenops lindborgi 127 Selenops Puerto Rico: Susua State Forest, Sabana sel_443, sel_444, sel_445, sel_446, lindborgi Grande, 18°04′15.0″N, 66°54′31.6″W, sel_447, sel_448, sel_449, Selenops 10.VI.2006, SCC06_046 sel_450, sel_451, sel_452, sel_453 insularis 128 Selenops Puerto Rico: Quebradillas, Merendero de sel_423, sel_424, sel_425, sel_426, lindborgi Guajataca, 18°29′23.7″N, 66°56′59.4″W, sel_427, sel_428, sel_429, Selenops 9.VI.2006, SCC06_043 sel_430, sel_431, sel_432 insularis 129 Selenops Puerto Rico: Arecibo, Arenalejos, Carretera sel_392, sel_393, sel_394, sel_395, lindborgi 657, km 1.9, 18°25′15.9″N, 66°40′35.2″W, sel_396, sel_397, sel_398 Selenops 7.VI.2006, SCC06_035 insularis 130 Selenops Puerto Rico: between Barceloneta and Arecibo, sel_419, sel_420, sel_421, sel_422 lindborgi Bosque Cambalacheo, 18°27′07.0″N, Selenops 66°35′49.9″W, 9.VI.2006, SCC06_041 insularis 131 Selenops Puerto Rico: Ciales, Bosque Fronton, Carretera sel_503 insularis 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_060 132 Selenops Puerto Rico: Coamo, Baños de Coamo, sel_462, sel_463 insularis 18°02′19.4″N, 66°22′27.0″W, 11.VI.2006, SCC06_048 133 Selenops Puerto Rico: Salina, Reserva Jobos, Parque sel_464 lindborgi Jagueys, 17°57′13.9″N, 66°15′03.5″W, 11.VI.2006, SCC06_049 134 Selenops Puerto Rico: Toa Baja, Bosque Media Luna, sel_461 insularus PR-2, km 21–6, 18°24′38.30″N, 66°14′44.36″W, 14.VI.2006, SCC06_058 135 Selenops Puerto Rico: Manuabo, Mariani Creek, sel_465, sel_466, sel_467, sel_468 lindborgi 18°00′29.7″N, 65°52′17.0″W, 11.VI.2006, Selenops SCC06_050 insularis 136 Selenops Puerto Rico: Humacao, Barrio Collores, sel_469, sel_470, sel_471, sel_472, insularis 18°09′44.82″N, 65°49′06.50″W, 11.VI.2006, sel_473, sel_474, sel_475, SCC06_051 sel_476, sel_477, sel_478, sel_479 137 Selenops Puerto Rico: Loiza, Punta Vacia, Talega, sel_399, sel_400, sel_401, sel_401, lindborgi 18°27′03.8″N, 65°54′16.7″W, 8.VI.2006, sel_402, sel_403 SCC06_038 138 Selenops Puerto Rico: Ceiva, Los Corchos, 18°12′13.8″N, sel_416, sel_417, sel_418 lindborgi 65°40′06.5″W, 8.VI.2006, SCC06_040 Selenops insularis

APPENDIX Continued

Locality number Species Collection information Voucher numbers

139 Selenops Puerto Rico: Fajardo, Seven Seas Public Beach, sel_405, sel_406, sel_407, sel_408, lindborgi 18°22′03.7″N, 65°38′04.9″W, 8.VI.2006, sel_409, sel_410, sel_411, Selenops SCC06_039 sel_412, sel_413, sel_414, sel_415 insularis 140 Selenops Bahamas: Andros Island: Owens Town, sel_302, sel_303, sel_304, sel_305, submaculosus 24°52′30.1″N, 78°02′03.6″W, 13.V.2006, sel_306, sel_307, sel_308 SCC06_002 141 Selenops Bahamas: Andros Island: Morgan’s Cave at sel_309, sel_310, sel_311, sel_666 submaculosus Morgan’s Bluff, 25°10′30.1″N, 78°01′26.2″W, 13.V.2006, SCC06_003 142 Selenops Bahamas: Andros Island: International Field sel_286, sel_287, sel_288, sel_289, submaculosus Station, 24°53′51.1″N, 77°55′50.1″W, sel_290, sel_291, sel_292, 12.V.2006, SCC06_001 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_836 143 Selenops Bahamas: Pigeon Cay, near IFS on Andros sel_312 submaculosus Island, 24°52′54.4″N, 77°53′53.5″W, 13.V.2006, SCC06_004 144 Selenops Bahamas: Andros Island: Cargill Creek, sel_668 submaculosus 24°30′00.37″N, 77°43′15.61″W, 13.V.2006 145 Selenops aissus Bahamas: Abaco: Ralph’s Chimney off Queen’s sel_315 (Abaco) Highway, 26°14′58.2N, 77°11′25.4″W, 14.V.2006, SCC06_006 146 Selenops aissus Bahamas: Abaco: Abaco National Park, sel_313 26°03′44.0″N, 77°12′46.2″W, 14.V.2006, SCC06_005 147 Selenops Bahamas: Great Exuma: Bahamas Sound sel_332 submaculosus Subdivision near old airport, 23°27′56.0″N, 75°46′24.8″W, 18.V.2006, SCC06_011 148 Selenops aissus Bahamas: Great Exuma: Regatta Point, sel_319, sel_320, sel_321, sel_322, 23°30′24.7″N, 75°45′58.0″W, 18.V.2006, sel_323, sel_324 SCC06_009 149 Selenops aissus Bahamas: Stocking Island, near Great Exuma, sel_325, sel_326, sel_327, sel_328, 23°32′08.9″N, 75°46′29.6″W, 18.V.2006, sel_329, sel_330, sel_331 SCC06_010 150 Selenops aissus Bahamas: San Salvador: Gerace Field Station, sel_333, sel_334, sel_335, sel_336, trails behind ﬁeld station, 24°06.9′N, sel_337, sel_338, sel_339, 74°27.8′W, 19.V.2006, SCC06_012 sel_340, sel_341, sel_342, sel_343 151 Selenops Bahamas: Great Inagua: Man o’War Bay, sel_316 lindborgi 21°04′30.2″N, 73°38′36.7″W, 16.V.2006, SCC06_007 152 Selenops Bahamas: Great Inagua: Old Aerostat Base, sel_317, sel_318 lindborgi 21°06.06.7″N, 73°39′01.9″W, 16.V.2006, SCC06_228 153 Selenops n. sp. 11 Turks and Caicos: Providenciales, North-West sel_689, sel_690, sel_691, sel_692 Point Pond Nature Reserve, 21°50′32.1″N, 72°19′43.7″W, 8.II.2007, SCC07_010 and SCC07_011

APPENDIX Continued

Locality number Species Collection information Voucher numbers

154 Selenops n. sp. 11 Turks and Caicos: Providenciales, Turtle Cove, sel_346, sel_347, sel_348, sel_349, Third Turtle Drive, 21°47′01.1″N, sel_694, sel_695 72°13′45.4″W, 22.V.2006 and 10.II.2007, SCC06_017 and SCC07_013 155 Selenops n. sp. 11 Turks and Caicos: Providenciales, The Bight, sel_344, sel_345 21°47′00.6″N, 72°13′06.4″W, 22.V.2006, SCC06_015 156 Selenops n. sp. 11 Turks and Caicos: North Caicos, Wade’s Green sel_675, sel_680, sel_681, sel_682, Plantation, 21°55′13.36″N, 72°01′12.45″W, sel_683, sel_684, sel_685, 2–3.II.2007, SCC07_001 and SCC07_005 sel_686, sel_687, sel_688, sel_689 157 Selenops n. sp. 11 Turks and Caicos: Middle Caicos, Garden Pond sel_676, sel_677, sel_678, sel_679 Field Road, 21°48′24.76″N, 71°45′42.78″W, 3.II.2007, SCC07_004 158 Selenops n. sp. 11 Turks and Caicos: Providenciales, South View sel_693 Drive off of Leeward Highway, 21°46′45.7″N, 72°13′45.4″W, 10.II.2007, SCC07_012 159 Selenops debils USA: California, San Diego Co., Upper Otay sel_002 gp. species 1 River Valley, 32°59′50.33″N, 116°19′09.27″W, 19.V.2003 160 Selenops debils USA: California, San Diego Co., Jamul, Lyons sel_837 gp. species 1 Valley, north of Lyons Peak, 32°44′03.93″N, 116°53′51.23″W, 29.VII.2007 161 Selenops debils USA: California, San Diego Co., Anza Borrego sel_021, sel_214 gp. species 1 Desert State Park, Carrizo Palm Grove, 32°44′31.29″N, 116°12′51.11″W 162 Selenops debils Mexico: Baja California Sur, 2 km east of sel_210, sel_213 gp. species 1 Ballena between San Ignacio and San Juanico, 26°27′10.60″N, 111°34′53.39″W 163 Selenops debils Mexico: Baja California Sur, 3 km west of Villa sel_212 gp. species 1 Insurgentes on road to San Miguel de Comundu, 25°16′50.46″N, 111°50′03.75″W 164 Selenops debils Mexico: Baja California Sur, Cuevas Pintas, sel_009 gp. species 1 26°01′38.63″N, 111°30′24.45’W 165 Selenops debils USA: Arizona, Coconino Co., Monument Trail, sel_211 gp. species 1 ﬂat near archaeological site, 36°25.309′N, 112°27.483′W, 13.VIII.2004, SCC04_013 166 Selenops debils USA: Arizona: Santa Cruz Co., Madera sel_264, sel_270, sel_271, sel_272 gp. species 2 Canyon, Mt Wrightson Trail, 31°43′06.86’N, 110°52′22.45’W 167 Selenops debils Mexico: Sonora, Sonoran Highway on road to sel_851 gp. species 3 Yecora, 28°23′15.01’N, 108°55′41.42’W 168 Selenops debils USA: Texas: Val Verde Co., Seminole Canyon, sel_208 gp. species 3 Highway 90, under bridge, 29°42′21.12’N, 101°18′28.48’W 169 Selenops Mexico: Hidalgo, Villa Flores, Ejido ′El Rayo’, sel_1008 mexicanus 24°14′1.00’N, 99°1′4.50’W, 2.XI.2007 170 Selenops abyssus Mexico: Colima, Manzanillo, Municipio sel_1004 Manzanillo, 1.2 to 1.4 km east La Central, 19°8′56.4’N, 104°25′35.3’W, X.2005 171 Selenops abyssus Mexico: Colima, Municipio Ixtlahuacan, sel_1013 Tamala, 19°5′2.13’N, 103°47′26.21″

APPENDIX Continued

Locality number Species Collection information Voucher numbers

172 Selenops abyssus Mexico: Michoacan, Municipio Coalcomán, sel_1006, sel_1012 Coalcomán, 18°24′12.81″N, 103°07′58.82″W, X.1005 173 Selenops n. sp. 4 Mexico: Morelos, Cuernavaca, Colonia sel_1010 Chamilpa, Instituto de Biotecnología, UNAM, 18°55′51.86″N, 99°14′16.60″W 174 Selenops gracilis Mexico: Guerrero, Arcelia, Campo Morado, sel_1014, sel_1015 17°34′60.00″N, 100°4′60.00″W 175 Selenops Mexico: Guerrero, Omiltemi, Chilpancingo, sel_1007 nigromaculatus? Camino al Omiltemi, 17°33′3.76″N, This is an 99°30′21.83″W immature specimen, but is found near the type locality of S. nigromaculatus, hence the ‘?’ 176 Selenops n. sp. 4 Mexico: Puebla, Municipio Zapotitlan de las sel_1002, sel_1005 Salinas, Cerro el Pajarito, 18°22′48.3″N, 97°30′26.9″W, V.2005 177 Selenops Mexico: Chiapas, Tuxtla-Gutierrez, Cañon 1016, 1017 mexicanus deSumidero, 16°49′41.46″N, 93°6′22.22″W 178 Selenops Mexico: Chiapas, Berriozabal, dirt road from sel_848 mexicanus Efrain A. Gutierrez, approx. 8 km north Berriozabal, 16°52′27.45″N, 93°17′28.20″W 179 Selenops n. sp. Mexico: Chiapas, Pueblo Nueva Solistahuacan, sel_043 19 17°11.550′N, 92°54.875′W, 17.IX.2004, SCC04_018 180 Selenops Mexico: Chiapas, La Reforma, Municipio La sel_044 mexicanus Concordia, 15°54.212′N, 92°40.157′W, 18.IX.2004, SCC04_018b 181 Selenops Mexico: Chiapas, Huixtla, Las Golindrinas, sel_031, sel_034, sel_035, sel_036, mexicanus 15°25.747′N, 92°39.270′W, 23.IX.2004, sel_037, sel_038, sel_039, Selenops n. sp. SCC04_020 sel_040, sel_041, sel_045, 19 sel_1011 182 Selenops Mexico: Chiapas, Municipio Motozintla de sel_029, sel_030, sel_032, sel_033, mexicanus Mendoza, Chevolcan, 15°20′52.4″N, sel_038 Selenops n. sp. 92°19′25.4″W, 21.XI.2004, SCC04_019 19 183 Selenops Mexico: Chiapas: road to Roberto Barrio, sel_849 mexicanus ~4 km south-west Nuevo Sonora, 17°23′41.10″N, 91°54′10.70″W 184 Selenops Mexico: Veracruz, Municipio Tamalin, El sel_1018, sel_1019, sel_1020 mexicanus Mamey, 21°31′13.63″N, 97°38′31.75″W 185 Selenops Guatemala: Petén, Sta Elena de la Cruz, sel_865, sel_866, sel_867, sel_868 mexicanus Colonia del Bosque, near Flores, Cueva Selenops n. sp. Actun Kan, 16°54′10.9″N, 89°53′44.3″W, 18 1.I.2008, SCC08_001 186 Selenops Guatemala: Zacatán, Las Guacamayas, sel_869, sel_870, sel_871, sel_872, bifurcatus Carretera Sta Rosalia Marmol, sel_873, sel_874, sel_875, Hídroelectrica Pasabíen, 15°01′39.7″N, sel_876, sel_877, sel_878, sel_879 89°41′41.2″W, 1.I.2008, SCC08_002

APPENDIX Continued

Locality number Species Collection information Voucher numbers

187 Selenops El Salvador: Dep. Chaletenango, Chaletenango sel_927 bifurcatus off Carretera Troncal del Norte, Hotel Maya, 14°16′29.1″N, 89°08′32.7″W, 5-6.I.2008, SCC08_009 188 Selenops El Salvador: Dep. Chaletenango, Mun. sel_925, sel_926, sel_927, sel_928, bifurcatus Chaletenango, La Cueva del Corridor, sel_929 6.I.2008, SCC08_010 189 Selenops El Salvador: Dep. Chaletenango, Chaletenango, sel_921, sel_922, sel_923, sel_924 bifurcatus Municipio Tejutto, Rest. Eucalyptos, Selenops 14°12′20.4″N, 89°06′43.9″W, 5.I.2008, mexicanus SCC08_008 190 Selenops El Salvador: San Salvador, Museo Nacional de sel_881, sel_882, sel_883, sel_884, bifurcatus Historia Natural grounds, 13°40′23.4″N, sel_885, sel_886, sel_887, 89°11′53.6″W, 3.I.2008, SCC08_003 sel_888, sel_889 191 Selenops El Salvador: near San Vicente, Mun. Tepetitán sel_897, sel_898, sel_899, sel_900, bifurcatus vic. Finca El Carmen, 13°37′53.0″N, sel_901, sel_902, sel_903, Selenops 88°50′19.5″W, 4.I.2008, SCC08_005 sel_904, sel_905, sel_906, mexicanus sel_907, sel_908, sel_909, sel_910, sel_911 192 Selenops El Salvador: Dep. San Vicente, vic. San sel_890, sel_891, sel_892, sel_893, bifurcatus Vicente, road to Zacatecoluca, behind strip sel_894, sel_895, sel_896 club ‘Dreamed Girl’, 13°37′43.4″N, 88°46′49.6″W, 3.I.2008, SCC08_004 193 Selenops El Salvador: Dept La Union, Mun. El Carmen, sel_916, sel_917, sel_918, sel_919, bifurcatus Lotiﬁcacion Amaya, 13°21.44.9″N, sel_920 87°59′58.2″W, 5.I.2008, SCC08_007 194 Selenops El Salvador: Dept La Union, Conchagua, sel_912, sel_913, sel_914, sel_915 bifurcatus Volcán Conchagua near La Union, 13°18′14.1″N, 87°51′19.6″W, 4.I.2008, SCC08_006 195 Selenops Nicaragua: Dept Nuevo Segovia, Alc. Ocotal, sel_930 bifurcatus Barrio Roberto Gomez above Rio Coco, 13°37′05.8″N, 86°27′57.3″W, 11.I.2008, SCC08_012 196 Selenops Nicaragua: Dept Madríz, Alc. Ocotal, sel_931, sel_932, sel_933, sel_934, bifurcatus Totogalpa, 13°33′49.5″N, 86°29′54.6″W, sel_935, sel_936, sel_937, Selenops 11.I.2008, SCC08_013 sel_938, sel_939 mexicanus 197 Selenops Nicaragua: Dept Leon, Alc. San Jacinto, Mina sel_954, sel_955, sel_956, sel_957, bifurcatus El Límon, Rancho Las Brisas, 12°37′03.8″N, sel_958, sel_959, sel_960 86°44′34.3″W, 14.I.2008, SCC08_016 198 Selenops Nicaragua: Dept Leon, Alc. El Jicaral, Camino sel_961, sel_962, sel_963, sel_964, bifurcatus Santa Rosa, Puente La Guayabita, sel_965, sel_966, sel_967, 12°44′31.2″N, 86°22′44.6″W, SCC08_017, sel_968, sel_969, sel_970 SCC08_017 199 Selenops Nicaragua: Dept Matagalpa, Alc. San Ramon, sel_971, sel_972, sel_973, sel_974 mexicanus Mata Palo, 12°56′16.5″N, 85°51′12.2″W, 14.I.2008, SCC08_018 200 Selenops Nicaragua: Dept Boaco, Aguas calientes, Alc. sel_963, sel_975, sel_976, sel_977, bifurcatus Teustepe, Camino La Cuesta, 12°22′57.8″N, sel_978, sel_979, sel_980, Selenops 85°47′30.7″W, 15.I.2008, SCC08_020 sel_981, sel_982, sel_983, mexicanus sel_985, sel_986

APPENDIX Continued

Locality number Species Collection information Voucher numbers

201 Selenops Nicaragua: Lago Nicaragua, Isla Ometepe, sel_940, sel_941, sel_942, sel_943, bifurcatus Volcán Concepción, Charco Verde, Hotel sel_944, sel_945, sel_946, Selenops Finca Vincenzia and up hill across the sel_947, sel_948, sel_949, mexicanus street, 11°28′42.6″N, 35°38′20.6″W and sel_950, sel_951, sel_952, sel_953 11°29′31.2″N, 85°38′14.1″W, 12-13.I.2008, SCC08_014, SCC08_015 202 Selenops Costa Rica: Guanacaste, Palo Verde National sel_989, sel_990 mexicanus Park, Cueva Las Tigres, 10°21′58.9″N, 85°21′14.2″W, 17.I.2008, SCC08_022 203 Selenops Costa Rica: Guanacaste: Palo Verde Field sel_987, sel_988 bifurcatus Station, hill behind OTES office, Selenops 10°20′42.5″N, 85°20′19.1″W, 17.I.2008, mexicanus SCC08_021 204 Selenops Costa Rica: Guanacaste: Nicoya Peninsula, sel_991, sel_992, sel_993, sel_994 mexicanus near Loma Bonita, 10°15′04.0″N, 85°17′30.5″W, 18.I.2008, SCC08_023 205 Selenops banksi Panamá: Barro Colorado Island, I.2008 sel_1000, sel_1001 206 Selenops Panamá: STRI, Galeta, Plot F, 2004 sel_265, sel_266, sel_267, sel_268 mexicanus 207 Selenops Japan: Kyoto, Shugakuin, 23.VIII.2007 sel_861, sel_862, sel_863 bursarius 208 Selenops radiatus Namibia: 12.VI.2006 sel_998, 999 209 Selenops radiatus Tanzania: Iringa, Lutheran House Hostel sel_997 210 Anyphops South Africa: Guateng, Roodeport, Ruimsig sel_547, sel_548 barnardi Butterﬂy Reserve 211 Anyphops South Africa: Tsitsikamma National Park, sel_549 parvulus 78 km E Knysna 212 Anyphops South Africa: Grahmstown Municipal. Caravan sel_550 tugelanus Park 213 Anyphops South Africa: Eastern Cape, Kai Mouth, 58 km sel_551 stauntoni NE East London 214 Anyphops kraussi South Africa: Table Mountain National Park, sel_552 Newland’s Forest 215 Garcorops Madagascar: Reserve Nat. Integrale de Lokobe, sel_553 madagascarensis 3.61 km ESE Hellville 216 Hovops sp. Madagascar: Park National Montagne d’Ambre sel_275 montane rainforest 217 New genus sp. 1 Australia: Western Australia: Ravensthorpe, T80881, T80996 Australia Ravensthorpe Ranges South, WAM 10, 33°38′16.03″S, 120°10′46.01″E, 17.V.2007, under rock, MCLeng, ML Moir 218 New genus sp. 2 Australia: Western Australia: Two Peoples Bay T78485, T78489 Australia Nature Reserve, granite outcrop, site 6, 34°59′18″S, 118°44″E, 14.X.2006, under granite rock, ML Moir, JM Waldock 219 New genus sp. 3 Australia: Western Australia: Fiztgerald River T78500, T78498 Australia National Park, East Mt Barren, site 7, 33°55′28″S, 120°01′13″E, 25.XI.2006, under rock ML Moir, KEC Brennan