Biological Journal of the Linnean Society. 101, 288-322
Transcription
Biological Journal of the Linnean Society. 101, 288-322
Biological Journal of the Linnean Society, 2010, 101, 288–322. With 9 figures Molecular systematics of Selenops spiders (Araneae: Selenopidae) from North and Central America: implications for Caribbean biogeography SARAH C. CREWS1,2* and ROSEMARY G. GILLESPIE1 1 University of California Berkeley, Department of Environmental Sciences Policy and Management, 137 Mulford Hall, Berkeley, CA 94720-3114, USA 2 Berkeley City College, Department of Science and Biotechnology, 2050 Center Street, Berkeley, CA 94704, USA Received 16 February 2010; revised 3 May 2010; accepted for publication 3 May 2010 bij_1494 288..322 The Caribbean region includes a geologically complex mix of islands, which have served as a backdrop for some significant studies of biogeography, mostly with vertebrates. Here, we use the tropical/subtropical spider genus Selenops (Selenopidae) to obtain a finer resolution of the role of geology in shaping patterns of species diversity. We obtained a broad geographic sample from over 200 localities from both the islands and American mainland. DNA sequence data were generated for three mitochondrial genes and one nuclear gene for eleven outgroup taxa and nearly 60 selenopid species. Phylogenetic analysis of the data revealed several biogeographic patterns common to other lineages that have diversified in the region, the most significant being: (1) a distinct biogeographic break between Northern and Southern Lesser Antilles, although with a slight shift in the location of the disjunction; (2) diversification within the islands of Jamaica and Hispaniola; (3) higher diversity of species in the Greater Antilles relative to the Lesser Antilles. However, a strikingly unique pattern in Caribbean Selenops is that Cuban species are not basal in the Caribbean clade. Analyses to test competing hypotheses of vicariance and dispersal support colonization through GAARlandia, an Eocene–Oligocene land span extending from South America to the Greater Antilles, rather than over-water dispersal. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322. ADDITIONAL KEYWORDS: Bayesian phylogenetics – island biogeography – likelihood analysis of geographic range evolution. INTRODUCTION Remote islands form the basis for many biological studies because of their ability to act as a laboratory, with repeated sets of ecological and/or evolutionary experiments occurring within a circumscribed time frame (Cronk, 1997; Losos et al., 1998; Gillespie & Roderick, 2002; Gillespie, 2004; Ricklefs & Bermingham, 2008). While the Hawaiian Islands have served as a model system for processes of in situ diversification, the long history of studies on the biota of the Caribbean has provided some of the most important *Corresponding author. E-mail: [email protected] 288 insights into the complex interaction between colonization and diversification. In particular, the Caribbean has served as the setting for the establishment of most of the central tenets in the equilibrium theory of island biogeography (Munroe, 1948), the arguments being formulated independently by MacArthur and Wilson (1963, 1967) much later (Lomolino & Brown, 2009). More recent research on the islands has allowed an understanding of the interplay between ecological and evolutionary processes in shaping species diversity (Losos & Schluter, 2000; Schoener, Spiller & Losos, 2001). The primary feature of the Caribbean region that makes it particularly useful for examining the interaction between colonization and diversification is its © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS long and complex geological history. The Caribbean Basin began forming nearly 140 Mya. Islands in the basin consist of four different types: (1) land-bridge islands which were connected to each other or to the mainland at times of lower sea level; (2) continental islands which broke off from the mainland through tectonic displacement; (3) uplifted limestone islands; and (4) volcanic islands (MacPhee & Iturralde-Vinent, 2005; Robertson, 2009). Despite their limited isolation, the age and geologic complexity of the area have provided ‘well-defined paths of entry by which immigrants may reach’ the islands (Munroe, 1948). Moreover, the islands have served as the setting for adaptive radiation among lineages with limited dispersal ability, in particular lizards of the genus Anolis (Losos, 1992, 1994, 2009), frogs of the genus Eleutherodactylus (Hedges, 1989; Heinicke, Duellman & Hedges, 2007), some lineages of insects [e.g. beetles (Liebherr, 1988b), flies (Wilder & Hollocher, 2003)] and plants [e.g. lineages within the Melastomaceae (Michelangeli et al., 2008) and Asteraceae (FranciscoOrtega et al., 2008)]. Although studies to date have provided insights into how the individual lineages have colonized and subsequently diversified within the island system, notable controversies remain, including the source of colonists and the means by which they colonized the islands, biogeographic patterns within lineages and whether these patterns might be expected to be shared across multiple lineages (Guyer & Savage, 1986; Williams, 1989; Hedges, Hass & Maxon, 1992; Crother & Guyer, 1996; Hedges, 1996a,b). A particular focus of debate has been the role of vicariance vs. dispersal in shaping the Caribbean biota. Hedges and colleagues (Hedges et al., 1992; Hedges, 1996a,b; Hedges & Heinicke, 2007; Heinicke et al., 2007), working with herpetofauna, have suggested that the absence of lineages older than the break-up of the proto-Antilles (a contiguous land mass between North and South America) precludes a vicariant origin and they argue for the initial colonization of most taxa via over-water dispersal on flotsam. A similarly dominant role for dispersal has been suggested for multiple lineages of plants, such as Miconieae (Michelangeli et al., 2008). In contrast, other studies have suggested that vicariance has played a larger role than dispersal in the initial colonization of the Caribbean; for example, in lizards (Crother & Guyer, 1996; Iturralde-Vinent & MacPhee, 1999; MacPhee & Iturralde-Vinent, 2005) and some plants [e.g. Euphorbiaceae (van Ee et al., 2008)]. A related controversy focuses on the hypothesis of GAARlandia (Greater Antilles + Aves Ridge), first proposed by Iturralde-Vinent & MacPhee (1999), who used geological data and fossil evidence to demonstrate the likely existence of a land span connecting 289 South America to the Greater Antilles during the Eocene–Oligocene transition 35–33 Mya. The land span, although probably short-lived, may have provided an avenue for terrestrial organisms to colonize the Greater Antilles from South America. Among mammals, molecular phylogenies of primates and hystricognath rodents are consistent with the model, while sloths and insectivorans are not (Dávalos, 2004). The pattern in plants is similarly mixed. Molecular phylogenetic data from the genera Croton (Euphorbiaceae) (van Ee et al., 2008) and Styrax (Styracaceae) (Fritsch, 2003) show that the timing of divergence of lineages is consistent with the GAARlandia hypothesis. However, similar data from endemic legume radiations in the Greater Antilles, although initially thought to indicate ancient splitting between lineages consistent with the GAARlandia hypothesis (Lavin et al., 2001), show more recent diversification (Lavin & Beyra-Matos, 2008), which is likely to hold also for lineages of Asteraceae (Francisco-Ortega et al., 2008). Clearly, the timing and frequency of dispersal and vicariance, and the interplay between the two, varies across biotic assemblages. The challenge, then, is to understand the circumstances dictating the relative roles of each and how they interact. Arthropods, because they can provide a fine-scale resolution of biogeographic patterns (Ferrier et al, 2004), are ideal candidates for elucidating the nature of these relationships. Although the biogeography of terrestrial invertebrates in the Caribbean has been examined in some detail (see Liebherr, 1988a and chapters therein), few recent studies have been attempted, with little molecular information on the timing and nature of the interplay between colonization and diversification. However, there are some notable exceptions (Davies & Bermingham, 2002; Wilder & Hollocher, 2003; Brisson, Wilder & Hollocher, 2006). In particular, recent studies on spiders (Sicariidae: Loxosceles) support the GAARlandia hypothesis in the colonization of the lineage of North from South America (Binford et al., 2008), while crickets show a more mixed pattern of both vicariance and dispersal, coupled with intra-island diversification (Oneal, 2009). In this study, we combine molecular and morphological methods to examine the phylogenetic relationships and biogeographic history of the cursorial and dispersal-limited spider genus Selenops (Araneae: Selenopidae) in the Caribbean. These primarily tropical and subtropical spiders (Muma, 1953; Corronca, 1998; Alayón, 2005) are distinctive in that they are extremely dorsoventrally flattened and exceedingly fast. They are found in a variety of habitats and microhabitats (Crews, Wienskoski & Gillespie, 2008). Although the genera and species groups have © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 290 S. C. CREWS and R. G. GILLESPIE Figure 1. Map of the study area. The Americas; the boxed region shows the primary study area. undergone several revisions (Muma, 1953; Corronca, 1998; Alayón, 2005), there is no phylogenetic framework for the family or for any of the component genera. They were chosen for the current study because of their high diversity and abundance in the Caribbean, where they occur in both the Greater and Lesser Antilles, as well as on the adjacent mainland (southern North America and throughout South America) (Muma, 1953; Crews, 2005; Crews et al., 2008, 2009;) (Figs 1, 2). Accordingly, they provide the potential to reveal fine-scale biogeographic patterns across the islands of the Caribbean. The current study uses the genus to infer the relative importance of the following two processes in dictating the biogeographic history of the lineage in the Caribbean: (1) the frequency of colonization to the Caribbean region from a mainland source and between islands within the Caribbean; and (2) whether within-island diversification has occurred through a single radiation or through dispersal and multiple radiations. We also test the hypotheses of dispersal and vicariance in the framework of the GAARlandia hypothesis and likelihood biogeographic analysis. MATERIAL AND METHODS TAXON SAMPLING A comprehensive geographic sample of the genus was obtained from the Caribbean region, including most islands and several sites throughout Mexico, Central America and the South American mainland (see also Supporting Information, Figs S1, S2). Political reasons prohibited us from obtaining permits to collect several endemic species from Cuba and the single species from Navassa Island. The implications for these omissions are discussed at the end of this paper. Outgroups included other genera in the family Selenopidae from all major geographic locations where the family is found, in particular the type of the genus (Selenops radiatus Latreille) from Africa, Selenops bursarius Karsch from Japan, Selenops montigenus Simon from Nepal/India and representatives of the three other genera described from Africa (six species of Anyphops, one species of Hovops and one species of Garcorops), as well as an undescribed Australian genus. Chosen representatives outside of the family include a broad sample of eight genera from two families, the © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 291 Figure 2. Map of the study area showing the number of localities per region (the first number), the number of total specimens per region (the second number) and the number of species collected out of possible known species per region (the third and fourth numbers, respectively). For more detailed collection information, see the Appendix and Supporting Information (Figs S1, S2). Table 1. Genetic loci and primer pairs used for PCR amplification CO1 LCO1_1490 C1N_2568 5′-GGTCAACAAATCATAAAGATATTG-3′ – Folmer et al. 1994 5′-GCTACAACAATAATAAGTATCATG-3′ – Hedin & Maddison, 2001 16S–ND1 12350mod 13398 Histone H3aF H3aR 5′-TTDGNTACCAAGCAGACVGC-3′ – this study 5′-CGCCTGTTTAACAAAAACAT-3′ – Simon et al. 1994 5′-ATGGCTCGTACCAAGCAGACVGC-3′ – Colgan et al. 1998 5′-ATATCCTTRGGCATRATRGTGAC-3′ – Colgan et al. 1998 CO1, cytochrome oxidase I; 16S, ribosomal DNA; ND1, NADH dehydrogenase I. Sparassidae and the Ctenidae, and were based on unpublished data (M. Ramirez, pers. comm.) (Table 4). The genus Selenops has also been found in Dominican amber and one of these specimens is an adult male, described by Schawaller (1984) as Selenops beynai. The specimen was scanned using X-ray computed tomography, as in Penney et al. (2007); however, the poor preservation of the genitalia prohibited even tentative incorporation into the phylogenetic framework. MOLECULAR METHODS Four gene fragments were amplified – three mitochondrial [cytochrome oxidase I (CO1), 16S ribosomal DNA (16S) and the intervening leucine tRNA and NADH dehydrogenase I (ND1)] and one nuclear [histone 3a (H3)] (see Table 1). The respective lengths of the amplification products were ~850, ~800 and ~330 base pairs (bp). DNA sequences can be found on GenBank (GU109549–GU110746, HM575429–HM576623, and HM576658). These markers were chosen as they have become a standard in spider molecular phylogenetics, with several primers available for each gene (Hedin & Maddison, 2001; Arnedo et al., 2004; Crews & Hedin, 2006). Also, the chosen genes evolve at different rates and contain both protein and non-protein coding regions. DNA was extracted from a portion of a leg using a Qiagen DNeasy Tissue Kit following the manufacturer’s protocol. Each new specimen used in this study was given an individual number (e.g. sel_001) and has been deposited in the Essig Museum of Entomology at the University of California, Berkeley and the California Academy of Sciences. Remaining genomic DNA is stored at -80 °C in the Gillespie and Roderick Laboratories, University of California, © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 292 S. C. CREWS and R. G. GILLESPIE Berkeley. Primer pairs used are given in Table 1. In some cases, primarily with outgroup taxa, amplification was difficult and, in such instances, the Epicentre FailSafe PCR kit was used. In the majority of cases, sequence data were obtained for all gene fragments for multiple representatives of each species. In one situation with the species Selenops insularis Keyserling, there was evidence for multiple copies of H3a in some specimens, thus these sequences were not analysed for these individuals. PHYLOGENETIC METHODS Alignments of the protein-coding loci CO1, ND1 and H3a were performed manually using Mesquite ver. 2.5 (Maddison & Maddison, 2008), with the aminoacid translations used as a guide. The 16S data were aligned using secondary structure based on the model from Masta (2000). While there were some length differences between taxa, alignment was straightforward. Data were partitioned by codon position for protein coding genes, by stems and loops for ribosomal DNA and by gene for both the maximum likelihood and Bayesian analyses to improve the fit of the substitution model to the data (Nylander et al., 2004; Brandley, Schmitz & Reeder, 2005). The doublet model of nucleotide substitution was used for the stem regions of 16S and the tRNA (Schöniger & von Haeseler, 1994; Kjer, 2004). Maximum likelihood analyses were performed with RAxML ver. 7.0.4 (Stamatakis, 2006) and Bayesian analyses were performed using MrBayes ver. 3.1.2 (Huelsenbeck & Ronquist, 2001; Ronquist & Huelsenbeck, 2003; Altekar et al., 2004). RAxML is able to analyse partitioned data, but only under the generalised time reversible (GTR) model, thus, while the same partitioning regime was used in both Bayesian and likelihood analyses, this was the model that was used in the maximum likelihood analyses. To determine the models for each partition in the Bayesian analysis, MrModeltest ver. 2.3 (Nylander, 2004) was used. Models were chosen using the Akaike information criterion (AIC; Akaike, 1973; see Posada & Buckley, 2004) and are listed in Table 2. LIKELIHOOD ANALYSES RAxML maximum likelihood analyses were conducted in a variety of ways following the suggestions of the author (Stamatakis, 2006). First, one analysis was conducted which included 893 terminals after identical haplotypes were removed. However, to ease the computational strain for more intensive analyses, terminals that were ⱕ 0.3% different were removed from the analysis (sensu McGuire et al., 2007). This truncated data set contained 306 terminals. Table 2. Partitions used in likelihood and Bayesian analyses and selected models for each partition used in Bayesian analyses Partition Selected model 16S stems 16S half stems 16S loops Leucine tRNA stems Leucine tRNA loops ND1 postion 1 ND1 position 2 ND1 position 3 CO1 position 1 CO1 position 2 CO1 position 3 H3a position 1 H3a position 2 H3a position 3 GTR + G + doublet GTR + G GTR + I + G HKY + G + doublet HKY + G GTR + I + G GTR + I + G GTR + G GTR + I + G GTR + I + G GTR + G GTR + I JC + I K80 + G CO1, cytochrome oxidase I; 16S, ribosomal DNA; ND1, NADH dehydrogenase I. The RAxML manual suggests two ways to analyse data – the ‘fast and easy way’ and the ‘hard and slow way’ (Stamatakis, 2006). The fast and easy way was used to analyse the full data set because of its large size (~900 terminals and ~2000 bps). The hard and slow way was used to analyse the smaller data set and allows the program to find ‘good’ settings particular to an individual data set. The user’s manual was followed exactly for the analysis of the truncated data set using the ‘hard and slow’ method. First, five randomized maximum parsimony trees were generated and then each tree was inferred using a fixed setting of ten for the initial rearrangement. Next, this setting was automatically determined for the same five starting trees and whichever settings yielded the best likelihood scores were used for subsequent analyses. The second part of the ‘hard and slow’ method involves the number of rate categories. For this, the number of rate categories is increased by 15, from 10 to 55 for each of the five starting trees, using whichever setting worked best from the initial rearrangement analyses. Finally, ten analyses were run using the best settings from the above experiments and bootstraps from 500 iterations were then added to the tree with the best likelihood. For all RAxML analyses, the rapid bootstrap algorithm was used (Stamatakis, Hoover & Rougemong, 2008). BAYESIAN ANALYSES Several analyses were run using MrBayes-mpi on the cluster at the Museum of Vertebrate Zoology, University of California, Berkeley, as well as on the CIPRES © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS cluster at the San Diego Supercomputer Center. Despite using the truncated data set and running the jobs in parallel, analyses required months to near completion. Programs were run using the default settings for 40 million generations and, if convergence was not met, the generations were increased in increments up to 100 million, saving every 1000th tree. Convergence was assessed using Are We There Yet? (AWTY) (Wilgenbusch, Warren & Swofford, 2004; Nylander et al., 2008). LAGRANGE ANALYSES The program Lagrange (Ree et al., 2005; Ree & Smith, 2008) was used to test hypotheses of vicariance and dispersal. Lagrange uses likelihood models to test geographic range evolution and allows changes in dispersal and extinction parameters at different times in the past, allowing the incorporation of external information such as geological data and dispersal capabilities. For example, if a land mass did not exist at a particular time period, because it had not yet emerged or was inundated, the rate of dispersal to the land mass would be 0 during this time and could increase during the time period(s) the land mass was available for colonization. In an area as geologically complex as the Caribbean, there are nearly endless ways to parameterize the models, but simplicity was maintained throughout each analysis. Lagrange requires a tree and a matrix of range data for the included taxa. We analysed a truncated data set, selecting one specimen from each species, along with the outgroups, using a partitioned RAxML search for the best tree. We then pruned the outgroups before conducting the Lagrange analyses to make the computational load smaller, and because the focus of the questions concerns only the ingroup. We ensured the tree had the same basic structure as trees from the more complete analyses and that all relationships supported in those analyses also appeared in this tree. We divided the range of the Selenopids in North and Central America into five areas: C (Central America and Mexico), S (South America), G (Greater Antilles), N (Northern Lesser Antilles), A (Southern Lesser Antilles). Although certain parts of these regions were not available for colonization throughout particular time periods (i.e. some of the Greater Antilles have been emergent longer than others, etc.), we simply used the maximum times from their first appearance. We set the age of the root node of the tree to 130 Myr, as it is assumed a split between the ingroup, i.e. American selenopids, and the outgroup, i.e. African selenopids, was caused by the separation of Africa from South America. Dating vicariance events by the initiation of mid-ocean ridge spreading 293 Table 3. Correspondence between time slices and geographical ranges defined for models used in Lagrange analyses Time slice Land availability 3.0 5.0 Closing of the Isthmus of Panamá Most recent appearance of Northern Lesser Antilles Most recent appearance of Southern Lesser Antilles Disappearance of GAARlandia Appearance of GAARlandia From 55–50 Mya, a part of Jamaica was connected to Central America via the Nicaraguan Rise Time after which land was available in the Greater Antilles region Age of root node, corresponds to separation of Africa and South America 12.0 33.0 35.0 50.0 55.0 130.0 is problematic, in that in some cases this upper bound is too old as a result of chance transoceanic interchange after actual separation. However, the existence of distinct clades of Selenops on the different continents would argue for little genetic exchange between the continental land masses (Smith & Peterson, 2002) and would therefore indicate that it is indeed appropriate to use the separation of Africa and America to date Selenops. Throughout all analyses, we focused on six time periods which correspond to the availability of land for colonization (Table 3). The following three analyses consisted of two models each, one representing each of three scenarios with, and without, GAARlandia. The three scenarios were: (1) a dispersal-based scenario where distance between land masses determines the probability of colonization; (2) a dispersal-based scenario in which the ability to colonize an available land mass is not dependent on distance, thus the colonization of any one land mass from another is equiprobable; (3) a vicariance-based scenario, with little to no over-water dispersal. This means that colonization of one area from another could occur only through connections of one land mass to another. In some cases, certain areas were never connected to other land masses, such as the Lesser Antilles. In this case, the probability of dispersal is not set to zero, but rather a very low probability, as the presence of the spiders indicates colonization at some point in the past. It is possible to set different dispersal probabilities for each direction, so that the probability of moving from one region to another can be lower or higher than in the opposite direction. However, to maintain © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 294 S. C. CREWS and R. G. GILLESPIE simplicity, bidirectional probabilities were set as equal. Within each model, the only parameter changed between the two analyses was the probability of colonization with and without the presence of GAARlandia. The maximum range size was set to two areas and the areas G, N and A were excluded from the root (> 130 Myr) as they were not available for colonization at this time (Table 3). RESULTS SAMPLING We obtained over 1000 specimens from over 200 localities within the area of primary focus for this study. In total, we have 29 out of 41 Caribbean island species, half of the known Mexican and Central American species and one fifth of the described South American species (Appendix). PHYLOGENETIC ANALYSES Likelihood analysis The tree from the analysis of the full data set is shown in Figure 3 and has a likelihood score of -61 544.60. Nodes with bootstrap (BS) values ⱖ 70% are considered to be supported. There is no support (BS < 70%) for many basal nodes. Further discussion of the results from this tree is given below where compared with trees from the other analyses. In the analysis of the truncated data set, the best likelihood score came from the trial with a fixed setting of 10 for the initial rearrangement, rather than the automatic setting (Table 5). The best likelihood from the experiment to determine a good setting for the number of rate categories occurred when this setting was at 25 (Table 6). Thus, the initial rearrangement setting was fixed at 10 (-i 10) and the number of rate categories was set to 25 (-c 25). The best overall likelihood with these settings from the MultTrees analysis came from the second run (Table 7) and the results are shown in Figure 4. The overall structure is similar to the tree obtained from the analysis of the full data set, in which many basal nodes are not supported, while nodes above these are. This tree is discussed in more detail below. Bayesian analysis The analyses were run for 64 million generations (the maximum possible given limits of storage space for our output files). According to the cumulative plot from AWTY (Wilgenbusch et al., 2004; Nylander et al., 2008), the run reached convergence near 55 million generations. Because convergence was only reached very late in the analysis, the first 90% of trees were eliminated as burn-in, leaving ~12 000 trees from which to compute a consensus. This tree is shown in Figure 5 and nodes with posterior probability values ⱖ 0.95 are considered to be supported. The branch lengths are longer than in the likelihood analyses and, while a few more basal nodes are supported than in the likelihood analyses, the overall pattern is the same. The similarities and differences among all three trees are discussed below. Comparison of trees All three trees are very similar with many of the minor differences not supported. The remainder of the basal nodes occurring below the Selenopids of North and Central America is only supported as monophyletic in the Bayesian analysis. The focal taxa of the study, the Selenopids of North and Central America, are monophyletic and further subdivided into a well-supported strictly Caribbean clade (Fig. 6, clade A) and the remaining taxa, supported as a clade in the Bayesian tree only; (Fig. 6, clade B), including taxa from the south-western USA, Mexico, Central America, the Southern Lesser Antilles (SLA) and South America. Within this clade B, although basal relationships are not supported, all analyses support a southern Caribbean basin clade (Fig. 6, clade C) consisting of taxa from Aruba, Bonaire, Curaçao, Trinidad and Tobago. Selenops n. sp. 5 from Aruba is always sister to Selenops curazao from Bonaire and Curaçao and, this clade (Fig. 6, clade D), is always sister to Selenops willinki from northern South America and Tobago + S. geraldinae from Trinidad (Fig. 6, clade E). Also within clade B, another wellsupported clade in all analyses consists of Selenops banksi, found in Panama and South America, and Selenops micropalpus, found in the Southern Lesser Antilles from Dominica to St Vincent and the Grenadines (Fig. 6, clade F). There is support for a sister group relationship between the South American taxa + the Central and North American taxa in the Bayesian tree only. Both the Bayesian analysis and the likelihood analysis of the truncated data set support a Central American + North American clade (Fig. 6, clade G), as well as one between the widespread Selenops mexicanus, Selenops gracilis and a new species found only in Mexico (Fig. 6, clade H). In the Bayesian tree, S. mexicanus is paraphyletic. There is little support for any other relationships in clade B, other than the species from the Selenops debilis group of the southwestern USA and Northern Mexico (Fig. 6, clade I). The Caribbean clade (Fig. 6, clade A) consists only of taxa from Caribbean islands and is strongly supported in all analyses, but, again, with little support for basal nodes, the exception being the widespread Selenops lindborgi and its sister species, S. n. sp. 3, which are supported as sister to the rest of the Caribbean taxa (Fig. 6, clade J). © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 295 Figure 3. Likelihood tree resulting from the RAxML analysis of the full data set. The map above the tree depicts the Caribbean islands and the colours correspond to branches in the tree and indicate on which island the species is found. Multiple colours along a branch indicate that the species is found on multiple islands. A branch outlined in black indicates the species is found in Cuba. Selenops radiatus (highlighted in blue) is the type of the genus. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 296 S. C. CREWS and R. G. GILLESPIE Table 4. Outgroup taxa used to root trees, collection localities, voucher numbers and location of vouchers Family Genus and species Locality Voucher number and locality Ctenidae Ctenidae Ctenidae Madagascar, Ranomafana French Guiana, Tresor Nature Reserve French Guiana, Emerald Jungle Village CASENT9024024 – CAS CASENT9021738 – CAS CASENT9021735 – CAS Ctenidae Sparassidae Sparassidae Vulsor sp. Phoneutria fera Cupiennius ca. granadensis Acanthoctenus sp. Olios sp. 1 Olios sp. 2 ARAMR000556 – MACN In author’s personal collection In author’s personal collection Sparassidae Sparassidae Sparassidae Sparassidae Sparassidae Polybetes pythagoricus Heteropoda sp. Heteropoda sp. Damastes sp. 1 Damastes sp. 2 Argentina, Parque Nacional Cope, límte NE USA, California, Esparto México, Baja California, north of Guerrero Negro Argentina, Buenos Aires Prov., José Mármol Nepal, near Sauraha Tanzania Madagascar, Toliara Madagascar, Ambohitantely Ar 10876 – MACN In author’s personal collection In author’s personal collection sel_554 – CAS CASENT9015896 – CAS CAS, California Academy of Sciences; MACN, Museo Argentino de Ciencias Naturales. Table 5. Likelihoods from the ‘hard and slow’ RAxML analyses to determine the best initial rearrangement setting for the data -ln(L) for initial rearrangement setting fixed at 10 -ln(L) for automatic initial rearrangement setting 49 876.238442* 49 881.090042 49 880.995332 49 881.483263 49 879.897717 49 880.146295 49 877.349169 49 893.148152 49 879.924317 49 897.163460 *The best score is denoted. A relationship consisting of the four Jamaican species, four species endemic to Hispaniola + S. insularis, from throughout the Greater Antilles, is represented in all three analyses, although not supported in the full data set (Fig. 6, clade K). The Jamaican species are monophyletic, with well-supported interrelationships in all analyses (Fig. 6, clade L). The sister clade, consisting primarily of Hispaniolan endemics (three of which are undescribed), is also well supported (Fig. 6, clade M). However, Hispaniola has several species outside this clade. Nodes on the branches subtending other major Caribbean lineages (clades N, O, P and Q in Fig. 6) are unsupported, although many sister group relationships and one small subclade consisting of three undescribed species from Hispaniola and one from the Turks and Caicos Islands (Fig. 6, clade N) are supported in all analyses. Lagrange analyses The results of the Lagrange analyses are given in Table 8. Shown in Figures 7–9 are the maximum like- lihood reconstructions of range evolution under each of the six models. The best likelihood score overall (-121.90) was from model 3B, the vicariance-based model that includes GAARlandia (Fig. 9B). In models 2A–3B, the best likelihood scores were produced from those that included GAARlandia. The first two analyses (using models 1A and 1B), which take distance between islands into account, produced very similar likelihood scores and maximum likelihood reconstructions, although the model without GAARlandia had an insignificantly greater likelihood score. However, in the other two analyses, the differences in likelihood scores were significant and, in the analyses modelled with no GAARlandia, there was much more uncertainty in the reconstructions (Figs 7–9 – grey branches indicate that alternative reconstructions fall within two log-likelihood units of the scenario that is depicted). Likelihood ratio tests were used to compare nested models and, when scenarios were not nested (e.g. – scenario 2A and scenario 3A), the highest likelihood score is taken as the best. DISCUSSION Unique and shared biogeographic patterns are summarized in Table 9. SOUTHERN CARIBBEAN BASIN Members of the well-supported Southern Caribbean Basin clade (Aruba, Bonaire, Curaçao, Trinidad and Tobago, Fig. 6, clade C) are never found within the larger well-supported Caribbean clade (Fig. 6, clade A). Geological data often suggest a relationship between these southern islands, known as the Aruba– Tobago Belt (Iturralde-Vinent & MacPhee, 1999) and © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 297 Table 6. Likelihoods from the ‘hard and slow’ RAxML analyses to determine the best setting for the number of rate categories for the data Rate categories = 10 Rate categories = 25 Rate categories = 40 Rate categories = 55 Starting tree -ln(L) -ln(L) -ln(L) -ln(L) 1 2 3 4 5 49 876.825172 49 909.100516 49 884.108152 49 878.378649 49 885.284097 49 880.146295 49 877.349169* 49 893.148152 49 879.924317 49 897.163460 49 878.266732 49 877.528777 49 882.377547 49 884.766631 49 877.756447 49 881.349421 49 880.668732 49 885.827844 49 884.610743 49 891.633712 *The best score is denoted. Table 7. Likelihoods from the MultTrees analyses with the initial rearrangement setting at 10 and the number of rate categories set to 25 Tree -ln(L) 1 2 3 4 5 6 7 8 9 10 49 890.352743 49 875.885667* 49 877.963666 49 876.018067 49 884.591963 49 881.261966 49 879.547932 49 883.400698 49 897.114073 49 879.548262 *The best score is denoted. indeed the affinities are not surprising given the proximity of the islands to each other and to the South American continent. The amphibian and reptile assemblages on each of these islands are largely continental and also distinct from the primary Caribbean elements (Hedges, 2006). In Selenops, this Southern Caribbean clade is apparently not closely related to other Caribbean taxa, a pattern found in many other groups, including mammals (Dávalos, 2004) and plants [orchids (TrejoTorres & Ackerman, 2001)]. However, a contrasting pattern has been found in Anolis lizards in which the Southern Caribbean Basin taxa show stronger affinities with the Antilles (Jackman et al., 1999; Creer et al., 2001); these affinities are hypothesized to have arisen as a result of the Lesser Antilles being much further west, and thus closer to Bonaire, in the past (Creer et al., 2001). NORTHERN VS. SOUTHERN LESSER ANTILLES A pattern that the Selenops spiders share with several insects (Wilder & Hollocher, 2003), Anolis lizards (Gorman & Atkins, 1969; Jackman et al., 1999; 2002; Creer et al., 2001; Schneider, Losos & de Queiroz, 2001) and Eleutherodactylus frogs (Kaiser, Sharbel & Green, 1994), is that species in the Northern Lesser Antilles are only distantly related to species in the Southern Lesser Antilles. The species S. n. sp. 7 is found in the Northern Lesser Antilles from Les Saintes northward to Montserrat and Antigua, while the species occurring in the Southern Lesser Antilles, from Dominica south to St Vincent and the Grenadines (at least to Mayreau) is S. micropalpus. The northern species is nested well within the strictly Caribbean clade, while S. micropalpus shares a relationship with S. banksi found from Panamá to Peru to Guyana. The precise location where northern and southern lineages are separated is variable, being slightly to the south in other lineages. For example, in Anolis, it is between Dominica and Martinique (Losos & Thorpe, 2004); among Lygaeid bugs (Slater, 1988), carabid beetles (Liebherr, 1988b), butterflies (Davies & Bermingham, 2002), Eleutherodactylus frogs (Kaiser et al., 1994) and populations of the bananaquit (Seutin et al., 1994), it is between St Vincent and St Lucia. Differences in the location of the boundary between northern and southern lineages may occur as a result of the timing of colonization of the different groups, which is likely related to the timing of emergence of the individual islands. Interestingly, anoles from the Southern Lesser Antilles, like the spiders, show affinities with Central and South American anoles (Jackman et al., 1999; Creer et al., 2001). ORIGIN OF TAXA The basal taxa for the larger Caribbean clade (Fig. 6, clade A) are the widely distributed S. lindborgi (Puerto Rico, Culebra, Vieques, all of the Virgin Islands, St Kitts, Nevis, eastern Hispaniola and Great Inagua in the Bahamas, see also Supporting Information, Fig. S1E–G) and the very narrowly distributed © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 298 S. C. CREWS and R. G. GILLESPIE Figure 4. Likelihood tree resulting from the RAxML analysis of the truncated data set. The map above the tree in Figure 3 depicts the Caribbean islands and the colours correspond to branches in the tree and indicate on which island the species is found. Multiple colours along a branch indicate that the species is found on multiple islands. A branch outlined in black indicates the species is found in Cuba. Selenops radiatus (highlighted in blue) is the type of the genus. (Isla Mona and Puerto Rico, see also Supporting Information, Fig. S1F) S. n. sp. 3 (Fig. 6 clade J). A similar pattern is found among Anolis, with Puerto Rico endemic Anolis occultus also basal (Jackman et al., 1999). Likewise, the most basal iguana of the genus Cyclura is also located on the Puerto Rican bank (Malone et al., 2000). This pattern, which indicates a common origin of Caribbean diversity for these groups, is in contrast to data from geckos, frogs, colubrid snakes and butterflies, which suggest Hispaniola as a centre of diversity (Liebherr, 1988a, and references therein). © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 299 Figure 5. Tree resulting from the Bayesian analysis of the truncated data set. The map above the tree in Figure 3 depicts the Caribbean islands and the colours correspond to branches in the tree and indicate on which island the species is found. Multiple colours along a branch indicate that the species is found on multiple islands. A branch outlined in black indicates the species is found in Cuba. Selenops radiatus (highlighted in blue) is the type of the genus. UNIQUE BIOGEOGRAPHY OF JAMAICA Jamaica is one of the oldest islands of the Greater Antilles, with areas that may have had some parts continuously above sea level for many millions of years longer than other islands (Iturralde-Vinent & MacPhee, 1999; Iturralde-Vinent & Gahagan, 2002). It is also more isolated than other islands as its last probable connection with a land mass was likely with Central America through the Nicaraguan Rise 55 Mya. Our data reflect this isolated history, as Jamaican species of Selenops form a monophyletic group of endemics (Fig. 6, clade L). Monophyly of Jamaican taxa is also present in anoles (Jackman et al., 1999; Nicholson et al., 2005) and Eleutherodactylus frogs (Hedges, 1996a,b). However, affinities of the Jamaican clade differ between spider and vertebrate groups: The Jamaican clade of Selenops is supported in the Bayesian and truncated likelihood analyses as being sister to a clade of primarily Hispaniolan species (Fig. 6, clade K). In contrast, the Jamaican clade of Eleutherodactylus frogs is most © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 300 S. C. CREWS and R. G. GILLESPIE Figure 6. Bayesian tree with species symbols and asterisks indicative of support removed for clarity. The outgroup taxa have also been removed. Letters on the nodes indicate clades discussed in the text and in Table 9. closely related to species from Cuba, while the Jamaican lineages of Anolis lizards (Nicholson et al., 2005) and short-faced bats (Dávalos, 2007) are sister to clades from the mainland. Overall, Jamaica’s history has been quite different from that of the other Greater Antillean islands and its fauna may have accumulated via dispersal and in situ speciation rather than vicariance (Buskirk, 1985; Crother & Guyer, 1996). BIOGEOGRAPHICALLY DERIVED POSITION OF CUBA Cuba has often been depicted as a basal locality in area cladograms (Buskirk, 1985; Crother & Guyer, 1996). In contrast, although not always supported, Selenops species from Cuba appear not to be basal, at least based on morphology and our limited molecular sampling. Only one species (Selenops aissus – collected from the Bahamas, but that also occurs in © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 301 Table 8. Results of the Lagrange analyses for each of the six proposed models Model -ln(L) Dispersal Extinction 1A 1B 2A 2B 3A 3B 135.5 135.9 144.3 128.1* 260.8 121.9* 0.01634 0.01637 0.02604 0.02137 0.4647 0.1189 0.0082 0.008387 0.008411 0.008094 0.006629 0.008262 – – – – – – dispersal where distance is important; without GAARlandia dispersal where distance is important; with GAARlandia dispersal where distance is not important; without GAARlandia dispersal where distance is not important; with GAARlandia little to no over-water dispersal; without GAARlandia little to no over-water dispersal; with GAARlandia Model 1 is a dispersal-based model in which the distance between land masses is considered. Model 2 is a dispersal-based model in which the distance between land masses is not taken into account. Model 3 is a vicariance-based model in which dispersal probabilities are very low if dispersal must occur over water. The ‘A’ portion of each model was run without GAARlandia, while the ‘B’ portion was run with GAARlandia. The dispersal and extinction values are the maximum likelihoods estimates for the rate of each process and represent the mean number of events per unit of branch length. *Hypotheses that were statistically different from the null hypothesis of ‘no GAARlandia’ are marked. Figure 7. Maximum likelihood reconstruction of geographic range evolution under a dispersal-based model where distance between land masses is taken into account. Single-area ancestral ranges are shown at nodes. Grey branches indicate that alternative reconstructions fall within two log-likelihood units of the scenario that is depicted. Range transitions along branches show sequences of dispersal and extinction events. C, Central America; S, South America; G, Greater Antilles; N, Northern Lesser Antilles; A, Southern Lesser Antilles. (A) without GAARlandia; (B) with GAARlandia. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 302 S. C. CREWS and R. G. GILLESPIE Figure 8. Maximum likelihood reconstruction of geographic range evolution under a dispersal-based model where distance between land masses is ignored. Single-area ancestral ranges are shown at nodes. Grey branches indicate that alternative reconstructions fall within two log-likelihood units of the scenario that is depicted. Range transitions along branches show sequences of dispersal and extinction events. C, Central America; S, South America; G, Greater Antilles; N, Northern Lesser Antilles; A, Southern Lesser Antilles. (A) without GAARlandia; (B) with GAARlandia. Cuba) occurs at the base of an internal clade, while all other sampled species which occur in Cuba (although all but one – Selenops submaculosus – were collected from other islands) are nested high within the trees (S. submaculosus, Selenops simius, Selenops inuslaris) and it is inferred based on morphology (S. C. Crews & R. G. Gillespie, unpubl. data) that most of the Cuban endemics are closely related to S. simius and S. submaculosus (Fig. 6, clade O). GREATER ANTILLES and St Maarten and Anguilla are nested well within a clade of Hispaniolan animals (Fig. 6, clade O). This suggests that S. n. sp. 7 and S. n. sp. 8 colonized the Northern Lesser Antilles region from Hispaniola and thus support the Greater Antilles as a centre of species diversity via dispersal events in Anolis (Glor, Losos & Larson, 2005). Also, many species of Selenops in Hispaniola have very small ranges that mirror those of many endemic anoles from the Anolis cybotes group (Glor et al., 2003), indicating similar patterns of speciation between the two groups. AS A CENTRE OF SPECIES DIVERSITY There are two additional patterns that appear to be shared between Selenops and Anolis. In Selenops spiders, the species from the Northern Lesser Antilles SPECIES–AREA RELATIONSHIPS In many taxa as diverse as fungi (Lodge, Baroni & Cantrell, 2002), vertebrates (Ricklefs & Lovette, © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 303 Figure 9. Maximum likelihood reconstruction of geographic range evolution under a vicariance-based model where there are very low probabilities of over-water dispersal. Single-area ancestral ranges are shown at nodes. Grey branches indicate that alternative reconstructions fall within two log-likelihood units of the scenario that is depicted. Range transitions along branches show sequences of dispersal and extinction events. C, Central America; S, South America; G, Greater Antilles; N, Northern Lesser Antilles; A, Southern Lesser Antilles. (A) without GAARlandia; (B) with GAARlandia. 1999) and invertebrates (Nichols, 1988), the Greater Antilles harbour more species than the Lesser Antilles. This can be attributed largely to island area (MacArthur & Wilson, 1963) and associated habitat diversity and age (Losos, 1996; Ricklefs & Bermingham, 2002, 2008). In Selenops, the same pattern is found, with larger, older islands (Greater Antilles) having more species than smaller, younger, less habitat-diverse islands (Lesser Antilles). In the Greater Antilles there is often a pattern of number of species in Cuba > Hispaniola > Jamaica > Puerto Rico, based on island size. This pattern also prevails in Selenops. In this genus, 17 species occur in Cuba with 12 endemics (Alayón, 2005), while in Hispaniola there are at least 16 species with 11 endemics and, in Jamaica, at least five species are known, with four endemics. However, the Bahamas have no known endemic species of Selenops. HYPOTHESIS TESTING In the maximum likelihood analyses of range expansion, likelihood ratio tests of scores for the scenarios that include the existence of the GAARlandia land span are either equally probable or more favourable than those that do not. This does not mean that over-water dispersal has not occurred, but rather that land bridges hold a stronger signature on the phylogeny. These results contrast to those for mammals in which there was little to no support for a land span between the Greater Antilles and northern South America (Dávalos, 2004). Likewise, Hedges and © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 304 S. C. CREWS and R. G. GILLESPIE Table 9. Biogeographic patterns in Caribbean Selenops species Shared with other taxa? Pattern 1. Distinct, distantly related SLA and NLA clades Yes 2. Monophyly of Jamaican taxa 3. More than one colonization and diversification in Hispaniola 4. Patterns of endemism throughout the Caribbean (GA harbour more endemics than LA) 5. Endemic species in Hispaniola with distributions that overlap endemics of other taxa 6. Southern Netherlands Antilles form a clade with Trinidad and Tobago exlcusive of other Caribbean taxa 7. Location of the split between the NLA and SLA is between Dominica and Les Saintes, Guadeloupe 8. Cuban species are not basal in the Caribbean clade Yes Yes Clades that show particular patterns Which taxa? Lygaeid bugs, Carabid beetles, fruitflies, butterflies, Anolis, Eleutherodactylus, bananaquit Anolis, Eleutherodactylus Anolis F and O L A, J, M, N Yes Fungi, Anolis, birds, carabid beetles LA – F,O GA – J,K,L,M,N,P,Q Yes Anolis N, P, others in clade A No – C No – SLA – F NLA – O No – Q, S. aissus If a pattern is shared with other taxa, the taxa are noted. Clades referenced are those that display the patterns mentioned here and are depicted in Figure 6. GA, Greater Antilles; LA, Lesser Antilles; NLA, Northern Lesser Antilles, SLA, Southern Lesser Antilles. others (Hedges, Hass & Maxon, 1992; Hedges, 1996a,b; Hedges & Heinicke, 2007; Heinicke et al., 2007) found that molecular clock estimates of divergence times precluded a major role of land bridges in the origin of Caribbean herpetofauna. The suitability Dispersal–Extinction–Cladogenesis model of geographic range evolution used here, in which dispersal events cause range expansion, local extinction events cause range contraction and the probability of each kind of event is proportional to the branch length, has been questioned for island fauna, as terminal taxa may be restricted to single islands (Ree & Smith, 2008). However, in our models, islands were either grouped together or several species were spread across multiple islands and thus the model is reasonable in this particular case. CONCLUSIONS The current study provides a basis for biogeographic comparison across different lineages in the Caribbean. It is one of the most extensive data sets for Caribbean fauna and the most comprehensive molecular data set of any spider group. While inclusion of taxa currently missing from our analyses, and possibly the use of other markers, may help resolve basal relationships, it should also be noted that deep, short branches, such as those found here, may be very difficult if not impossible to resolve (Degnan & Salter, 2005; Kubatko & Degnan, 2007). The results reveal several patterns common to other disparate taxa, as well as many unique patterns which warrant further study. Moreover, the data set provides the groundwork for behavioural, ecological and population-level studies similar to lineages such as Anolis lizards (Losos, 2009) and passerine birds (Ricklefs & Bermingham, 2007). ACKNOWLEDGEMENTS We would like to acknowledge members of S.C.C.’s dissertation committee for their guidance: George Roderick, Jim McGuire and Charles Griswold. We would like to thank the following museums, curators and collection managers for specimen loans: American Museum of Natural History – Norman I. Platnick and Louis Sorkin; Museum of Comparative Zoology – Laura Leibensperger; California Academy of Sciences – Charles Griswold; National Museum of Natural History – Jonathan Coddington; British Museum of Natural History – Janet Beccaloni; Peabody Musem at Yale – Raymond Pupedis; Essig Museum of Ento- © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS mology – Cheryl Barr; Museo Nacional de Histora Natural, Santo Domingo – Sardis Medrano Cabral, Carmelo Nuñez. We would also like to thank Jim McGuire for use of the MVZ cluster and Mark Miller and Lucie Chan for use of the SGE cluster and the CIPRES portal and the San Diego Supercomputer Center. We are extremely grateful to Richard Ree for quickly responding and helping us to understand and implement Lagrange. We would also like to thank Matt Brandley for phylogenetic methodology discussions. We are grateful to all of the many people that aided us in obtaining permits and collecting: Kelvin Guerrero, Denia Veloz, Eladio Fernandez, Alberto Puente-Rolón, Beverly Mae Nisbeth, Adriel Thibou, Germain George, Brian Riggs, Brian Manco, Margaret Jones, Renata Platenberg, Chris Niebuhr, Abel Pérez-González, G. B. Edwards, Oscar Francke, Alejandro Mondragon, Mark da Silva, Facundo Franken, Roy Croes, Gijs Van Hoorn, Adolphe O. Debrot, Mark Vermeij, Fred the Abaco Caveman, Raveen Gibson, Daniel Palmer, Jim Starrett, Marshal Hedin, Nicole VanderSal, Sean Schoville, Luke Mahler, Uri García, Beto Mendoza, Adrian Nieto Montes de Oca, Rebecca Duncan, Pierre Paquin, Matthew Cottam, Jan den Dulk, Joey Slowik, Nicole Esteban, Arturo Herrera, Nancy Bottomley, Inilek Wilmot, Lauren Esposito, Stephen Touissant, Arlington James, Ferdinand Tripoli, Daniel Memia Zolo, Nouree-Yvon, Martín Ramírez, Mark Harvey, Volker Framenau, Jeremy Miller, Hannah Wood, Yuri Marusik, Caroline Chaboo, Cheryl Barr, Bill Shepard, Akio Tanikawa, C. J. Hayden, Aaron Abdel, Dan Warren. Finally, we would like to acknowledge Matjaz Kunter and an anonymous reviewer for their straightforward and constructive reviews of this paper. Funding was provided by the Schlinger Foundation, with additional support from the Margaret C. Walker Fund. REFERENCES Akaike H. 1973. Information theory and an extension of the maximum likelihood principle. In: Petrov BN, Caski F, eds. Second international symposium on information theory. Budapest, Hungary: Akademiai Kiado, 267–281. Alayón G. 2005. La familia Selenopidae (Arachnidae: Araneae) en Cuba. Solenodon 5: 10–52. Altekar G, Dwarkadas S, Huelsenbeck JP, Ronquist F. 2004. Parallel Metropolis coupled Markov chain Monte Carlo for Bayesian phylogenetic inference. Bioinformatics 20: 407–415. Arnedo MA, Coddington JA, Agnarsson I, Gillespie RG. 2004. From a comb to a tree: phylogenetic relationships of the comb-footed spiders (Araneae, Theridiidae) inferred from nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution 31: 225–245. 305 Binford GJ, Callahan MS, Bodner MR, Rynerson MR, Berea Nuñez P, Ellison CE, Duncan RP. 2008. Phylogenetic relationships of Loxosceles and Sicarius are consistent with Western Gondwanan vicariance. Molecular Phylogenetics and Evolution 49: 538–553. Brandley MC, Schmitz A, Reeder TW. 2005. Partitioned Bayesian analyses, partition choice, and the phylogenetic relationships of scincid lizards. Systematic Biology 54: 373– 390. Brisson JA, Wilder J, Hollocher H. 2006. Phylogenetic analysis of the cardini group of Drosophila with respect to changes in pigmentation. Evolution 60: 1228–1241. Buskirk RE. 1985. Zoogeographic patterns and tectonic history of Jamaica and the Northern Caribbean. Journal of Biogeography 12: 445–461. Colgan DJ, McLauchlan A, Wilson GDF, Livingston SP, Edgecombe GD, Macaranas J, Cassis G, Gray MR. 1998. Histone H3 and U2 snRNA DNA sequences and arthropod molecular evolution. Australian Journal of Zoology 46: 419–437. Corronca JA. 1998. A taxonomic revision of the afrotropical species of Selenops Latreille, 1819 (Araneae, Selenopidae). Zootaxa 107: 1–35. Creer DA, de Queiroz K, Jackman TR, Losos JB, Larson A. 2001. Systematics of the Anolis roquet series of the Southern Lesser Antilles. Journal of Herpetology. 35: 428– 441. Crews SC. 2005. Selenopidae. In: Ubick D, Paquin P, Cushing PE, Roth V, eds. Spiders of North America: an identification manual. Keene, NH: American Arachnological Society, 221. Crews SC, Hedin MC. 2006. Studies of morphological and molecular phylogenetic divergence in spiders (Araneae: Homalonychus) from the American southwest, including divergence along the Baja California Peninsula. Molecular Phylogenetics and Evolution 38: 470–487. Crews SC, Puente-Rolón AR, Rutstein E, Gillespie RG. 2009. A comparison of populations of island and adjacent mainland species of Caribbean Selenops (Araneae: Selenopidae) spiders. Molecular Phylogenetics and Evolution 54: 970–983. Crews SC, Wienskoski E, Gillespie RG. 2008. Life history of the spider Selenops occultus Mello-Leitão (Araeae, Selenopidae) from Brazil with notes on the natural history of the genus. Journal of Natural History 42-43: 2747–2761. Cronk QCB. 1997. Islands: stability, diversity, conservation. Biodiversity and Conservation 6: 447–493. Crother BI, Guyer C. 1996. Caribbean historical biogeography: was the dispersal vicariance debate eliminated by an extraterrestrial bolide? Herpetologica 52: 440–465. Dávalos LM. 2004. Phylogeny and biogeography of Caribbean mammals. Biological Journal of the Linnean Society 81: 373–394. Dávalos LM. 2007. Short-faced bats (Phyllostomidae: Stenodermatina): a Caribbean radiation of strict frugivores. Journal of Biogeography 34: 364–375. Davies N, Bermingham E. 2002. The historical biogeography of two Caribbean butterflies (Lepidoptera: Heliconiidae) © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 306 S. C. CREWS and R. G. GILLESPIE as inferred from genetic variation at multiple loci. Evolution 56: 573–589. Degnan JH, Salter LA. 2005. Gene tree distributions under the coalescent process. Evolution 59: 24–37. Ferrier S, Powell GVN, Richardson KS, Manion G, Overton JM, Allnutt TF, Cameron SE, Mantle K, Burgess ND, Faith DP, Lamoreux JF, Kier G, Hijmans RJ, Funk VA, Cassis GA, Fisher BL, Flemons P, Lees D, Lovett JC, Van Rompaey RSAR. 2004. Mapping more of terrestrial biodiversity for global conservation assessment. BioScience 54: 1101–1109. Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R. 1994. DNA primers for amplification of mitochondrial cytochrome C oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3: 294–299. Francisco-Ortega J, Ventosa I, Oviedo R, Jiménez F, Herrera P, Maunder M, Panero JL. 2008. Caribbean island Asteraceae: systematics, molecules, and conservation on a biodiversity hotspot. The Botanical Review 74: 112–131. Fritsch PW. 2003. Multiple geographic origins of Antillean Styrax. Systematic Botany 28: 421–430. Gillespie RG. 2004. Community assembly through adaptive radiation in Hawaiian spiders. Science 16: 356–359. Gillespie RG, Roderick GK. 2002. Arthropods on islands: colonization, speciation, and conservation. Annual Review of Entomology 47: 595–632. Glor RE, Kolbe JK, Powell R, Larson A, Losos JB. 2003. Phylogenetic analysis of ecological and morphological diversification in Hispaniolan trunk-ground anoles (Anolis cybotes group). Evolution 58: 2383–2397. Glor RE, Losos JB, Larson A. 2005. Out of Cuba: overwater dispersal and speciation among lizards in the Anolis caroliinensis subgroup. Moleular Ecology 14: 2419–2432. Gorman GC, Atkins L. 1969. The zoogeography of Lesser Antillean Anolis lizards – an analysis based upon chromosomes and lactic dehydrogenases. Bulletin of the Museum of Comparative Zoology 138: 554–579. Guyer C, Savage JM. 1986. Cladistic relationships among anoles (Sauria:Iguanidae). Systematic Zoology 35: 509–531. Hedges SB. 1989. Evolution and biogeography of West Indian frogs of the genus Eleutherodactylus: slow-evolving loci and the major groups. In: Woods CA, ed. Biogeography of the West Indies. Past, present and future. Gainesville, FL: Sandhill Crane Press, 305–370. Hedges SB. 1996a. The origin of West Indian amphibians and reptiles. In: Powell R, Henderson RW, eds. Contributions to West Indian herpetology: a tribute to Albert Schwartz. Ithaca, NY: Society For The Study of Amphibians and Reptiles, 95–128. Hedges SB. 1996b. Historical biogeography of West Indian vertebrates. Annual Review of Ecology and Systematics 27: 163–196. Hedges SB. 2006. Paleogeography of the Antilles and the origin of West Indian amphibians and reptiles. Applied Herpetology 3: 281–292. Hedges SB, Hass CA, Maxon LR. 1992. Caribbean biogeography: molecular evidence for dispersal in West Indian terrestrial vertebrates. Proceedings of the National Academy of Sciences of the United States of America 89: 1909–1913. Hedges SB, Heinicke MP. 2007. Molecular phylogeny and biogeography of West Indian frogs of the genus Leptodactylus. Molecular Phylogenetics and Evolution 44: 308–314. Hedin MC, Maddison WP. 2001. A combined molecular approach to phylogeny of the jumping spider subfamily Dendryphantinae (Araneae: Salticidae. Molecular Phylogenetics and Evolution 18: 386–403. Heinicke MP, Duellman WE, Hedges SB. 2007. Major Caribbean and Central American frog faunas originated by ancient oceanic dispersal. Proceedings of the National Academy of Sciences of the United Sates of America 104: 10092–10097. Huelsenbeck JP, Ronquist FR. 2001. MrBayes: Bayesian inference of phylogeny. Bioinformatics 17: 754–755. Iturralde-Vinent MA, Gahagan L. 2002. Late Eocene to Middle Miocene tectonic evolution of theCaribbean: Some principles and their implications for plate tectonic modeling. In: Jackson TA, ed. Caribbean geology into the third millennium: transactions of the Fifteenth Caribbean Geological Conference. Mona, Jamaica: University of the West Indies Press, 47–62. Iturralde-Vinent MA, MacPhee RDE. 1999. Paleogeography of the Caribbean region: implications for Cenzoic biogeography. Bulletin of the American Museum of Natural History 238: 1–95. Jackman TR, Irschick DI, de Queiroz K, Losos JB, Larson A. 2002. Molecular phylogenetic perspective on evolution of lizards of the Anolis grahami series. Journal of Experimental Zoology Part B 294: 1–16. Jackman TR, Larson A, de Queiroz K, Losos JB. 1999. Phylogenetic relationships and tempo of early diversification in Anolis lizards. Systematic Biology 48: 254–285. Kaiser H, Sharbel TF, Green DM. 1994. Systematics and biogeography of eastern Caribbean Eleutherodactylus (Anura: Leptodactylidae): evidence from allozymes. Amphibia. Reptilia 15: 375–394. Kjer KM. 2004. Aligned 18S and insect phylogeny. Systematic Biology 53: 506–514. Kubatko LS, Degnan JH. 2007. Inconsistency of phylogenetic estimates from concatenated data under coalescence. Systematic Biology 56: 17–24. Lavin M, Beyra-Matos A. 2008. The impact of ecology and biogeography on legume diversity and phylogeny in the Caribbean region: a new direction in historical biogeography. The Botanical Review 74: 178–196. Lavin M, Wojciechowski MF, Richman A, Sanderson M. 2001. Identifying Tertiary radiations of Fabaceae in the Greater Antilles: alternatives to cladistic vicariance analysis. International Journal of Plant Science 162: S53– S76. Liebherr JK. 1988a. Zoogeography of Caribbean insects. Ithaca, NY: Cornell University Press. 285p. Liebherr JK. 1988b. Biogeographic patterns of West Indian Platynus carabid beetles (Coleoptera). In: Liebherr JK, ed. Zoogeography of Caribbean insects. Ithaca, NY: Cornell University Press, 121–152. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS Lodge DJ, Baroni TJ, Cantrell SA. 2002. Basidiomycetes of the Greater Antilles Project. In: Watling R, Frankland JC, Ainsworth AM, Isaac S, Robinson CH, eds. Tropical mycology V. 1 macromycetes. Engham UK: CABI Publishing, 45–60. Lomolino MV, Brown JH. 2009. The reticulating phylogeny of island biogeography theory. Quarterly Review of Biology 84: 357–390. Losos JB. 1992. The evolution of convergent community structure in Caribbean Anolis communities. Systematic Biology. 41: 403–420. Losos JB. 1994. Historical contingency and lizard community ecology. In: Vitt LJ, Pianka ER, eds. Lizard ecology: historical and experimental perspectives. Princeton, NJ: Princeton University Press, 319–333. Losos JB. 1996. Ecological and evolutionary determinants of the species-area relation in Caribbean anoline lizards. Philosophical Transactions of the Royal Society B 351: 847–854. Losos JB. 2009. Lizards in an evolutionary tree: ecology and adaptive radiation of anoles. Berkeley, CA: University of California Press. Losos JB, Jackman TR, Larson A, de Queiroz K, Rodríguez-Schettino L. 1998. Historical contingency and determinism in replicated adaptive radiations of island lizards. Science 279: 2115–2118. Losos JB, Schluter D. 2000. Analysis of an evolutionary species-area relationship. Nature 408: 847–850. Losos JB, Thorpe RS. 2004. Evolutionary diversification of Anolis lizards: introduction. In: Dieckmann U, Metz HAJ, Doebeli M, Tautz D, eds. Adaptive speciation. Cambridge: Cambridge University Press, 343–344. MacArthur RH, Wilson EO. 1963. An equilibrium theory of insular zoogeography. Evolution 17: 373–387. MacArthur RH, Wilson EO. 1967. The theory of island biogeography. Princeton, NJ: Princeton University Press, 224p. MacPhee RDE, Iturralde-Vinent MA. 2005. The interpretation of Caribbean paleogeography: reply to Hedges. In: Alcover JA, Bover P, eds. Proceedings of the International Symposium ‘Insular Vertebrate Evolution: The Palaeontological Approach’. Monografies de la Societat d’Història Natural de les Balears 12. Maddison WP, Maddison DR. 2008. Mesquite: a modular system for evolutionary analysis. Version 2.5. Available at http://mesquiteproject.org Malone CL, Wheeler TC, Davis SK, Taylor JF. 2000. Biogeography and systematic of the Caribbean rock iguana (Cyclura): implications for conservation and insights into the biogeographic history of the West Indies. Molecular Phylogenetics and Evolution 17: 269–279. Masta SE. 2000. Mitochondrial sequence evolution in spiders: intraspecific variation in tRNAs lacking the TYC arm. Molecular Biology and Evolution 17: 1091–1100. McGuire JA, Linkem CW, Koo MS, Hutcison DW, Lappin AK, Orange DI, Lemos-Espinal J, Riddle BR, Jaeger JR. 2007. Mitochondrial introgression and incomplete lineage sorting through space and time: phylogenetics of crotophytid lizards. Evolution 61: 2879–2897. 307 Michelangeli FA, Judd WS, Penneys DS, Skean Jr. JD, Bécquer-Granados ER, Goldenberg R, Martin CV. 2008. Multiple events of dispersal and radiation of the tribe Miconieae (Melastomataceae) in the Caribbean. Botanical Review 74: 53–77. Muma MH. 1953. A study of the spider family Selenopidae in North and Central America and the West Indies. American Museum Novitates 1619: 1–55. Munroe EG. 1948. The geographical distribution of butterflies in the West Indies. PhD Dissertation. Ithaca, NY: Cornell University. Nichols SW. 1988. Kaleidoscopic biogeography of West Indian Scaritinae (Coleoptera: Carabidae). In: Liebherr JK, ed. Zoogeography of Caribbean insects. Ithaca, NY: Cornell University Press, 71–121. Nicholson KE, Glor RE, Kolbe JJ, Larson A, Hedges SB, Losos JB. 2005. Mainland colonization by island lizards. Journal of Biogeography 32: 929–938. Nylander JAA. 2004. MrModeltest v2. Program distributed by the author. Evolutionary Biology Centre, Uppsala University. Nylander JAA, Ronquist F, Huelsenbeck JP, NievesAldrey JL. 2004. Bayesian phylogenetic analysis of combined data. Systematic Biology 53: 47–67. Nylander JAA, Wilgenbusch JC, Warren DL, Swofford DL. 2008. AWTY (are we there yet?): a system for graphical exploration of MCMC convergence in Bayesian phylogenetics. Bioinformatics 24: 581–583. Oneal E. 2009. Biogeographic and evolutionary mechanisms driving diversification in Caribbean ground crickets (genus Amphiacusta). PhD dissertation. Ann Arbor, MI: University of Michigan. Penney D, Dierick M, Cnudde V, Masschaele B, Vlassenbroeck J, Van Hoorebeke L, Jacobs P. 2007. The first Micropholcommatidae (Araneae), imaged in Eocene Paris amber using X-ray computed tomography. Zootaxa 1623: 47–53. Posada D, Buckley TR. 2004. Advantages of AIC and Bayesian approaches over likelihood ratio tests for model selection in phylogenetics. Systematic Biology 53: 793–808. Ree RH, Moore BR, Webb CO, Donoghue MJ. 2005. A likelihood framework for inferring the evolution of geographic range on phylogenetic trees. Evolution 59: 2229– 2311. Ree RH, Smith SA. 2008. Maximum-likelihood inference of geographic range evolution by dispersal, local extinction, and cladogenesis. Systematic Biology 57: 4–414. Ricklefs RE, Bermingham E. 2002. The concept of the taxon cycle in biogeography. Global Ecology and Biogeography 11: 353–361. Ricklefs RE, Bermingham E. 2007. Evolutionary radiations of passerine birds in archipelagoes. The American Naturalist 169: 285–297. Ricklefs RE, Bermingham E. 2008. The West Indies as a laboratory of biogeography and evolution. Philosophical Transactions of the Royal Society B 363: 2393–2413. Ricklefs RE, Lovette IJ. 1999. The roles of island area per se and habitat diversity in the species–area relationships of © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 308 S. C. CREWS and R. G. GILLESPIE four Lesser Antillean faunal groups. Journal of Animal Ecology 68: 1142–1160. Robertson REA. 2009. Antilles geology. In: Gillespie RG, Clague DA, eds. Encyclopedia of islands. Berkeley, CA: University of California Press, 29–35. Ronquist F, Huelsenbeck JP. 2003. MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19: 1572–1574. Schawaller W. 1984. Die families Selenopidae in Dominikanischem Berrnstein (Arachnida, Araneae). Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie) 103: 1–8. Schneider CJ, Losos JB, de Queiroz K. 2001. Evolutionary relationships of the Anolis bimaculatus group from the Northern Lesser Antilles. Journal of Herpetology 35: 1–12. Schoener TW, Spiller DA, Losos JB. 2001. Natural restoration of the species–area relation for a lizard after a hurricane. Science 294: 1525–1528. Schöniger M, von Haeseler A. 1994. A stochastic model and the evolution of autocorrelated DNA sequences. Molecular Phylogenetics and Evolution 3: 240–247. Seutin G, Klein NK, Ricklefs RE, Bermingham E. 1994. Historical biogeography of the bananaquit (Coereba flaveola) in the Caribbean region: a mitochondrial DNA assessment. Evolution 48: 1041–1061. Simon C, Frati F, Bechenbach A, Crespi B, Liu H, Flook P. 1994. Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America 87: 651–701. Slater J. 1988. Zoogeography of West Indian Lygaeidae (Hemiptera). In: Liebherr JK, ed. Zoogeography of Caribbean insects. Ithaca, NY: Cornell University Press, 38–60. Smith AB, Peterson KJ. 2002. Dating the time of origin of major clades: molecular clocks and the fossil record. Annual Review of Earth and Planetary Sciences 30: 65–88. Stamatakis A. 2006. RAxML-VI-HPC: maximum likelihoodbased phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22: 2688–2690. Stamatakis A, Hoover P, Rougemong J. 2008. A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology 57: 758–771. Trejo-Torres JC, Ackerman D. 2001. Biogeography of the Antilles based on a parsimony analysis of orchid distributions. Journal of Biogeography 28: 775–794. van Ee B, Berry PE, Riina R, Gutiérrez Amaro JE. 2008. Molecular phylogenetics and biogeography of the Caribbean-centered Croton subgenus Moacroton (Euphorbiaceae s.s.). Botanical Review 74: 132–165. Wilder JA, Hollocher H. 2003. Recent radiation of endemic Caribbean Drosophila of the dunni subgroup inferred from multilocus DNA sequence variation. Evolution 57: 2566– 2579. Wilgenbusch JC, Warren DL, Swofford DL. 2004. AWTY: a system for graphical exploration of MCMC convergence in Bayesian phylogenetic inference. Available at http://king2. scs.fsu.edu/CEBProjects/awty/awty_start.php Williams EE. 1989. A critique of Guyer and Savage (1986): cladistic relationships among anoles (Sauria: Iguanidae): are the data available to reclassify the anoles? In: Woods CA, ed. Biogeography of the West Indies: past, present, and future. Gainesville, FL: Sandhill Crane Press, 433– 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. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 309 APPENDIX Collecting localities and voucher numbers of all animals used in this study. Locality numbers refer to numbers in the Supporting Information (Figs S1 and S2). Locality number Species Collection information Voucher numbers 1 Selenops n. sp. 5 sel_069, sel_070 2 Selenops n. sp. 5 3 Selenops n. sp. 5 4 Selenops curazao 5 Selenops curazao 6 Selenops curazao 7 Selenops curazao 8 Selenops curazao 9 Selenops willinki Aruba: Bringamosa, house of Roy Croes, 12°39.547′N, 69°58.077′W, 14.X.2004, SCC04_041 Aruba: Luela, shooting range, 12°29.023′N, 69°57.778′W, 16.X.2004, SCC04_044 Aruba:N. sp. 5 National Park, near hill Gran Tonel and Valley Rooi Coashati, 12°29.356′N, 69°55.461′W, 16.X.2004, SCC04_045 Curaçao: Carmabi Institute, 12°07.351′N, 68°58.132′W, 7.X.2004, SCC04_026 Curaçao: Girouette Plantation, E of Schottegat Harbour, house of A. DeBrot, 12°09′03.20″N, 68°54′56.35″W, 9.X.2004, SCC04_032 Bonaire: Sabadaco, near cave, 12°11.587′N, 68°17.765′W, 11.X.2004, SCC04_033 Bonaire: Altamira Ungu, 12°13.949′N, 68°20.703′W, 12.X.2004, SCC04_038 Bonaire: Nort di Saliña, Kaya Otomac, 12°10′55.92″N, 68°16′28.39″W, 11.X.2004, SCC04_039 Trinidad and Tobago: Little Tobago, 11°18′03.7″N, 60°30′11.1″W, 16.VII.2005, SCC05_041 10 Selenops geraldinae 11 Selenops geraldinae 12 Selenops geraldinae 13 Selenops geraldinae 14 Selenops geraldinae 15 Selenops hebraicus Selenops occultus 16 Trinidad and Tobago: St George Co., Point Gourde Road near Trinidad Military Base, near Chaguaramas off Chaguaramas Main Road, 10°40′47.1″N, 61°37′30.9″W, 11.VII.2005, SCC05_035 Trinidad and Tobago: Gaspar Grande Island south of Chaguaramas, trails around island, 10°39′46.5″N, 61°38′58.0″W, 12.VII.2005, SCC05_037 Trinidad and Tobago: Monos Island, South Sea, 10°40′54.2″N, 61°41′21.6″W, 13.VII.2005, SCC05_038 Trinidad and Tobago: Huevos Island, 10°41′28.3″N, 61°42′55.0″W, 13.VII.2005, SCC05_039 Trinidad and Tobago: Chacachacare Island, 10°41′24.2″N, 61°44′53.7″W, 13.VII.2005, SCC05_040 Argentina: Parque Nacional Chaco, Sendero peatonal. 25.II.2004. Brazil: São Paulo, Universidade São Paulo, 23°34′27.96″S, 46°40′21.29″W sel_072 sel_068 sel_047, sel_058, sel_217 sel_048, sel_049, sel_050, sel_051, sel_052 sel_053 sel_059, sel_060, sel_082 sel_054, sel_056, sel_057, sel_061, sel_062, sel_063, sel_064, sel_065 sel_230, sel_231, sel_232, sel_233, sel_234, sel_235, sel_236, sel_238, sel_240, sel_242, sel_243, sel_245, sel_251, sel_252, sel_253, sel_255, sel_258, sel_259, sel_261, sel_262, sel_263 sel, 218, sel_219, sel_220, sel_221, sel_222, sel_223 sel_224, sel_225, sel_226, sel_227, sel_228, sel_229, sel_237, sel_241, sel_257 sel_244, sel_247, sel_248, sel_249, sel_250 sel_239 sel_246 MACN-Ar#12782 sel_995 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 310 S. C. CREWS and R. G. GILLESPIE APPENDIX Continued Locality number Species Collection information Voucher numbers 17 Selenops occultus sel_284 18 Selenops occultus 18 Selenops melanurus Selenops micropalpus Brazil: Rio de Janeiro, Sierra da Carioca, 22°57′52.22″S, 43°16′31.34″W Brazil: Rio de Janeiro, Pao de Açucar, 22°55′22.05″S, 43°09′33.11″W Brazil: Rio de Janeiro, Pao de Açucar, 22°55′22.05″S, 43°09′33.11″W St Vincent and the Grenadines: Young Island, South of Villa Beach, 13°07.895′N, 61°12.142′W, 24.X.2004, SCC04_053 St Vincent and the Grenadines: King’s Hill Forest Reserve, 13°08.825′N, 61°10.021′W, 27.X.2004, SCC04_055 St Lucia: Vieux Fort, on hill above airport, 13°44′20.2″N, 60°56′40.8″W, 13.III.2007 St Lucia: Anse la Ray, Ti-Kaye, 13°55′29.6″N, 61°02′41.3″W, 13.III.2007, SCC07_046 St Lucia: Dennery: south of Dennery on east coast road, eastern nature trail, heritage tourism site, 13°53′50.8″N, 60°52′51.2″W, 13.III.2007, SCC07_047 St Lucia: Gros Islet, Pigeon Island, on top of hill, 14°05′31.3″N, 60°57′03.8″W, 12.III.2007, SCC07_044 St Lucia: Gros Islet, Beausejour, past cricket sponsor’s office, 14°04′43.1″N, 60°56′31.1″W, 12.III.2007, SCC07_045 Martinique: Le Diamant, Grand Anse du Diamant, off of road D37, 14°28′32.9″N, 61°02′13.4″W, 10.III.2007, SCC07_043 Martinique: La Caravelle Reserve Naturelle, trail to Pointe Caricoli, 14°46′09.3″N, 60°53′24.7″W, 8.III.2007, SCC07_041 Martinique: Anse Ceran, off of road D-10, 14°50′01.5″N, 61°13′24.7″W, 9.III.2007, SCC07_042 Dominica: Roseau: Botanical Park, top of hill near shrine, 15°17.998′N, 61°22.754′W, 1.XI.2004, SCC04_059 Dominica: Jimmit-Warner, on top of hill with cellular phone antenna, 15°22.690′N, 061°24.003′W, 2.XI.2004, SCC04_061 Dominica: Cabrits National Park, near trail to fort, 15°35.049′N, 61°28.371′W, 2.XI.2004, SCC04_060 Guadeloupe: Les Saintes, top of Le Chameau, 15°51′28.1″N, 61°35′39.8″W, 6.III.2007, SCC07_039 Guadeloupe: Basse-Terre, Parc Archélogique des Roches Gravées, near Trois-Rivieres, 15°58.394′N, 61°38.347′W, 10.XI.2004, SCC04_064 19 20 Selenops micropalpus 21 Selenops micropalpus Selenops micropalpus Selenops micropalpus 22 23 24 Selenops micropalpus 25 Selenops micropalpus 26 Selenops micropalpus 27 Selenops micropalpus 28 Selenops micropalpus 29 Selenops micropalpus 30 Selenops micropalpus 31 Selenops micropalpus 32 Selenops n. sp. 7 33 Selenops n. sp. 7 sel_283 sel_277, sel_278, sel_279, sel_280, sel_281, sel_282 sel_091, sel_113 sel_088, sel_090, sel_092, sel_093, sel_094, sel_095, sel_112 sel_832, sel_833, sel_834 sel_820, sel_821, sel_822, sel_823 sel_825, sel_826, sel_827, sel_828, sel_829, sel_830, sel_831 sel_811, sel_812, sel_813, sel_814 sel_815, sel_816, sel_817, sel_818, sel_819 sel_804, sel_805, sel_806, sel_807, sel_808, sel_809, sel_810 sel_791, sel_792, sel_793, sel_794, sel_795, sel_797, sel_798, sel_799 sel_800, sel_801, sel_802, sel_803 sel_097, sel_098, sel_099, sel_100, sel_101 sel_102, sel_103, sel_104, sel_105 sel_106, sel_107, sel_108, sel_110, sel_111, sel_135 sel_783, sel_784, sel_785 sel_114 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 311 APPENDIX Continued Locality number Species Collection information Voucher numbers 34 Selenops n. sp. 7 sel_778, sel_779, sel_780, sel_781 35 Selenops n. sp. 7 36 Selenops n. sp. 7 37 Selenops n. sp. 7 Guadeloupe: Basse-Terre, Trois Rivieres, end of Trail Sentier de l′Acomat off Rue Nelson Mandela, 15°58′03.0″N, 61°37′50.1″W, 5.III.2007, SCC04_038 Guadeloupe: Basse-Terre, near Vieux Fort on the D6, along road at Forêt Domaniale du Litoral, 15°57.943′N, 061°42.517′W, 11.XI.2004, SCC04_065 Guadeloupe: Gran-Terre, Pointe du Chateaux, 16°14′51.6″N, 61°11′02.6″W, 7.III.2007, SCC07_040 Montserrat: Jack Boy Hill, 17°46′02.1″N, 62°10′17.0″W, 2.III.2007, SCC07_035 38 Selenops n. sp. 7 39 Selenops n. sp. 7 40 Selenops n. sp. 7 41 Selenops n. sp. 7 42 Selenops lindborgi 43 Selenops lindborgi 44 Selenops lindborgi 45 Selenops lindborgi 46 Selenops n. sp. 8 47 Selenops n. sp. 8 48 Selenops n. sp. 8 49 Selenops n. sp. 8 Montserrat: Sweet Water Ghaut, 16°47′07.2″N, 62°10′59.8″W, 2.III.2007, SCC07_036 Montserrat: Silver Hills, north side of Silver Hill, 16°48′41.3″N, 62°11′28.7″W, 3.III.2007, SCC07_037 Antigua: Nelson’s Dockyard National Park, Shirley’s Heights lookout, 17°00′06.7″N, 61°44′57.6″W, 27.II.2007, SCC07_034 Antigua: Indian Town, east of Veranda Resort, 17°05′50.2″N, 61°40′53.0″W, 27.II.2007, SCC07_033 St Kitts and Nevis: Nevis, Tamarind Bay, Galliput Restaurant, 17°09′48.53″N, 62°37′50.02″W, 23.II.2007, SCC07_030 St Kitts and Nevis: Nevis, Round Hill entrance to Mt Nevis, 17°11′13.83″N, 62°36′00.96″W, 23.II.2007, SCC07_029 St Kitts and Nevis: St Kitts, Major’s Bay, 17°13′37.9″N, 62°38′49.3″W, 24.II.2007, SCC07_032 St Kitts and Nevis: St Kitts, Sand Bank Bay, 17°14′59.1″N, 62°38′40.8″W, 24.II.2007, SCC07_031 Saba: Giles Quarter Trail, 17°36′54.36″N, 63°14′35.52″W, 12.III.2008 St Maarten: Mullet Bay, abandoned Mullet Bay Resort, 18°02′48.0″N, 63°027′29.7″W, 21.III.2007, SCC07_027 St Maarten: Emilio Wilson Estate and Park, 18°02′32.7″N, 63°03′53.1″W, 20.II.2007, SCC07_026 St Maarten: Upper Princess Quarter, 18°01′48.0″N, 63°02′08.0″W, 21.III.2007, SCC07_028 sel_115 sel_786, sel_787, sel_788, sel_789, sel_790 sel_758, sel_759, sel_760, sel_761, sel_762, sel_763, sel_764, sel_765, sel_766, sel_767, sel_768 sel_769 sel_771, sel_772, sel_773, sel_774, sel_775, sel_776, sel_777 sel_754, sel_755, sel_756, sel_757 sel_753 sel_740, sel_741, sel_742 sel_735, sel_736, sel_737, sel_738, sel_739 sel_750, sel_751, sel_752 sel_743, sel_745, sel_746, sel_747, sel_748, sel_749 sel_1021, sel_1022 sel_712, sel_713, sel_718, sel_719, sel_720, sel_721, sel_722, sel_723, sel_724, sel_725, sel_726, sel_727, sel_728, sel_729, sel_730 sel_711, sel_714, sel_715, sel_716, sel_717 sel_731, sel_732 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 312 S. C. CREWS and R. G. GILLESPIE APPENDIX Continued Locality number Species Collection information Voucher numbers 50 Selenops n. sp. 8 sel_116, sel_119, sel_120, sel_121, sel_708, sel_709, sel_710 51 Selenops mexicanus 52 Selenops n. sp. 8 53 Selenops n. sp. 8 54 Selenops n. sp. 8 55 Selenops n. sp. 8 56 Selenops n. sp. 8 57 Selenops lindborgi 58 Selenops lindborgi 59 Selenops lindborgi 60 Selenops lindborgi 61 Selenops lindborgi 62 Selenops lindborgi Selenops insularis Selenops lindborgi Selenops insularis Selenops lindborgi Selenops insularis Selenops lindborgi St Maarten: south-east side of island, trail from Back Bay to Geneve Bay, 18°00.929′N, 63°01′840W, 12–13.XI.2004 and 20.II.2007, SCC04_066, 068, SCC07_025 St Maarten: Philipsburg, Front Street, near entrance to cruise ship dock, 18°00.906′N, 63°02.587′W, 12.XI.2004, SCC04_067 Anguilla: Shoal Bay West, 18°09′52.8″N, 63°09′21.3″W, 13.II.2007, SCC07_016 Anguilla: The Cove, 18°10′14.1″N, 63°07′52.6″W, 13.II.2007, SCC07_017 Anguilla: Long Bay, Long Bay Beach, 18°11′29.3″N, 63°07′49.7″W, 13.II.2007, SCC07_015 Anguilla: Blowing Point, 18°10′18.0″N, 63°05′28.7″W, 13.II.2007, SCC07_018 Anguilla: Windward Point, 18°16′18.2″N, 62°58′05.3″W, 12.II.2007, SCC07_014 USVI: St Croix: Fredericksted, Sprat Hall Beach, Rte.63, 17°44′09.8″N, 64°53′24.0″W, 14.VI.2006, SCC06_056 USVI: St Croix: Sprat Hall Hill off West Shore Road, first right after subtracking station, 17°44′38.4″N, 64°53′22.3″W,14.VI.2006 SCC06_055 USVI: St Croix: Butler Bay, West Shore Road, 17°45′49.7″N, 64°52′58.8″W, 14.VI.2006, SCC06_054 USVI: St Croix: intersecting road from Creque Dam to Mahogany Road (intersects Mount Victory Camp) 17°44′27.1″N, 64°51′25.4″W, 14.VI.2006, SCC06_057 USVI: St Croix: Radio Telescope Station, east island, 17°45.398′N, 64°35.045′W, 18.XI.2004, SCC04_072 Puerto Rico: Vieques: Laguna Kiani, 18°07′02.2″N, 65°33′41.4″W, 19.VI.2006, SCC06_065 63 64 65 sel_117, sel_118 sel_697 sel_703, sel_704 sel_698, sel_699, sel_700, sel_701, sel_702 sel_705, sel_706, sel_707 sel_696 sel_497, sel_498, sel_499, sel_500 sel_494, sel_495, sel_496 sel_490, sel_491, sel_492, sel_493 sel_501 sel_527 sel_522, sel_523, sel_524, sel_525, sel_526 Puerto Rico: Vieques: Ruinas Central Playa Grande, 18°05′43.2″N, 65°31′13.2″W, 19.VI.2006, SCC06_064 sel_516, sel_517, sel_518, sel_519, sel_520, sel_521 Puerto Rico: Vieques: Cano Hondo, Cerca de Puerto Mosquito, 18°06′11.0″N, 65°27′05.5″W, 19.VI.2006, SCC06_061 sel_504, sel_505, sel_506, sel_507, sel_508, sel_509, sel_510, sel_511, sel_512 Puerto Rico: Vieques: Refugio Nacional de Vida Silvestre, road to Playa Caracas, Laguna Puerto Ferro, 18°06′24″N, 65°25′25.8″W, 19.VI.2006, SCC06_063 sel_513, sel_514, sel_515 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 313 APPENDIX Continued Locality number Species Collection information Voucher numbers 66 Selenops lindborgi sel_480, sel_481, sel_482, sel_483, sel_484, sel_485 67 Selenops lindborgi 68 Selenops lindborgi 69 Selenops lindborgi 70 Selenops lindborgi 71 Selenops lindborgi 72 Selenops lindborgi 73 Selenops lindborgi 74 Selenops lindborgi 75 Selenops lindborgi 76 Selenops lindborgi 77 Selenops lindborgi 78 Selenops lindborgi 79 Selenops lindborgi 80 81 Selenops submaculosus Selenops simius Puerto Rico: Culebra: Monte Resaca, 18°19′30.7″N, 65°18′10.5″W, 12.VI.2006, SCC06_052 Puerto Rico: Culebra: Brava Beach Trail, 18°19′38.9″N, 65°16′54.1″W, 12.VI.2006, SCC06_053 USVI: St Thomas, Estate Perserverance, Perserverance Bay Trail, 18°21.463′N, 64°59.753′W, 22.X.2004, SCC04_050 USVI: St Thomas, St Peter, house of R. Platenberg, 18°21′22.17″N, 64°56′49.53″W, 23.X.2004, SCC04_075 USVI: St Thomas, Magen’s Bay Trail, 18°21.350′N, 64°55.231′W, 22.X.2004 and 9.XI.2004, SCC04_052 and SCC04_073 USVI: St Thomas, Benner Hill, above armory, 18°19.533′N, 64°51.703′W, 19.XI.2004, SCC04_074 USVI: St Thomas, East End, Estate Nazareth, Dolphin House, 18°19.128′N, 64°51.567′W, 19.XI.2004, SCC04_076 USVI: St John, Bordeaux Ridge Road, 18°20.125′N, 64°43.672′W, 17.XI.2004, SCC04_071 USVI: St John, Cinnamon Bay Loop Trail, 18°21.226′N, 64°45.259′W, 16-17.XI.2004, SCC04_070 USVI: St John, Leinster Bay Trail, 18°21.825′N, 64°43.743′W, 16.XI.2004, SCC04_069 BVI: Tortola: vic. Sage Mountain, 18°24′46.44″N, 64°39′18.43″W, 20.X.2004, SCC04_049 BVI: Guana Island: north side near beach house, 18°28.793′N, 64°34.473′W, 18.X.2004, SCC04_047 BVI: Guana Island: south side near salt pond, 18°28.619′N, 64°34.475′W, 18.X.2004, SCC04_046 BVI: Virgin Gorda, lower trail up Gorda Peak, 18°28.774′N, 64°24.210′W, 19.X.2004, SCC04_048 Cuba: Sierra de Mesa, Pinar del Rio 82 Selenops simius Cayman Islands: Grand Cayman, Queen Elizabeth II Botanic gardens, storage facility south of main building, 19°19.055′N, 81°09.527′W, 30.IX.2004, SCC04_021 Cayman Islands: Grand Cayman, Queen Elizabeth II Botanic gardens, tree trail behind iguanas, 19°19.042′N, 81°10.081′W, 2.X.2004, SCC04_022 sel_486, sel_487, sel_488, sel_489 sel_076 sel_131 sel_124, sel_125, sel_128, sel_129, sel_130, sel_164 sel_123, sel_132, sel_133, sel_134 sel_143, sel_144, sel_145, sel_146, sel_147 sel_140, sel_141, sel_142, sel_157 sel_126, sel_127 sel_136, sel_137, sel_138, sel_139 sel_078 sel_071, sel_081, sel_085, sel_089 sel_079, sel_084, sel_086, sel_087 sel_073, sel_074, sel_075, sel_083, sel_089, sel_096, sel_109 sel_276 sel_046, sel_066, sel_067 sel_080 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 314 S. C. CREWS and R. G. GILLESPIE APPENDIX Continued Locality number Species Collection information Voucher numbers 83 Selenops simius sel_022 84 Selenops simius 85 Selenops candidus 86 Selenops candidus 87 Selenops candidus 88 Selenops n. sp. 17 89 Selenops candidus 90 Selenops candidus 91 Selenops candidus 92 Selenops petrunkevitchi Cayman Islands: Little Cayman, on road across street from Pirate’s Point Resort, 19°39.754′N, 80°06.032′W, 3.X.2004, SCC04_023 Cayman Islands: Cayman Brac, National Trust House off West End Road, 19°42.019′N, 79°52.084′W, 3.X.2004, SCC04_025 Jamaica: Westmoreland Paris, near New Hope on road toward Savanna-la-Mar, 18°14′55.4″N, 78°14′41.0″W, 29.V.2006, SCC06_027 Jamaica: St Ann Parish, North Coast Highway, between Discovery Bay and Rio Bueno, 18°28′31.3″N, 77°25′49.0″W, 28.V.2006, SCC06_024 Jamaica: Clarendon Parish, off road to Lluidasvale, 18°07′50.8″N, 77°10′05.0″W, 31 May 2006, SCC06_028 Jamaica: St Catherine Parish, Hellshire Hills, A2 Depression, 17°51′59.3″N, 76°57′54.0″W, 3.VI.2006, SCC06_031 Jamaica: St Mary Parish, near Mango Valley, 1.6 km off North Coast Highway, 18°24′23.4″N, 77°02′37.6″W, 28.V.2006, SCC06_023 Jamaica: St Andrew Parish, Castleton Botanic Gardens, 18°10′20.3″N, 76°49′27.6″W, 27.V.2006, SCC06_022 Jamaica: St Andrew Parish, Hermitage Dam Road, 2–6 km from junction with Stony Hill, 18°04′25.4″N, 76°47′01.3″W, 5.VI.2006, SCC06_033 Jamaica: St Thomas Parish, Blue Mountains National Park, Whitfield Hall, 18°02′54.8″N, 76°37′03.7″W, 1.VI.2006, SCC06_029 93 Selenops n. sp. 16 94 Selenops insularis Selenops pensilis Selenops n. sp. 13 95 96 Selenops marcanoi Jamaica: St Thomas Parish, near 12 mile Bull Bay, on left side of road heading east, 17°55′32.5″N, 76°38′31.0″W, 5.VI.2006, SCC06_034 Hispaniola: Haiti: Jacmel, St Cyr 72°31′41.2″N, 18°14′16.6″W, 23.X.2006, SCC06_078 Hispaniola: Dominican Republic: Prov. Pedernales: road to Aguacate from Rio Mulito, 18°13.895′N, 71°45.190′W, 25.XI.2004, SCC04_082 Hispaniola: Dominican Republic: Prov. Pedernales, Rio Mulito (El Banano), 18°09.165′N, 071°45.388′W, 25.XI.2004, SCC04_081 sel_023, sel_024, sel_025, sel_026, sel_027, sel_028 sel_362 sel_357, sel_358, sel_359, sel_360 sel_363, sel_364 sel_376, sel_377, sel_378, sel_379, sel_380, sel_381, sel_382, sel_383, sel_384 sel_352, sel_353, sel_354, sel_355, sel_356 sel_350, sel_351 sel_385 sel_365, sel_366, sel_367, sel_368, sel_369, sel_370, sel_371, sel_372, sel_373, sel_374, sel_375, sel_376 sel_386, sel_387, sel_388, sel_389, sel_390, sel_391 sel_655, sel_656, sel_657, sel_658, sel_659, sel_660, sel_661, sel_662, sel_663, sel_664, sel_665 sel_156 sel_150, sel_151, sel_152, sel_153, sel_154, sel_155 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 315 APPENDIX Continued Locality number Species Collection information Voucher numbers 97 Selenops phaselus sel_158, sel_160, sel_632, sel_633, sel_634 98 Selenops phaselus 99 Selenops n. sp. 14 Hispaniola: Dominican Republic: Prov. Pedernales, Parque Nacional Sierra de Baoruco, Las Abejas, 18°08.804′N, 71°37.164′W, 24-Nov-04 and 10-Oct-06, SCC04_077 and SCC06_072 Hispaniola: Dominican Republic: Prov. Pedernales, 26 km north of Cabo Rojo, Sierra de Baoruco, 18°06.490′N, 71°37.316′W, 24.XI.2004, SCC04_078 Hispaniola: Dominican Republic: Prov. Pedernales, 13–14 km north of Cabo Rojo on Carretera ALCOA, 18°01.962′N, 71°38.748′W, 24.XI.2004, SCC04_079 Hispaniola: Dominican Republic: Prov. Pedernales, Parque Jaragua, VII.2006 Hispaniola: Dominican Republic: Prov. Pedernales, Boca de la Cañada, Pedernales-N. sp. 15 Road, 9.X.2006, SCC06_071 Hispaniola: Dominican Republic: Prov. Pedernales, N. sp. 15, Fondo de Mama Cocoño, 25.X.2003, SCC03_021 Hispaniola: Dominican Republic, Prov. Pedernales, Laguna N. sp. 15, El Cajuil, 9.X.2006, SCC06_070 Hispaniola: Dominican Republic: Prov. Barahona, Carretera Higuero-Polo, 26.XI.2004, SCC04_083 Hispaniola: Dominican Republic: Prov. Barahona, Polo Coffee Plantation, July 2006 Hispaniola: Dominican Republic: Prov. Barahona, Barahona, CoralSol Resort, San Rafael Beach, July 2006 sel_537, sel_538, sel_539, sel_540, sel_541, sel_542, sel_543, sel_544, sel_545, sel_561, sel_563, sel_564, sel_567, sel_573, sel_574, sel_575, sel_576, sel_587, sel_588 sel_175 100 101 102 Selenops insularis Selenops n. sp. 14 Selenops n. sp. 15 Selenops insularis 103 Selenops n. sp. 15 104 Selenops insularis s. n. sp. 1 Selenops phaselus Selenops n. sp. 13 Selenops insularis 105 106 107 Selenops insularis 108 Selenops insularis 109 Selenops insularis 110 Selenops lindborgi Hispaniola: Dominican Republic: Prov. Peravia, Bani, road from Bani to Manaclar, past La Laguna, 18°21.343′N, 70°21.077′W, 27.XI.2004, SCC04_086 Hispaniola: Dominican Republic: Prov. Peravia, Bani, Rio Nizao, 18°16.915′N, 70°12.101′W, 27.XI.2004, SCC04_087 Hispaniola: Dominican Republic: Prov. San Cristóbal, Engombe, farm of autonomous university, 18°27.360′N, 70°00.306′W, 27.X.2003, SCC03_023 Hispaniola: Dominican Republic: Prov. La Altagracia, Parque del Este, Guaraguao, 18°19.968′N, 68°48.709′W, 30.XI.2004, SCC04_090 sel_148, sel_159, sel_215 sel_161, sel_162, sel_163, sel_165, sel_166, sel_167 sel_586, sel_589 sel_630, sel_631, sel_635, sel_636, sel_637 sel_018 sel_621, sel_622, sel_633, sel_634, sel_635, sel_636, sel_637, sel_638 sel_169, sel_170, sel_171, sel_172, sel_173 sel_562, sel_565, sel_566, sel_568, sel_585 sel_176, sel_177, sel_178, sel_179 sel_010, sel_011, sel_012, sel_013, sel_014, sel_015, sel_016, sel_017 sel_200, sel_201, sel_202, sel_203, sel_204, sel_205, sel_206, sel_207 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 316 S. C. CREWS and R. G. GILLESPIE APPENDIX Continued Locality number 111 112 113 114 115 116 117 118 Species Collection information Voucher numbers Selenops lindborgi Selenops insularis Selenops lindborgi Selenops insularis Selenops n. sp. 9 Selenops insularis Hispaniola: Dominican Republic: Prov. La Altagracia, Parque del Este, Boca de Yuma, 18°21.875′N, 68°37.080′W, 29-30.XI.2004, SCC04_089 Hispaniola: Domnican Republic: Prov. La Altagracia, Punta Cana Resort, 18°30′55.53″N, 68°22′28.73″W, 5-9.VII.2006, SCC04_066 sel_191, sel_192, sel_193, sel_194, sel_195, sel_196, sel_197, sel_199, sel_216 Hispaniola: Dominican Republic: Prov: Hato Major, Los Haitises, 1 km south of El Valley near Hato Mayor, 18°58′24.01″N, 69°22′34.58″W, VI.2006 Hispaniola: Dominican Republic: Prov. Samaná, Las Terrenas, 19°19′28.88″N, 69°32′50.77″W, VI.2006 sel_555 Hispaniola: Dominican Republic: Prov. La Vega, Constanza, Alto Cerro Hotel, 18°54′14.16″N, 70°44′41.14″W, VI.2006 Hispaniola: Dominican Republic: Prov. Puerto Plata, Sosua, VI.2006 Hispaniola: Dominican Republic: Prov. Puerto Plata, entrance to Loma de Isabel Torres, 19°46′41.9″N, 70°42′01.1″W, 7.X.2006, SCC06_068 Hispaniola: Dominican Republic, Prov. Santiago, Mata Grande, 19°11′43.0″N, 70°59′42.0″W, 14–15.X.2006, SCC06_075 Hispaniola: Dominican Republic, Prov. Santiago, Armando Bermudéz Park, ranger station at trail to Loma del Oro, 19°12′05.2″N, 71°00′04.8″W, 13.X.2006, SCC06_074 Hispaniola: Dominican Republic, Prov. Monti Cristi, Monte Cristi, El Morro, 19°53′42.44″N, 71°39′14.48″W, 8.X.2006, SCC06_069 sel_546 Selenops lindborgi Selenops insularis Selenops n. sp. 2 Selenops insularis Selenops insularis Selenops n. sp. 12 Selenops n. sp. 13 119 Selenops n. sp. 13 120 Selenops insularis Selenops n. sp. 10 121 122 Selenops insularis Selenops pensilis Selenops bani Selenops phaselus 123 Selenops n. sp. 3 124 Selenops insularis Hispaniola: Dominican Republic: Prov. Independencia, La Descubierta, El Azufrada, north side of Lago Enriquillo, 18°33.751′N, 71°41.853′W, 26.XI.2004, SCC04_084 Hispaniola: Haiti: Kenskoff, Belot-Montcel, 18°27′11.3″N, 72°21′06.4″W, 20-21.X.2006, SCC06_076 Puerto Rico: Isla Mona: Bajura de Empalme, 18°06′25.07″N, 67°53′10.41″W Puerto Rico: Isla Mona: Sardinera, 18°05′46.76″N, 67°56′12.35″W sel_528, sel_529, sel_530, sel_531, sel_532, sel_533, sel_534, sel_535, sel_536 sel_577, sel_590, sel_591, sel_592, sel_593, sel_594 sel_578 sel_604, sel_605, sel_606, sel_607, sel_608, sel_609 sel_640, sel_641, sel_642, sel_643, sel_644, sel_645, sel_646 sel_639 sel_579, sel_580, sel_581, sel_582, sel_583, sel_584, sel_610, sel_611, sel_612, sel_613, sel_614, sel_615, sel_616, sel_617, sel_618 sel_180, sel_181, sel_182, sel_183, sel_184, sel_185, sel_186, sel_187, sel_188, sel_189, sel_190, sel_569, sel_570, sel_571, sel_572, sel_599 sel_647, sel_648, sel_649, sel_650, sel_651, sel_652, sel_653, sel_654 sel_846, sel_847 sel_838, sel_839, sel_840, sel_841, sel _842, sel_843, sel_844, sel_845 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 317 APPENDIX Continued Locality number Species Collection information Voucher numbers 125 Selenops n. sp. 3 sel_502 126 Selenops n. sp. 3 Selenops insularis Selenops lindborgi Selenops lindborgi Selenops insularis Selenops lindborgi Selenops insularis Selenops lindborgi Selenops insularis Selenops lindborgi Selenops insularis Selenops insularis Puerto Rico: Isla Mona: Camino de los Cobros, 18°04′02.68″N, 67°52′45.75″W, VII.2006 Puerto Rico: Maricao, Bosque Estatl de Maricao, 18°08′51.2″N, 66°59′35.0″W, 10.VI.2006, SCC06_045 sel_434, sel_435, sel_436, sel_437, sel_438, sel_439, sel_440, sel_441, sel_442 Puerto Rico: Susua State Forest, Sabana Grande, 18°04′15.0″N, 66°54′31.6″W, 10.VI.2006, SCC06_046 sel_443, sel_444, sel_445, sel_446, sel_447, sel_448, sel_449, sel_450, sel_451, sel_452, sel_453 Puerto Rico: Quebradillas, Merendero de Guajataca, 18°29′23.7″N, 66°56′59.4″W, 9.VI.2006, SCC06_043 sel_423, sel_424, sel_425, sel_426, sel_427, sel_428, sel_429, sel_430, sel_431, sel_432 Puerto Rico: Arecibo, Arenalejos, Carretera 657, km 1.9, 18°25′15.9″N, 66°40′35.2″W, 7.VI.2006, SCC06_035 sel_392, sel_393, sel_394, sel_395, sel_396, sel_397, sel_398 Puerto Rico: between Barceloneta and Arecibo, Bosque Cambalacheo, 18°27′07.0″N, 66°35′49.9″W, 9.VI.2006, SCC06_041 sel_419, sel_420, sel_421, sel_422 Puerto Rico: Ciales, Bosque Fronton, Carretera 146, km 16.3 Interior Camino Maximo Nuñez, Sector Los Gonzalez, 18°18′33.8″N, 66°32′42.8″W, 15.VI.2006, SCC06_060 Puerto Rico: Coamo, Baños de Coamo, 18°02′19.4″N, 66°22′27.0″W, 11.VI.2006, SCC06_048 Puerto Rico: Salina, Reserva Jobos, Parque Jagueys, 17°57′13.9″N, 66°15′03.5″W, 11.VI.2006, SCC06_049 Puerto Rico: Toa Baja, Bosque Media Luna, PR-2, km 21–6, 18°24′38.30″N, 66°14′44.36″W, 14.VI.2006, SCC06_058 Puerto Rico: Manuabo, Mariani Creek, 18°00′29.7″N, 65°52′17.0″W, 11.VI.2006, SCC06_050 sel_503 127 128 129 130 131 132 Selenops insularis 133 Selenops lindborgi 134 Selenops insularus 135 Selenops lindborgi Selenops insularis Selenops insularis 136 137 Selenops lindborgi 138 Selenops lindborgi Selenops insularis Puerto Rico: Humacao, Barrio Collores, 18°09′44.82″N, 65°49′06.50″W, 11.VI.2006, SCC06_051 Puerto Rico: Loiza, Punta Vacia, Talega, 18°27′03.8″N, 65°54′16.7″W, 8.VI.2006, SCC06_038 Puerto Rico: Ceiva, Los Corchos, 18°12′13.8″N, 65°40′06.5″W, 8.VI.2006, SCC06_040 sel_462, sel_463 sel_464 sel_461 sel_465, sel_466, sel_467, sel_468 sel_469, sel_470, sel_471, sel_472, sel_473, sel_474, sel_475, sel_476, sel_477, sel_478, sel_479 sel_399, sel_400, sel_401, sel_401, sel_402, sel_403 sel_416, sel_417, sel_418 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 318 S. C. CREWS and R. G. GILLESPIE APPENDIX Continued Locality number 139 140 Species Collection information Voucher numbers Selenops lindborgi Selenops insularis Selenops submaculosus Puerto Rico: Fajardo, Seven Seas Public Beach, 18°22′03.7″N, 65°38′04.9″W, 8.VI.2006, SCC06_039 sel_405, sel_406, sel_407, sel_408, sel_409, sel_410, sel_411, sel_412, sel_413, sel_414, sel_415 Bahamas: Andros Island: Owens Town, 24°52′30.1″N, 78°02′03.6″W, 13.V.2006, SCC06_002 Bahamas: Andros Island: Morgan’s Cave at Morgan’s Bluff, 25°10′30.1″N, 78°01′26.2″W, 13.V.2006, SCC06_003 Bahamas: Andros Island: International Field Station, 24°53′51.1″N, 77°55′50.1″W, 12.V.2006, SCC06_001 sel_302, sel_303, sel_304, sel_305, sel_306, sel_307, sel_308 141 Selenops submaculosus 142 Selenops submaculosus 143 Selenops submaculosus 144 145 Selenops submaculosus Selenops aissus 146 Selenops aissus 147 Selenops submaculosus 148 Selenops aissus 149 Selenops aissus 150 Selenops aissus 151 Selenops lindborgi 152 Selenops lindborgi 153 Selenops n. sp. 11 Bahamas: Pigeon Cay, near IFS on Andros Island, 24°52′54.4″N, 77°53′53.5″W, 13.V.2006, SCC06_004 Bahamas: Andros Island: Cargill Creek, 24°30′00.37″N, 77°43′15.61″W, 13.V.2006 Bahamas: Abaco: Ralph’s Chimney off Queen’s (Abaco) Highway, 26°14′58.2N, 77°11′25.4″W, 14.V.2006, SCC06_006 Bahamas: Abaco: Abaco National Park, 26°03′44.0″N, 77°12′46.2″W, 14.V.2006, SCC06_005 Bahamas: Great Exuma: Bahamas Sound Subdivision near old airport, 23°27′56.0″N, 75°46′24.8″W, 18.V.2006, SCC06_011 Bahamas: Great Exuma: Regatta Point, 23°30′24.7″N, 75°45′58.0″W, 18.V.2006, SCC06_009 Bahamas: Stocking Island, near Great Exuma, 23°32′08.9″N, 75°46′29.6″W, 18.V.2006, SCC06_010 Bahamas: San Salvador: Gerace Field Station, trails behind field station, 24°06.9′N, 74°27.8′W, 19.V.2006, SCC06_012 Bahamas: Great Inagua: Man o’War Bay, 21°04′30.2″N, 73°38′36.7″W, 16.V.2006, SCC06_007 Bahamas: Great Inagua: Old Aerostat Base, 21°06.06.7″N, 73°39′01.9″W, 16.V.2006, SCC06_228 Turks and Caicos: Providenciales, North-West Point Pond Nature Reserve, 21°50′32.1″N, 72°19′43.7″W, 8.II.2007, SCC07_010 and SCC07_011 sel_309, sel_310, sel_311, sel_666 sel_286, sel_287, sel_288, sel_289, sel_290, sel_291, sel_292, sel_293, sel_294, sel_295, sel_296, sel_297, sel_298, sel_299, sel_300, sel_301, sel_556, sel_557, sel_558, sel_595, sel_596, sel_667, sel_669, sel_671, sel_835, sel_836 sel_312 sel_668 sel_315 sel_313 sel_332 sel_319, sel_320, sel_321, sel_322, sel_323, sel_324 sel_325, sel_326, sel_327, sel_328, sel_329, sel_330, sel_331 sel_333, sel_334, sel_335, sel_336, sel_337, sel_338, sel_339, sel_340, sel_341, sel_342, sel_343 sel_316 sel_317, sel_318 sel_689, sel_690, sel_691, sel_692 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 319 APPENDIX Continued Locality number Species Collection information Voucher numbers 154 Selenops n. sp. 11 sel_346, sel_347, sel_348, sel_349, sel_694, sel_695 155 Selenops n. sp. 11 156 Selenops n. sp. 11 157 Selenops n. sp. 11 158 Selenops n. sp. 11 159 Selenops debils gp. species 1 160 Selenops debils gp. species 1 161 Selenops debils gp. species 1 162 Selenops debils gp. species 1 163 Selenops debils gp. species 1 164 Selenops debils gp. species 1 Selenops debils gp. species 1 Turks and Caicos: Providenciales, Turtle Cove, Third Turtle Drive, 21°47′01.1″N, 72°13′45.4″W, 22.V.2006 and 10.II.2007, SCC06_017 and SCC07_013 Turks and Caicos: Providenciales, The Bight, 21°47′00.6″N, 72°13′06.4″W, 22.V.2006, SCC06_015 Turks and Caicos: North Caicos, Wade’s Green Plantation, 21°55′13.36″N, 72°01′12.45″W, 2–3.II.2007, SCC07_001 and SCC07_005 Turks and Caicos: Middle Caicos, Garden Pond Field Road, 21°48′24.76″N, 71°45′42.78″W, 3.II.2007, SCC07_004 Turks and Caicos: Providenciales, South View Drive off of Leeward Highway, 21°46′45.7″N, 72°13′45.4″W, 10.II.2007, SCC07_012 USA: California, San Diego Co., Upper Otay River Valley, 32°59′50.33″N, 116°19′09.27″W, 19.V.2003 USA: California, San Diego Co., Jamul, Lyons Valley, north of Lyons Peak, 32°44′03.93″N, 116°53′51.23″W, 29.VII.2007 USA: California, San Diego Co., Anza Borrego Desert State Park, Carrizo Palm Grove, 32°44′31.29″N, 116°12′51.11″W Mexico: Baja California Sur, 2 km east of Ballena between San Ignacio and San Juanico, 26°27′10.60″N, 111°34′53.39″W Mexico: Baja California Sur, 3 km west of Villa Insurgentes on road to San Miguel de Comundu, 25°16′50.46″N, 111°50′03.75″W Mexico: Baja California Sur, Cuevas Pintas, 26°01′38.63″N, 111°30′24.45’W USA: Arizona, Coconino Co., Monument Trail, flat near archaeological site, 36°25.309′N, 112°27.483′W, 13.VIII.2004, SCC04_013 USA: Arizona: Santa Cruz Co., Madera Canyon, Mt Wrightson Trail, 31°43′06.86’N, 110°52′22.45’W Mexico: Sonora, Sonoran Highway on road to Yecora, 28°23′15.01’N, 108°55′41.42’W USA: Texas: Val Verde Co., Seminole Canyon, Highway 90, under bridge, 29°42′21.12’N, 101°18′28.48’W Mexico: Hidalgo, Villa Flores, Ejido ′El Rayo’, 24°14′1.00’N, 99°1′4.50’W, 2.XI.2007 Mexico: Colima, Manzanillo, Municipio Manzanillo, 1.2 to 1.4 km east La Central, 19°8′56.4’N, 104°25′35.3’W, X.2005 Mexico: Colima, Municipio Ixtlahuacan, Tamala, 19°5′2.13’N, 103°47′26.21″ 165 166 Selenops debils gp. species 2 167 Selenops debils gp. species 3 Selenops debils gp. species 3 168 169 170 Selenops mexicanus Selenops abyssus 171 Selenops abyssus sel_344, sel_345 sel_675, sel_680, sel_681, sel_682, sel_683, sel_684, sel_685, sel_686, sel_687, sel_688, sel_689 sel_676, sel_677, sel_678, sel_679 sel_693 sel_002 sel_837 sel_021, sel_214 sel_210, sel_213 sel_212 sel_009 sel_211 sel_264, sel_270, sel_271, sel_272 sel_851 sel_208 sel_1008 sel_1004 sel_1013 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 320 S. C. CREWS and R. G. GILLESPIE APPENDIX Continued Locality number Species Collection information Voucher numbers 172 Selenops abyssus sel_1006, sel_1012 173 Selenops n. sp. 4 174 Selenops gracilis 175 Selenops nigromaculatus? This is an immature specimen, but is found near the type locality of S. nigromaculatus, hence the ‘?’ Selenops n. sp. 4 Mexico: Michoacan, Municipio Coalcomán, Coalcomán, 18°24′12.81″N, 103°07′58.82″W, X.1005 Mexico: Morelos, Cuernavaca, Colonia Chamilpa, Instituto de Biotecnología, UNAM, 18°55′51.86″N, 99°14′16.60″W Mexico: Guerrero, Arcelia, Campo Morado, 17°34′60.00″N, 100°4′60.00″W Mexico: Guerrero, Omiltemi, Chilpancingo, Camino al Omiltemi, 17°33′3.76″N, 99°30′21.83″W Mexico: Puebla, Municipio Zapotitlan de las Salinas, Cerro el Pajarito, 18°22′48.3″N, 97°30′26.9″W, V.2005 Mexico: Chiapas, Tuxtla-Gutierrez, Cañon deSumidero, 16°49′41.46″N, 93°6′22.22″W Mexico: Chiapas, Berriozabal, dirt road from Efrain A. Gutierrez, approx. 8 km north Berriozabal, 16°52′27.45″N, 93°17′28.20″W Mexico: Chiapas, Pueblo Nueva Solistahuacan, 17°11.550′N, 92°54.875′W, 17.IX.2004, SCC04_018 Mexico: Chiapas, La Reforma, Municipio La Concordia, 15°54.212′N, 92°40.157′W, 18.IX.2004, SCC04_018b Mexico: Chiapas, Huixtla, Las Golindrinas, 15°25.747′N, 92°39.270′W, 23.IX.2004, SCC04_020 sel_1002, sel_1005 176 177 178 Selenops mexicanus Selenops mexicanus 179 Selenops n. sp. 19 180 Selenops mexicanus 181 Selenops mexicanus Selenops n. sp. 19 Selenops mexicanus Selenops n. sp. 19 Selenops mexicanus 182 183 184 185 186 Selenops mexicanus Selenops mexicanus Selenops n. sp. 18 Selenops bifurcatus Mexico: Chiapas, Municipio Motozintla de Mendoza, Chevolcan, 15°20′52.4″N, 92°19′25.4″W, 21.XI.2004, SCC04_019 Mexico: Chiapas: road to Roberto Barrio, ~4 km south-west Nuevo Sonora, 17°23′41.10″N, 91°54′10.70″W Mexico: Veracruz, Municipio Tamalin, El Mamey, 21°31′13.63″N, 97°38′31.75″W Guatemala: Petén, Sta Elena de la Cruz, Colonia del Bosque, near Flores, Cueva Actun Kan, 16°54′10.9″N, 89°53′44.3″W, 1.I.2008, SCC08_001 Guatemala: Zacatán, Las Guacamayas, Carretera Sta Rosalia Marmol, Hídroelectrica Pasabíen, 15°01′39.7″N, 89°41′41.2″W, 1.I.2008, SCC08_002 sel_1010 sel_1014, sel_1015 sel_1007 1016, 1017 sel_848 sel_043 sel_044 sel_031, sel_034, sel_035, sel_036, sel_037, sel_038, sel_039, sel_040, sel_041, sel_045, sel_1011 sel_029, sel_030, sel_032, sel_033, sel_038 sel_849 sel_1018, sel_1019, sel_1020 sel_865, sel_866, sel_867, sel_868 sel_869, sel_870, sel_871, sel_872, sel_873, sel_874, sel_875, sel_876, sel_877, sel_878, sel_879 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 321 APPENDIX Continued Locality number Species Collection information Voucher numbers 187 Selenops bifurcatus sel_927 188 Selenops bifurcatus 189 Selenops bifurcatus Selenops mexicanus Selenops bifurcatus El Salvador: Dep. Chaletenango, Chaletenango off Carretera Troncal del Norte, Hotel Maya, 14°16′29.1″N, 89°08′32.7″W, 5-6.I.2008, SCC08_009 El Salvador: Dep. Chaletenango, Mun. Chaletenango, La Cueva del Corridor, 6.I.2008, SCC08_010 El Salvador: Dep. Chaletenango, Chaletenango, Municipio Tejutto, Rest. Eucalyptos, 14°12′20.4″N, 89°06′43.9″W, 5.I.2008, SCC08_008 El Salvador: San Salvador, Museo Nacional de Historia Natural grounds, 13°40′23.4″N, 89°11′53.6″W, 3.I.2008, SCC08_003 El Salvador: near San Vicente, Mun. Tepetitán vic. Finca El Carmen, 13°37′53.0″N, 88°50′19.5″W, 4.I.2008, SCC08_005 190 191 Selenops bifurcatus Selenops mexicanus 192 Selenops bifurcatus 193 Selenops bifurcatus 194 Selenops bifurcatus 195 Selenops bifurcatus 196 Selenops bifurcatus Selenops mexicanus Selenops bifurcatus 197 198 Selenops bifurcatus 199 Selenops mexicanus 200 Selenops bifurcatus Selenops mexicanus El Salvador: Dep. San Vicente, vic. San Vicente, road to Zacatecoluca, behind strip club ‘Dreamed Girl’, 13°37′43.4″N, 88°46′49.6″W, 3.I.2008, SCC08_004 El Salvador: Dept La Union, Mun. El Carmen, Lotificacion Amaya, 13°21.44.9″N, 87°59′58.2″W, 5.I.2008, SCC08_007 El Salvador: Dept La Union, Conchagua, Volcán Conchagua near La Union, 13°18′14.1″N, 87°51′19.6″W, 4.I.2008, SCC08_006 Nicaragua: Dept Nuevo Segovia, Alc. Ocotal, Barrio Roberto Gomez above Rio Coco, 13°37′05.8″N, 86°27′57.3″W, 11.I.2008, SCC08_012 Nicaragua: Dept Madríz, Alc. Ocotal, Totogalpa, 13°33′49.5″N, 86°29′54.6″W, 11.I.2008, SCC08_013 Nicaragua: Dept Leon, Alc. San Jacinto, Mina El Límon, Rancho Las Brisas, 12°37′03.8″N, 86°44′34.3″W, 14.I.2008, SCC08_016 Nicaragua: Dept Leon, Alc. El Jicaral, Camino Santa Rosa, Puente La Guayabita, 12°44′31.2″N, 86°22′44.6″W, SCC08_017, SCC08_017 Nicaragua: Dept Matagalpa, Alc. San Ramon, Mata Palo, 12°56′16.5″N, 85°51′12.2″W, 14.I.2008, SCC08_018 Nicaragua: Dept Boaco, Aguas calientes, Alc. Teustepe, Camino La Cuesta, 12°22′57.8″N, 85°47′30.7″W, 15.I.2008, SCC08_020 sel_925, sel_926, sel_927, sel_928, sel_929 sel_921, sel_922, sel_923, sel_924 sel_881, sel_882, sel_883, sel_884, sel_885, sel_886, sel_887, sel_888, sel_889 sel_897, sel_898, sel_899, sel_900, sel_901, sel_902, sel_903, sel_904, sel_905, sel_906, sel_907, sel_908, sel_909, sel_910, sel_911 sel_890, sel_891, sel_892, sel_893, sel_894, sel_895, sel_896 sel_916, sel_917, sel_918, sel_919, sel_920 sel_912, sel_913, sel_914, sel_915 sel_930 sel_931, sel_932, sel_933, sel_934, sel_935, sel_936, sel_937, sel_938, sel_939 sel_954, sel_955, sel_956, sel_957, sel_958, sel_959, sel_960 sel_961, sel_962, sel_963, sel_964, sel_965, sel_966, sel_967, sel_968, sel_969, sel_970 sel_971, sel_972, sel_973, sel_974 sel_963, sel_975, sel_976, sel_977, sel_978, sel_979, sel_980, sel_981, sel_982, sel_983, sel_985, sel_986 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 322 S. C. CREWS and R. G. GILLESPIE APPENDIX Continued Locality number Species Collection information Voucher numbers 201 Selenops bifurcatus Selenops mexicanus sel_940, sel_941, sel_942, sel_943, sel_944, sel_945, sel_946, sel_947, sel_948, sel_949, sel_950, sel_951, sel_952, sel_953 202 Selenops mexicanus 203 Selenops bifurcatus Selenops mexicanus Selenops mexicanus Nicaragua: Lago Nicaragua, Isla Ometepe, Volcán Concepción, Charco Verde, Hotel Finca Vincenzia and up hill across the street, 11°28′42.6″N, 35°38′20.6″W and 11°29′31.2″N, 85°38′14.1″W, 12-13.I.2008, SCC08_014, SCC08_015 Costa Rica: Guanacaste, Palo Verde National Park, Cueva Las Tigres, 10°21′58.9″N, 85°21′14.2″W, 17.I.2008, SCC08_022 Costa Rica: Guanacaste: Palo Verde Field Station, hill behind OTES office, 10°20′42.5″N, 85°20′19.1″W, 17.I.2008, SCC08_021 Costa Rica: Guanacaste: Nicoya Peninsula, near Loma Bonita, 10°15′04.0″N, 85°17′30.5″W, 18.I.2008, SCC08_023 Panamá: Barro Colorado Island, I.2008 Panamá: STRI, Galeta, Plot F, 2004 sel_1000, sel_1001 sel_265, sel_266, sel_267, sel_268 Japan: Kyoto, Shugakuin, 23.VIII.2007 sel_861, sel_862, sel_863 Namibia: 12.VI.2006 Tanzania: Iringa, Lutheran House Hostel South Africa: Guateng, Roodeport, Ruimsig Butterfly Reserve South Africa: Tsitsikamma National Park, 78 km E Knysna South Africa: Grahmstown Municipal. Caravan Park South Africa: Eastern Cape, Kai Mouth, 58 km NE East London South Africa: Table Mountain National Park, Newland’s Forest Madagascar: Reserve Nat. Integrale de Lokobe, 3.61 km ESE Hellville Madagascar: Park National Montagne d’Ambre montane rainforest Australia: Western Australia: Ravensthorpe, Ravensthorpe Ranges South, WAM 10, 33°38′16.03″S, 120°10′46.01″E, 17.V.2007, under rock, MCLeng, ML Moir Australia: Western Australia: Two Peoples Bay Nature Reserve, granite outcrop, site 6, 34°59′18″S, 118°44″E, 14.X.2006, under granite rock, ML Moir, JM Waldock Australia: Western Australia: Fiztgerald River National Park, East Mt Barren, site 7, 33°55′28″S, 120°01′13″E, 25.XI.2006, under rock ML Moir, KEC Brennan sel_998, 999 sel_997 sel_547, sel_548 204 205 206 207 208 209 210 211 212 213 214 215 216 Selenops banksi Selenops mexicanus Selenops bursarius Selenops radiatus Selenops radiatus Anyphops barnardi Anyphops parvulus Anyphops tugelanus Anyphops stauntoni Anyphops kraussi Garcorops madagascarensis Hovops sp. 217 New genus sp. 1 Australia 218 New genus sp. 2 Australia 219 New genus sp. 3 Australia sel_989, sel_990 sel_987, sel_988 sel_991, sel_992, sel_993, sel_994 sel_549 sel_550 sel_551 sel_552 sel_553 sel_275 T80881, T80996 T78485, T78489 T78500, T78498 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322