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

Molecular systematics of Selenops spiders (Araneae:Selenopidae) from North and Central America:implications for Caribbean biogeography

SARAH C. CREWS1,2* and ROSEMARY G. GILLESPIE1

1University of California Berkeley, Department of Environmental Sciences Policy and Management,137 Mulford Hall, Berkeley, CA 94720-3114, USA2Berkeley City College, Department of Science and Biotechnology, 2050 Center Street, Berkeley, CA94704, USA

Received 16 February 2010; revised 3 May 2010; accepted for publication 3 May 2010bij_1494 288..322

The Caribbean region includes a geologically complex mix of islands, which have served as a backdrop for somesignificant studies of biogeography, mostly with vertebrates. Here, we use the tropical/subtropical spider genusSelenops (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 taxaand nearly 60 selenopid species. Phylogenetic analysis of the data revealed several biogeographic patterns commonto other lineages that have diversified in the region, the most significant being: (1) a distinct biogeographic breakbetween 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 Antillesrelative to the Lesser Antilles. However, a strikingly unique pattern in Caribbean Selenops is that Cuban speciesare not basal in the Caribbean clade. Analyses to test competing hypotheses of vicariance and dispersal supportcolonization through GAARlandia, an Eocene–Oligocene land span extending from South America to the GreaterAntilles, rather than over-water dispersal. © 2010 The Linnean Society of London, Biological Journal of theLinnean Society, 2010, 101, 288–322.

ADDITIONAL KEYWORDS: Bayesian phylogenetics – island biogeography – likelihood analysis ofgeographic range evolution.

INTRODUCTION

Remote islands form the basis for many biologicalstudies because of their ability to act as a laboratory,with repeated sets of ecological and/or evolutionaryexperiments occurring within a circumscribed timeframe (Cronk, 1997; Losos et al., 1998; Gillespie &Roderick, 2002; Gillespie, 2004; Ricklefs & Berming-ham, 2008). While the Hawaiian Islands have servedas a model system for processes of in situ diversifica-tion, the long history of studies on the biota of theCaribbean has provided some of the most important

insights into the complex interaction between coloni-zation and diversification. In particular, the Carib-bean has served as the setting for the establishmentof most of the central tenets in the equilibrium theoryof island biogeography (Munroe, 1948), the argu-ments being formulated independently by MacArthurand Wilson (1963, 1967) much later (Lomolino &Brown, 2009). More recent research on the islandshas allowed an understanding of the interplaybetween ecological and evolutionary processes inshaping species diversity (Losos & Schluter, 2000;Schoener, Spiller & Losos, 2001).

The primary feature of the Caribbean region thatmakes it particularly useful for examining the inter-action between colonization and diversification is its*Corresponding author. E-mail: [email protected]

Biological Journal of the Linnean Society, 2010, 101, 288–322. With 9 figures

© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322288

long and complex geological history. The CaribbeanBasin began forming nearly 140 Mya. Islands in thebasin consist of four different types: (1) land-bridgeislands which were connected to each other or to themainland at times of lower sea level; (2) continentalislands which broke off from the mainland throughtectonic displacement; (3) uplifted limestone islands;and (4) volcanic islands (MacPhee & Iturralde-Vinent,2005; Robertson, 2009). Despite their limited isola-tion, the age and geologic complexity of the area haveprovided ‘well-defined paths of entry by which immi-grants may reach’ the islands (Munroe, 1948). More-over, the islands have served as the setting foradaptive radiation among lineages with limited dis-persal ability, in particular lizards of the genus Anolis(Losos, 1992, 1994, 2009), frogs of the genus Eleuth-erodactylus (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 (Francisco-Ortega et al., 2008)]. Although studies to date haveprovided insights into how the individual lineageshave colonized and subsequently diversified withinthe island system, notable controversies remain,including the source of colonists and the means bywhich they colonized the islands, biogeographic pat-terns within lineages and whether these patternsmight be expected to be shared across multiple lin-eages (Guyer & Savage, 1986; Williams, 1989;Hedges, Hass & Maxon, 1992; Crother & Guyer, 1996;Hedges, 1996a,b).

A particular focus of debate has been the role ofvicariance vs. dispersal in shaping the Caribbeanbiota. Hedges and colleagues (Hedges et al., 1992;Hedges, 1996a,b; Hedges & Heinicke, 2007; Heinickeet al., 2007), working with herpetofauna, have sug-gested that the absence of lineages older than thebreak-up of the proto-Antilles (a contiguous landmass between North and South America) precludes avicariant origin and they argue for the initial coloni-zation of most taxa via over-water dispersal onflotsam. A similarly dominant role for dispersal hasbeen suggested for multiple lineages of plants, suchas Miconieae (Michelangeli et al., 2008). In contrast,other studies have suggested that vicariance hasplayed a larger role than dispersal in the initialcolonization of the Caribbean; for example, in lizards(Crother & Guyer, 1996; Iturralde-Vinent & MacPhee,1999; MacPhee & Iturralde-Vinent, 2005) and someplants [e.g. Euphorbiaceae (van Ee et al., 2008)].

A related controversy focuses on the hypothesis ofGAARlandia (Greater Antilles + Aves Ridge), firstproposed by Iturralde-Vinent & MacPhee (1999), whoused geological data and fossil evidence to demon-strate the likely existence of a land span connecting

South America to the Greater Antilles during theEocene–Oligocene transition 35–33 Mya. The landspan, although probably short-lived, may have pro-vided an avenue for terrestrial organisms to colonizethe Greater Antilles from South America. Amongmammals, molecular phylogenies of primates andhystricognath 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 (Sty-racaceae) (Fritsch, 2003) show that the timing ofdivergence of lineages is consistent with the GAAR-landia hypothesis. However, similar data fromendemic legume radiations in the Greater Antilles,although initially thought to indicate ancient splittingbetween lineages consistent with the GAARlandiahypothesis (Lavin et al., 2001), show more recentdiversification (Lavin & Beyra-Matos, 2008), which islikely to hold also for lineages of Asteraceae(Francisco-Ortega et al., 2008).

Clearly, the timing and frequency of dispersal andvicariance, and the interplay between the two, variesacross biotic assemblages. The challenge, then, is tounderstand the circumstances dictating the relativeroles of each and how they interact. Arthropods,because they can provide a fine-scale resolution ofbiogeographic patterns (Ferrier et al, 2004), are idealcandidates for elucidating the nature of these rela-tionships. Although the biogeography of terrestrialinvertebrates in the Caribbean has been examined insome detail (see Liebherr, 1988a and chapterstherein), few recent studies have been attempted,with little molecular information on the timing andnature of the interplay between colonization anddiversification. However, there are some notableexceptions (Davies & Bermingham, 2002; Wilder &Hollocher, 2003; Brisson, Wilder & Hollocher, 2006).In particular, recent studies on spiders (Sicariidae:Loxosceles) support the GAARlandia hypothesis in thecolonization of the lineage of North from SouthAmerica (Binford et al., 2008), while crickets show amore mixed pattern of both vicariance and dispersal,coupled with intra-island diversification (Oneal,2009).

In this study, we combine molecular and morpho-logical methods to examine the phylogenetic relation-ships and biogeographic history of the cursorial anddispersal-limited spider genus Selenops (Araneae:Selenopidae) in the Caribbean. These primarily tropi-cal and subtropical spiders (Muma, 1953; Corronca,1998; Alayón, 2005) are distinctive in that they areextremely dorsoventrally flattened and exceedinglyfast. They are found in a variety of habitats andmicrohabitats (Crews, Wienskoski & Gillespie, 2008).Although the genera and species groups have

SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 289


undergone several revisions (Muma, 1953; Corronca,1998; Alayón, 2005), there is no phylogenetic frame-work for the family or for any of the componentgenera. They were chosen for the current studybecause of their high diversity and abundance in theCaribbean, where they occur in both the Greater andLesser Antilles, as well as on the adjacent mainland(southern North America and throughout SouthAmerica) (Muma, 1953; Crews, 2005; Crews et al.,2008, 2009;) (Figs 1, 2). Accordingly, they provide thepotential to reveal fine-scale biogeographic patternsacross the islands of the Caribbean. The currentstudy uses the genus to infer the relative importanceof the following two processes in dictating the biogeo-graphic history of the lineage in the Caribbean: (1)the frequency of colonization to the Caribbean regionfrom a mainland source and between islands withinthe Caribbean; and (2) whether within-island diver-sification has occurred through a single radiation orthrough dispersal and multiple radiations. We alsotest the hypotheses of dispersal and vicariance in theframework of the GAARlandia hypothesis and likeli-hood biogeographic analysis.

MATERIAL AND METHODSTAXON SAMPLING

A comprehensive geographic sample of the genus wasobtained from the Caribbean region, including mostislands and several sites throughout Mexico, CentralAmerica and the South American mainland (see alsoSupporting Information, Figs S1, S2). Political reasonsprohibited us from obtaining permits to collect severalendemic species from Cuba and the single species fromNavassa Island. The implications for these omissionsare discussed at the end of this paper. Outgroupsincluded other genera in the family Selenopidae fromall major geographic locations where the family isfound, in particular the type of the genus (Selenopsradiatus Latreille) from Africa, Selenops bursariusKarsch from Japan, Selenops montigenus Simon fromNepal/India and representatives of the three othergenera described from Africa (six species of Anyphops,one species of Hovops and one species of Garcorops),as well as an undescribed Australian genus. Chosenrepresentatives outside of the family include a broadsample of eight genera from two families, the

Figure 1. Map of the study area. The Americas; the boxed region shows the primary study area.

290 S. C. CREWS and R. G. GILLESPIE


Sparassidae and the Ctenidae, and were based onunpublished data (M. Ramirez, pers. comm.) (Table 4).

The genus Selenops has also been found in Domini-can amber and one of these specimens is an adultmale, described by Schawaller (1984) as Selenopsbeynai. The specimen was scanned using X-ray com-puted tomography, as in Penney et al. (2007);however, the poor preservation of the genitalia pro-hibited even tentative incorporation into the phyloge-netic framework.

MOLECULAR METHODS

Four gene fragments were amplified – three mitochon-drial [cytochrome oxidase I (CO1), 16S ribosomal DNA(16S) and the intervening leucine tRNA and NADHdehydrogenase I (ND1)] and one nuclear [histone 3a(H3)] (see Table 1). The respective lengths of the

amplification products were ~850, ~800 and ~330 basepairs (bp). DNA sequences can be found on GenBank(GU109549–GU110746, HM575429–HM576623, andHM576658). These markers were chosen as they havebecome 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 ratesand contain both protein and non-protein codingregions. DNA was extracted from a portion of a legusing a Qiagen DNeasy Tissue Kit following the manu-facturer’s protocol. Each new specimen used in thisstudy was given an individual number (e.g. sel_001)and has been deposited in the Essig Museum ofEntomology at the University of California, Berkeleyand the California Academy of Sciences. Remaininggenomic DNA is stored at -80 °C in the Gillespieand Roderick Laboratories, University of California,

Figure 2. Map of the study area showing the number of localities per region (the first number), the number of totalspecimens 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 SupportingInformation (Figs S1, S2).

Table 1. Genetic loci and primer pairs used for PCR amplification

CO1 LCO1_1490C1N_2568 5′-GGTCAACAAATCATAAAGATATTG-3′ – Folmer et al. 1994

5′-GCTACAACAATAATAAGTATCATG-3′ – Hedin & Maddison, 200116S–ND112350mod 5′-TTDGNTACCAAGCAGACVGC-3′ – this study13398 5′-CGCCTGTTTAACAAAAACAT-3′ – Simon et al. 1994HistoneH3aF 5′-ATGGCTCGTACCAAGCAGACVGC-3′ – Colgan et al. 1998H3aR 5′-ATATCCTTRGGCATRATRGTGAC-3′ – Colgan et al. 1998

CO1, cytochrome oxidase I; 16S, ribosomal DNA; ND1, NADH dehydrogenase I.



Berkeley. Primer pairs used are given in Table 1. Insome cases, primarily with outgroup taxa, amplifica-tion was difficult and, in such instances, the EpicentreFailSafe PCR kit was used. In the majority of cases,sequence data were obtained for all gene fragments formultiple representatives of each species. In one situa-tion with the species Selenops insularis Keyserling,there was evidence for multiple copies of H3a in somespecimens, thus these sequences were not analysed forthese individuals.

PHYLOGENETIC METHODS

Alignments of the protein-coding loci CO1, ND1 andH3a were performed manually using Mesquite ver.2.5 (Maddison & Maddison, 2008), with the amino-acid translations used as a guide. The 16S datawere aligned using secondary structure based on themodel from Masta (2000). While there were somelength differences between taxa, alignment wasstraightforward.

Data were partitioned by codon position for proteincoding genes, by stems and loops for ribosomal DNAand by gene for both the maximum likelihood andBayesian analyses to improve the fit of the substitu-tion model to the data (Nylander et al., 2004; Brand-ley, Schmitz & Reeder, 2005). The doublet model ofnucleotide substitution was used for the stem regionsof 16S and the tRNA (Schöniger & von Haeseler,1994; Kjer, 2004). Maximum likelihood analyses wereperformed with RAxML ver. 7.0.4 (Stamatakis, 2006)and Bayesian analyses were performed usingMrBayes ver. 3.1.2 (Huelsenbeck & Ronquist, 2001;Ronquist & Huelsenbeck, 2003; Altekar et al., 2004).RAxML is able to analyse partitioned data, but onlyunder the generalised time reversible (GTR) model,thus, while the same partitioning regime was used inboth Bayesian and likelihood analyses, this was themodel that was used in the maximum likelihoodanalyses. To determine the models for each partitionin the Bayesian analysis, MrModeltest ver. 2.3(Nylander, 2004) was used. Models were chosen usingthe Akaike information criterion (AIC; Akaike, 1973;see Posada & Buckley, 2004) and are listed in Table 2.

LIKELIHOOD ANALYSES

RAxML maximum likelihood analyses were con-ducted in a variety of ways following the suggestionsof the author (Stamatakis, 2006). First, one analysiswas conducted which included 893 terminals afteridentical haplotypes were removed. However, to easethe computational strain for more intensive analyses,terminals that were � 0.3% different were removedfrom the analysis (sensu McGuire et al., 2007). Thistruncated data set contained 306 terminals.

The RAxML manual suggests two ways to analysedata – the ‘fast and easy way’ and the ‘hard and slowway’ (Stamatakis, 2006). The fast and easy way wasused to analyse the full data set because of its largesize (~900 terminals and ~2000 bps). The hard andslow way was used to analyse the smaller data setand allows the program to find ‘good’ settings particu-lar to an individual data set. The user’s manual wasfollowed exactly for the analysis of the truncated dataset using the ‘hard and slow’ method. First, fiverandomized maximum parsimony trees were gener-ated and then each tree was inferred using a fixedsetting of ten for the initial rearrangement. Next, thissetting was automatically determined for the samefive starting trees and whichever settings yielded thebest likelihood scores were used for subsequent analy-ses. The second part of the ‘hard and slow’ methodinvolves the number of rate categories. For this, thenumber of rate categories is increased by 15, from 10to 55 for each of the five starting trees, using which-ever setting worked best from the initial rearrange-ment analyses. Finally, ten analyses were run usingthe best settings from the above experiments andbootstraps from 500 iterations were then added to thetree 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 thecluster at the Museum of Vertebrate Zoology, Univer-sity of California, Berkeley, as well as on the CIPRES

Table 2. Partitions used in likelihood and Bayesiananalyses and selected models for each partition used inBayesian analyses

Partition Selected model

16S stems GTR + G + doublet16S half stems GTR + G16S loops GTR + I + GLeucine tRNA stems HKY + G + doubletLeucine tRNA loops HKY + GND1 postion 1 GTR + I + GND1 position 2 GTR + I + GND1 position 3 GTR + GCO1 position 1 GTR + I + GCO1 position 2 GTR + I + GCO1 position 3 GTR + GH3a position 1 GTR + IH3a position 2 JC + IH3a position 3 K80 + G

CO1, cytochrome oxidase I; 16S, ribosomal DNA; ND1,NADH dehydrogenase I.



cluster at the San Diego Supercomputer Center.Despite using the truncated data set and running thejobs in parallel, analyses required months to nearcompletion. Programs were run using the defaultsettings for 40 million generations and, if convergencewas not met, the generations were increased in incre-ments 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 vicari-ance and dispersal. Lagrange uses likelihood modelsto test geographic range evolution and allows changesin dispersal and extinction parameters at differenttimes in the past, allowing the incorporation of exter-nal information such as geological data and dispersalcapabilities. For example, if a land mass did not existat a particular time period, because it had not yetemerged or was inundated, the rate of dispersal to theland mass would be 0 during this time and couldincrease during the time period(s) the land mass wasavailable for colonization. In an area as geologicallycomplex as the Caribbean, there are nearly endlessways to parameterize the models, but simplicity wasmaintained throughout each analysis.

Lagrange requires a tree and a matrix of rangedata for the included taxa. We analysed a truncateddata set, selecting one specimen from each species,along with the outgroups, using a partitioned RAxMLsearch for the best tree. We then pruned the out-groups before conducting the Lagrange analyses tomake the computational load smaller, and becausethe focus of the questions concerns only the ingroup.We ensured the tree had the same basic structure astrees from the more complete analyses and that allrelationships supported in those analyses alsoappeared in this tree.

We divided the range of the Selenopids in Northand Central America into five areas: C (CentralAmerica and Mexico), S (South America), G (GreaterAntilles), N (Northern Lesser Antilles), A (SouthernLesser Antilles). Although certain parts of theseregions were not available for colonization throughoutparticular time periods (i.e. some of the Greater Anti-lles have been emergent longer than others, etc.), wesimply used the maximum times from their firstappearance. We set the age of the root node of the treeto 130 Myr, as it is assumed a split between theingroup, i.e. American selenopids, and the outgroup,i.e. African selenopids, was caused by the separationof Africa from South America. Dating vicarianceevents by the initiation of mid-ocean ridge spreading

is problematic, in that in some cases this upper boundis too old as a result of chance transoceanic inter-change after actual separation. However, the exist-ence of distinct clades of Selenops on the differentcontinents would argue for little genetic exchangebetween the continental land masses (Smith & Peter-son, 2002) and would therefore indicate that it isindeed appropriate to use the separation of Africa andAmerica to date Selenops. Throughout all analyses,we focused on six time periods which correspond tothe availability of land for colonization (Table 3).

The following three analyses consisted of twomodels each, one representing each of three scenarioswith, and without, GAARlandia. The three scenarioswere: (1) a dispersal-based scenario where distancebetween land masses determines the probability ofcolonization; (2) a dispersal-based scenario in whichthe ability to colonize an available land mass is notdependent on distance, thus the colonization of anyone land mass from another is equiprobable; (3) avicariance-based scenario, with little to no over-waterdispersal. This means that colonization of one areafrom another could occur only through connections ofone land mass to another. In some cases, certainareas were never connected to other land masses,such as the Lesser Antilles. In this case, the probabil-ity of dispersal is not set to zero, but rather a very lowprobability, as the presence of the spiders indicatescolonization at some point in the past.

It is possible to set different dispersal probabilitiesfor each direction, so that the probability of movingfrom one region to another can be lower or higherthan in the opposite direction. However, to maintain

Table 3. Correspondence between time slices and geo-graphical ranges defined for models used in Lagrangeanalyses

Time slice Land availability

3.0 Closing of the Isthmus of Panamá5.0 Most recent appearance of Northern

Lesser Antilles12.0 Most recent appearance of Southern

Lesser Antilles33.0 Disappearance of GAARlandia35.0 Appearance of GAARlandia50.0 From 55–50 Mya, a part of Jamaica was

connected to Central America via theNicaraguan Rise

55.0 Time after which land was available inthe Greater Antilles region

130.0 Age of root node, corresponds toseparation of Africa and SouthAmerica



simplicity, bidirectional probabilities were set asequal. Within each model, the only parameterchanged between the two analyses was the probabil-ity of colonization with and without the presence ofGAARlandia. The maximum range size was set to twoareas and the areas G, N and A were excluded fromthe root (> 130 Myr) as they were not available forcolonization at this time (Table 3).

RESULTSSAMPLING

We obtained over 1000 specimens from over 200localities within the area of primary focus for thisstudy. In total, we have 29 out of 41 Caribbean islandspecies, half of the known Mexican and CentralAmerican species and one fifth of the described SouthAmerican species (Appendix).

PHYLOGENETIC ANALYSES

Likelihood analysisThe tree from the analysis of the full data set isshown in Figure 3 and has a likelihood score of-61 544.60. Nodes with bootstrap (BS) values � 70%are considered to be supported. There is no support(BS < 70%) for many basal nodes. Further discussionof the results from this tree is given below wherecompared with trees from the other analyses.

In the analysis of the truncated data set, the bestlikelihood score came from the trial with a fixedsetting of 10 for the initial rearrangement, ratherthan the automatic setting (Table 5). The best likeli-hood from the experiment to determine a good settingfor the number of rate categories occurred when thissetting was at 25 (Table 6). Thus, the initial rear-rangement setting was fixed at 10 (-i 10) and thenumber of rate categories was set to 25 (-c 25). Thebest overall likelihood with these settings fromthe MultTrees analysis came from the second run(Table 7) and the results are shown in Figure 4. Theoverall structure is similar to the tree obtained fromthe analysis of the full data set, in which many basalnodes are not supported, while nodes above these are.This tree is discussed in more detail below.

Bayesian analysisThe analyses were run for 64 million generations (themaximum possible given limits of storage space forour output files). According to the cumulative plotfrom AWTY (Wilgenbusch et al., 2004; Nylander et al.,2008), the run reached convergence near 55 milliongenerations. Because convergence was only reachedvery late in the analysis, the first 90% of trees wereeliminated as burn-in, leaving ~12 000 trees fromwhich to compute a consensus. This tree is shown in

Figure 5 and nodes with posterior probability values� 0.95 are considered to be supported. The branchlengths are longer than in the likelihood analysesand, while a few more basal nodes are supported thanin the likelihood analyses, the overall pattern is thesame. The similarities and differences among allthree trees are discussed below.

Comparison of treesAll three trees are very similar with many of theminor differences not supported. The remainder ofthe basal nodes occurring below the Selenopids ofNorth and Central America is only supported asmonophyletic in the Bayesian analysis. The focal taxaof the study, the Selenopids of North and CentralAmerica, are monophyletic and further subdividedinto a well-supported strictly Caribbean clade (Fig. 6,clade A) and the remaining taxa, supported as a cladein the Bayesian tree only; (Fig. 6, clade B), includingtaxa from the south-western USA, Mexico, CentralAmerica, the Southern Lesser Antilles (SLA) andSouth America. Within this clade B, although basalrelationships are not supported, all analyses supporta southern Caribbean basin clade (Fig. 6, clade C)consisting of taxa from Aruba, Bonaire, Curaçao,Trinidad and Tobago. Selenops n. sp. 5 from Aruba isalways sister to Selenops curazao from Bonaire andCuraçao and, this clade (Fig. 6, clade D), is alwayssister to Selenops willinki from northern SouthAmerica and Tobago + S. geraldinae from Trinidad(Fig. 6, clade E). Also within clade B, another well-supported clade in all analyses consists of Selenopsbanksi, found in Panama and South America, andSelenops micropalpus, found in the Southern LesserAntilles from Dominica to St Vincent and the Grena-dines (Fig. 6, clade F).

There is support for a sister group relationshipbetween the South American taxa + the Central andNorth American taxa in the Bayesian tree only. Boththe Bayesian analysis and the likelihood analysisof the truncated data set support a CentralAmerican + North American clade (Fig. 6, clade G), aswell as one between the widespread Selenops mexica-nus, Selenops gracilis and a new species found only inMexico (Fig. 6, clade H). In the Bayesian tree, S.mexicanus is paraphyletic. There is little support forany other relationships in clade B, other than thespecies from the Selenops debilis group of the south-western USA and Northern Mexico (Fig. 6, clade I).

The Caribbean clade (Fig. 6, clade A) consists onlyof taxa from Caribbean islands and is strongly sup-ported in all analyses, but, again, with little supportfor basal nodes, the exception being the widespreadSelenops lindborgi and its sister species, S. n. sp. 3,which are supported as sister to the rest of theCaribbean taxa (Fig. 6, clade J).



Figure 3. Likelihood tree resulting from the RAxML analysis of the full data set. The map above the tree depicts theCaribbean 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 indicatesthe species is found in Cuba. Selenops radiatus (highlighted in blue) is the type of the genus.



A relationship consisting of the four Jamaicanspecies, four species endemic to Hispaniola + S. insu-laris, from throughout the Greater Antilles, is repre-sented in all three analyses, although not supportedin the full data set (Fig. 6, clade K). The Jamaicanspecies are monophyletic, with well-supported inter-relationships in all analyses (Fig. 6, clade L). Thesister clade, consisting primarily of Hispaniolanendemics (three of which are undescribed), is alsowell supported (Fig. 6, clade M). However, Hispaniolahas several species outside this clade.

Nodes on the branches subtending other major Car-ibbean lineages (clades N, O, P and Q in Fig. 6) areunsupported, although many sister group relation-ships and one small subclade consisting of threeundescribed species from Hispaniola and one from theTurks and Caicos Islands (Fig. 6, clade N) are sup-ported in all analyses.

Lagrange analysesThe results of the Lagrange analyses are given inTable 8. Shown in Figures 7–9 are the maximum like-

lihood reconstructions of range evolution under eachof the six models. The best likelihood score overall(-121.90) was from model 3B, the vicariance-basedmodel that includes GAARlandia (Fig. 9B). In models2A–3B, the best likelihood scores were produced fromthose that included GAARlandia. The first two analy-ses (using models 1A and 1B), which take distancebetween islands into account, produced very similarlikelihood scores and maximum likelihood reconstruc-tions, although the model without GAARlandia hadan insignificantly greater likelihood score. However,in the other two analyses, the differences in likelihoodscores were significant and, in the analyses modelledwith no GAARlandia, there was much more uncer-tainty in the reconstructions (Figs 7–9 – greybranches indicate that alternative reconstructions fallwithin two log-likelihood units of the scenario that isdepicted). Likelihood ratio tests were used to comparenested models and, when scenarios were not nested(e.g. – scenario 2A and scenario 3A), the highestlikelihood score is taken as the best.

DISCUSSION

Unique and shared biogeographic patterns are sum-marized in Table 9.

SOUTHERN CARIBBEAN BASIN

Members of the well-supported Southern CaribbeanBasin clade (Aruba, Bonaire, Curaçao, Trinidad andTobago, Fig. 6, clade C) are never found within thelarger well-supported Caribbean clade (Fig. 6, cladeA). Geological data often suggest a relationshipbetween these southern islands, known as the Aruba–Tobago Belt (Iturralde-Vinent & MacPhee, 1999) and

Table 4. Outgroup taxa used to root trees, collection localities, voucher numbers and location of vouchers

Family Genus and species Locality Voucher number and locality

Ctenidae Vulsor sp. Madagascar, Ranomafana CASENT9024024 – CASCtenidae Phoneutria fera French Guiana, Tresor Nature Reserve CASENT9021738 – CASCtenidae Cupiennius ca.

granadensisFrench Guiana, Emerald Jungle Village CASENT9021735 – CAS

Ctenidae Acanthoctenus sp. Argentina, Parque Nacional Cope, límte NE ARAMR000556 – MACNSparassidae Olios sp. 1 USA, California, Esparto In author’s personal collectionSparassidae Olios sp. 2 México, Baja California, north of Guerrero

NegroIn author’s personal collection

Sparassidae Polybetes pythagoricus Argentina, Buenos Aires Prov., José Mármol Ar 10876 – MACNSparassidae Heteropoda sp. Nepal, near Sauraha In author’s personal collectionSparassidae Heteropoda sp. Tanzania In author’s personal collectionSparassidae Damastes sp. 1 Madagascar, Toliara sel_554 – CASSparassidae Damastes sp. 2 Madagascar, Ambohitantely CASENT9015896 – CAS

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

Table 5. Likelihoods from the ‘hard and slow’ RAxMLanalyses to determine the best initial rearrangementsetting for the data

-ln(L) for initialrearrangement settingfixed at 10

-ln(L) for automaticinitial rearrangementsetting

49 876.238442* 49 880.14629549 881.090042 49 877.34916949 880.995332 49 893.14815249 881.483263 49 879.92431749 879.897717 49 897.163460

*The best score is denoted.



indeed the affinities are not surprising given theproximity of the islands to each other and to theSouth American continent. The amphibian and reptileassemblages on each of these islands are largely con-tinental and also distinct from the primary Caribbeanelements (Hedges, 2006).

In Selenops, this Southern Caribbean clade isapparently not closely related to other Caribbeantaxa, a pattern found in many other groups, includingmammals (Dávalos, 2004) and plants [orchids (Trejo-Torres & Ackerman, 2001)]. However, a contrastingpattern has been found in Anolis lizards in which theSouthern Caribbean Basin taxa show stronger affini-ties with the Antilles (Jackman et al., 1999; Creeret al., 2001); these affinities are hypothesized to havearisen as a result of the Lesser Antilles being muchfurther 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 withseveral insects (Wilder & Hollocher, 2003), Anolis

lizards (Gorman & Atkins, 1969; Jackman et al.,1999; 2002; Creer et al., 2001; Schneider, Losos & deQueiroz, 2001) and Eleutherodactylus frogs (Kaiser,Sharbel & Green, 1994), is that species in the North-ern Lesser Antilles are only distantly related tospecies in the Southern Lesser Antilles. The speciesS. n. sp. 7 is found in the Northern Lesser Antillesfrom Les Saintes northward to Montserrat andAntigua, while the species occurring in the SouthernLesser Antilles, from Dominica south to St Vincentand the Grenadines (at least to Mayreau) is S. micro-palpus. The northern species is nested well within thestrictly Caribbean clade, while S. micropalpus sharesa relationship with S. banksi found from Panamá toPeru to Guyana. The precise location where northernand southern lineages are separated is variable, beingslightly to the south in other lineages. For example, inAnolis, 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 banan-aquit (Seutin et al., 1994), it is between St Vincentand St Lucia. Differences in the location of the bound-ary between northern and southern lineages mayoccur as a result of the timing of colonization of thedifferent groups, which is likely related to the timingof emergence of the individual islands. Interestingly,anoles from the Southern Lesser Antilles, like thespiders, show affinities with Central and SouthAmerican 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 VirginIslands, St Kitts, Nevis, eastern Hispaniola and GreatInagua in the Bahamas, see also Supporting Informa-tion, Fig. S1E–G) and the very narrowly distributed

Table 6. Likelihoods from the ‘hard and slow’ RAxML analyses to determine the best setting for the number of ratecategories for the data

Starting tree

Rate categories = 10 Rate categories = 25 Rate categories = 40 Rate categories = 55

-ln(L) -ln(L) -ln(L) -ln(L)

1 49 876.825172 49 880.146295 49 878.266732 49 881.3494212 49 909.100516 49 877.349169* 49 877.528777 49 880.6687323 49 884.108152 49 893.148152 49 882.377547 49 885.8278444 49 878.378649 49 879.924317 49 884.766631 49 884.6107435 49 885.284097 49 897.163460 49 877.756447 49 891.633712


Table 7. Likelihoods from the MultTrees analyses withthe initial rearrangement setting at 10 and the number ofrate categories set to 25

Tree -ln(L)

1 49 890.3527432 49 875.885667*3 49 877.9636664 49 876.0180675 49 884.5919636 49 881.2619667 49 879.5479328 49 883.4006989 49 897.114073

10 49 879.548262




(Isla Mona and Puerto Rico, see also SupportingInformation, Fig. S1F) S. n. sp. 3 (Fig. 6 clade J). Asimilar pattern is found among Anolis, with PuertoRico endemic Anolis occultus also basal (Jackmanet al., 1999). Likewise, the most basal iguana of thegenus Cyclura is also located on the Puerto Rican

bank (Malone et al., 2000). This pattern, which indi-cates a common origin of Caribbean diversity forthese groups, is in contrast to data from geckos, frogs,colubrid snakes and butterflies, which suggest His-paniola as a centre of diversity (Liebherr, 1988a, andreferences therein).

Figure 4. Likelihood tree resulting from the RAxML analysis of the truncated data set. The map above the tree inFigure 3 depicts the Caribbean islands and the colours correspond to branches in the tree and indicate on which islandthe species is found. Multiple colours along a branch indicate that the species is found on multiple islands. A branchoutlined 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 GreaterAntilles, with areas that may have had some partscontinuously above sea level for many millions ofyears longer than other islands (Iturralde-Vinent &MacPhee, 1999; Iturralde-Vinent & Gahagan, 2002).It is also more isolated than other islands as its lastprobable connection with a land mass was likely withCentral America through the Nicaraguan Rise55 Mya. Our data reflect this isolated history, as

Jamaican species of Selenops form a monophyleticgroup of endemics (Fig. 6, clade L). Monophyly ofJamaican taxa is also present in anoles (Jackmanet al., 1999; Nicholson et al., 2005) and Eleutherodac-tylus frogs (Hedges, 1996a,b). However, affinities ofthe Jamaican clade differ between spider and verte-brate groups: The Jamaican clade of Selenops is sup-ported in the Bayesian and truncated likelihoodanalyses as being sister to a clade of primarily His-paniolan species (Fig. 6, clade K). In contrast, theJamaican clade of Eleutherodactylus frogs is most

Figure 5. Tree resulting from the Bayesian analysis of the truncated data set. The map above the tree in Figure 3 depictsthe 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 indicatesthe species is found in Cuba. Selenops radiatus (highlighted in blue) is the type of the genus.



closely related to species from Cuba, while theJamaican lineages of Anolis lizards (Nicholson et al.,2005) and short-faced bats (Dávalos, 2007) are sisterto clades from the mainland. Overall, Jamaica’shistory has been quite different from that of the otherGreater Antillean islands and its fauna may haveaccumulated via dispersal and in situ speciationrather than vicariance (Buskirk, 1985; Crother &Guyer, 1996).

BIOGEOGRAPHICALLY DERIVED POSITION OF CUBA

Cuba has often been depicted as a basal locality inarea cladograms (Buskirk, 1985; Crother & Guyer,1996). In contrast, although not always supported,Selenops species from Cuba appear not to be basal, atleast based on morphology and our limited molecularsampling. Only one species (Selenops aissus – col-lected from the Bahamas, but that also occurs in

Figure 6. Bayesian tree with species symbols and asterisks indicative of support removed for clarity. The outgroup taxahave also been removed. Letters on the nodes indicate clades discussed in the text and in Table 9.



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

Model -ln(L) Dispersal Extinction

1A – dispersal where distance is important; without GAARlandia 135.5 0.01634 0.00821B – dispersal where distance is important; with GAARlandia 135.9 0.01637 0.0083872A – dispersal where distance is not important; without GAARlandia 144.3 0.02604 0.0084112B – dispersal where distance is not important; with GAARlandia 128.1* 0.02137 0.0080943A – little to no over-water dispersal; without GAARlandia 260.8 0.4647 0.0066293B – little to no over-water dispersal; with GAARlandia 121.9* 0.1189 0.008262

Model 1 is a dispersal-based model in which the distance between land masses is considered.Model 2 is a dispersal-based model in which the distance between land masses is not taken into account.Model 3 is a vicariance-based model in which dispersal probabilities are very low if dispersal must occur over water.The ‘A’ portion of each model was run without GAARlandia, while the ‘B’ portion was run with GAARlandia.The dispersal and extinction values are the maximum likelihoods estimates for the rate of each process and represent themean 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 wheredistance between land masses is taken into account. Single-area ancestral ranges are shown at nodes. Grey branchesindicate that alternative reconstructions fall within two log-likelihood units of the scenario that is depicted. Rangetransitions along branches show sequences of dispersal and extinction events. C, Central America; S, South America; G,Greater Antilles; N, Northern Lesser Antilles; A, Southern Lesser Antilles. (A) without GAARlandia; (B) with GAAR-landia.



Cuba) occurs at the base of an internal clade, while allother sampled species which occur in Cuba (althoughall but one – Selenops submaculosus – were collectedfrom other islands) are nested high within the trees(S. submaculosus, Selenops simius, Selenops inus-laris) and it is inferred based on morphology (S. C.Crews & R. G. Gillespie, unpubl. data) that most ofthe Cuban endemics are closely related to S. simiusand S. submaculosus (Fig. 6, clade O).

GREATER ANTILLES AS A CENTRE OF

SPECIES DIVERSITY

There are two additional patterns that appear to beshared between Selenops and Anolis. In Selenopsspiders, the species from the Northern Lesser Antilles

and St Maarten and Anguilla are nested well withina clade of Hispaniolan animals (Fig. 6, clade O). Thissuggests that S. n. sp. 7 and S. n. sp. 8 colonized theNorthern Lesser Antilles region from Hispaniola andthus support the Greater Antilles as a centre ofspecies diversity via dispersal events in Anolis (Glor,Losos & Larson, 2005). Also, many species of Selenopsin Hispaniola have very small ranges that mirrorthose of many endemic anoles from the Anolis cybotesgroup (Glor et al., 2003), indicating similar patternsof speciation between the two groups.

SPECIES–AREA RELATIONSHIPS

In many taxa as diverse as fungi (Lodge, Baroni &Cantrell, 2002), vertebrates (Ricklefs & Lovette,

Figure 8. Maximum likelihood reconstruction of geographic range evolution under a dispersal-based model wheredistance between land masses is ignored. Single-area ancestral ranges are shown at nodes. Grey branches indicate thatalternative reconstructions fall within two log-likelihood units of the scenario that is depicted. Range transitions alongbranches 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 GreaterAntilles harbour more species than the Lesser Anti-lles. This can be attributed largely to island area(MacArthur & Wilson, 1963) and associated habitatdiversity and age (Losos, 1996; Ricklefs & Berming-ham, 2002, 2008). In Selenops, the same pattern isfound, with larger, older islands (Greater Antilles)having more species than smaller, younger, lesshabitat-diverse islands (Lesser Antilles). In theGreater Antilles there is often a pattern of numberof species in Cuba > Hispaniola > Jamaica > PuertoRico, based on island size. This pattern also prevailsin Selenops. In this genus, 17 species occur in Cubawith 12 endemics (Alayón, 2005), while in Hispaniolathere are at least 16 species with 11 endemics and, inJamaica, at least five species are known, with four

endemics. However, the Bahamas have no knownendemic species of Selenops.

HYPOTHESIS TESTING

In the maximum likelihood analyses of range expan-sion, likelihood ratio tests of scores for the scenariosthat include the existence of the GAARlandia landspan are either equally probable or more favourablethan those that do not. This does not mean thatover-water dispersal has not occurred, but rather thatland bridges hold a stronger signature on the phylog-eny. These results contrast to those for mammals inwhich there was little to no support for a land spanbetween the Greater Antilles and northern SouthAmerica (Dávalos, 2004). Likewise, Hedges and

Figure 9. Maximum likelihood reconstruction of geographic range evolution under a vicariance-based model where thereare very low probabilities of over-water dispersal. Single-area ancestral ranges are shown at nodes. Grey branchesindicate that alternative reconstructions fall within two log-likelihood units of the scenario that is depicted. Rangetransitions along branches show sequences of dispersal and extinction events. C, Central America; S, South America; G,Greater Antilles; N, Northern Lesser Antilles; A, Southern Lesser Antilles. (A) without GAARlandia; (B) with GAAR-landia.



others (Hedges, Hass & Maxon, 1992; Hedges,1996a,b; Hedges & Heinicke, 2007; Heinicke et al.,2007) found that molecular clock estimates of diver-gence times precluded a major role of land bridges inthe origin of Caribbean herpetofauna. The suitabilityDispersal–Extinction–Cladogenesis model of geo-graphic range evolution used here, in which dispersalevents cause range expansion, local extinction eventscause range contraction and the probability of eachkind of event is proportional to the branch length, hasbeen questioned for island fauna, as terminal taxamay be restricted to single islands (Ree & Smith,2008). However, in our models, islands were eithergrouped together or several species were spreadacross multiple islands and thus the model is reason-able in this particular case.

CONCLUSIONS

The current study provides a basis for biogeographiccomparison across different lineages in the Carib-bean. It is one of the most extensive data sets forCaribbean fauna and the most comprehensivemolecular data set of any spider group. While inclu-sion of taxa currently missing from our analyses, andpossibly 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 verydifficult if not impossible to resolve (Degnan & Salter,2005; Kubatko & Degnan, 2007). The results revealseveral patterns common to other disparate taxa, aswell as many unique patterns which warrant furtherstudy. Moreover, the data set provides the ground-work for behavioural, ecological and population-levelstudies similar to lineages such as Anolis lizards(Losos, 2009) and passerine birds (Ricklefs & Ber-mingham, 2007).

ACKNOWLEDGEMENTS

We would like to acknowledge members of S.C.C.’sdissertation committee for their guidance: GeorgeRoderick, Jim McGuire and Charles Griswold. Wewould like to thank the following museums, curatorsand collection managers for specimen loans: AmericanMuseum of Natural History – Norman I. Platnick andLouis Sorkin; Museum of Comparative Zoology –Laura Leibensperger; California Academy of Sciences– Charles Griswold; National Museum of NaturalHistory – Jonathan Coddington; British Museum ofNatural History – Janet Beccaloni; Peabody Musemat Yale – Raymond Pupedis; Essig Museum of Ento-

Table 9. Biogeographic patterns in Caribbean Selenops species

PatternShared withother taxa? Which taxa?

Clades that showparticular patterns

1. Distinct, distantly related SLA andNLA clades

Yes Lygaeid bugs, Carabid beetles,fruitflies, butterflies, Anolis,Eleutherodactylus, bananaquit

F and O

2. Monophyly of Jamaican taxa Yes Anolis, Eleutherodactylus L3. More than one colonization and

diversification in HispaniolaYes Anolis A, J, M, N

4. Patterns of endemism throughout theCaribbean (GA harbour moreendemics than LA)

Yes Fungi, Anolis, birds, carabidbeetles

LA – F,OGA – J,K,L,M,N,P,Q

5. Endemic species in Hispaniola withdistributions that overlap endemics ofother taxa

Yes Anolis N, P, others in clade A

6. Southern Netherlands Antilles form aclade with Trinidad and Tobagoexlcusive of other Caribbean taxa

No – C

7. Location of the split between the NLAand SLA is between Dominica andLes Saintes, Guadeloupe

No – SLA – FNLA – O

8. Cuban species are not basal in theCaribbean clade

No – Q, S. aissus

If a pattern is shared with other taxa, the taxa are noted. Clades referenced are those that display the patterns mentionedhere and are depicted in Figure 6.GA, Greater Antilles; LA, Lesser Antilles; NLA, Northern Lesser Antilles, SLA, Southern Lesser Antilles.



mology – Cheryl Barr; Museo Nacional de HistoraNatural, Santo Domingo – Sardis Medrano Cabral,Carmelo Nuñez. We would also like to thank JimMcGuire for use of the MVZ cluster and Mark Millerand Lucie Chan for use of the SGE cluster and theCIPRES portal and the San Diego SupercomputerCenter. We are extremely grateful to Richard Ree forquickly responding and helping us to understand andimplement Lagrange. We would also like to thankMatt Brandley for phylogenetic methodology discus-sions. We are grateful to all of the many people thataided us in obtaining permits and collecting: KelvinGuerrero, Denia Veloz, Eladio Fernandez, AlbertoPuente-Rolón, Beverly Mae Nisbeth, Adriel Thibou,Germain George, Brian Riggs, Brian Manco, Marga-ret Jones, Renata Platenberg, Chris Niebuhr, AbelPérez-González, G. B. Edwards, Oscar Francke, Ale-jandro Mondragon, Mark da Silva, Facundo Franken,Roy Croes, Gijs Van Hoorn, Adolphe O. Debrot, MarkVermeij, Fred the Abaco Caveman, Raveen Gibson,Daniel Palmer, Jim Starrett, Marshal Hedin, NicoleVanderSal, Sean Schoville, Luke Mahler, Uri García,Beto Mendoza, Adrian Nieto Montes de Oca, RebeccaDuncan, Pierre Paquin, Matthew Cottam, Jan denDulk, Joey Slowik, Nicole Esteban, Arturo Herrera,Nancy Bottomley, Inilek Wilmot, Lauren Esposito,Stephen Touissant, Arlington James, FerdinandTripoli, Daniel Memia Zolo, Nouree-Yvon, MartínRamírez, Mark Harvey, Volker Framenau, JeremyMiller, Hannah Wood, Yuri Marusik, CarolineChaboo, Cheryl Barr, Bill Shepard, Akio Tanikawa, C.J. Hayden, Aaron Abdel, Dan Warren. Finally, wewould like to acknowledge Matjaz Kunter and ananonymous reviewer for their straightforward andconstructive reviews of this paper. Funding was pro-vided by the Schlinger Foundation, with additionalsupport from the Margaret C. Walker Fund.

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SUPPORTING INFORMATION

Additional Supporting Information may be found in the online version of this article:

Figure S1. Expansion of the boxed area in Figure 1 of the main text, divided into regions depicted in theFigure S2A–I, showing the detailed locality data.Figure S2. Collecting localities from the Caribbean region, including most islands and several sites throughoutMexico, Central America and the South American mainland.

Please note: Wiley-Blackwell are not responsible for the content or functionality of any supporting materialssupplied by the authors. Any queries (other than missing material) should be directed to the correspondingauthor for the article.



APPENDIX

Collecting localities and voucher numbers of all animals used in this study. Locality numbers refer to numbersin the Supporting Information (Figs S1 and S2).

Localitynumber Species Collection information Voucher numbers

1 Selenops n. sp. 5 Aruba: Bringamosa, house of Roy Croes,12°39.547′N, 69°58.077′W, 14.X.2004,SCC04_041

sel_069, sel_070

2 Selenops n. sp. 5 Aruba: Luela, shooting range, 12°29.023′N,69°57.778′W, 16.X.2004, SCC04_044

sel_072

3 Selenops n. sp. 5 Aruba:N. sp. 5 National Park, near hill GranTonel and Valley Rooi Coashati, 12°29.356′N,69°55.461′W, 16.X.2004, SCC04_045

sel_068

4 Selenops curazao Curaçao: Carmabi Institute, 12°07.351′N,68°58.132′W, 7.X.2004, SCC04_026

sel_047, sel_058, sel_217

5 Selenops curazao Curaçao: Girouette Plantation, E of SchottegatHarbour, house of A. DeBrot, 12°09′03.20″N,68°54′56.35″W, 9.X.2004, SCC04_032

sel_048, sel_049, sel_050, sel_051,sel_052

6 Selenops curazao Bonaire: Sabadaco, near cave, 12°11.587′N,68°17.765′W, 11.X.2004, SCC04_033

sel_053

7 Selenops curazao Bonaire: Altamira Ungu, 12°13.949′N,68°20.703′W, 12.X.2004, SCC04_038

sel_059, sel_060, sel_082

8 Selenops curazao Bonaire: Nort di Saliña, Kaya Otomac,12°10′55.92″N, 68°16′28.39″W, 11.X.2004,SCC04_039

sel_054, sel_056, sel_057, sel_061,sel_062, sel_063, sel_064, sel_065

9 Selenops willinki Trinidad and Tobago: Little Tobago,11°18′03.7″N, 60°30′11.1″W, 16.VII.2005,SCC05_041

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

10 Selenopsgeraldinae

Trinidad and Tobago: St George Co., PointGourde Road near Trinidad Military Base,near Chaguaramas off Chaguaramas MainRoad, 10°40′47.1″N, 61°37′30.9″W,11.VII.2005, SCC05_035

sel, 218, sel_219, sel_220, sel_221,sel_222, sel_223


Trinidad and Tobago: Gaspar Grande Islandsouth of Chaguaramas, trails around island,10°39′46.5″N, 61°38′58.0″W, 12.VII.2005,SCC05_037

sel_224, sel_225, sel_226, sel_227,sel_228, sel_229, sel_237,sel_241, sel_257


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

sel_239


Trinidad and Tobago: Chacachacare Island,10°41′24.2″N, 61°44′53.7″W, 13.VII.2005,SCC05_040

sel_246

15 Selenopshebraicus

Argentina: Parque Nacional Chaco, Senderopeatonal. 25.II.2004.

MACN-Ar#12782

16 Selenops occultus Brazil: São Paulo, Universidade São Paulo,23°34′27.96″S, 46°40′21.29″W

sel_995



APPENDIX Continued


17 Selenops occultus Brazil: Rio de Janeiro, Sierra da Carioca,22°57′52.22″S, 43°16′31.34″W

sel_284

18 Selenops occultus Brazil: Rio de Janeiro, Pao de Açucar,22°55′22.05″S, 43°09′33.11″W

sel_283

18 Selenopsmelanurus

Brazil: Rio de Janeiro, Pao de Açucar,22°55′22.05″S, 43°09′33.11″W

sel_277, sel_278, sel_279, sel_280,sel_281, sel_282

19 Selenopsmicropalpus

St Vincent and the Grenadines: Young Island,South of Villa Beach, 13°07.895′N,61°12.142′W, 24.X.2004, SCC04_053

sel_091, sel_113


St Vincent and the Grenadines: King’s HillForest Reserve, 13°08.825′N, 61°10.021′W,27.X.2004, SCC04_055

sel_088, sel_090, sel_092, sel_093,sel_094, sel_095, sel_112


St Lucia: Vieux Fort, on hill above airport,13°44′20.2″N, 60°56′40.8″W, 13.III.2007

sel_832, sel_833, sel_834


St Lucia: Anse la Ray, Ti-Kaye, 13°55′29.6″N,61°02′41.3″W, 13.III.2007, SCC07_046

sel_820, sel_821, sel_822, sel_823


St Lucia: Dennery: south of Dennery on eastcoast road, eastern nature trail, heritagetourism site, 13°53′50.8″N, 60°52′51.2″W,13.III.2007, SCC07_047



St Lucia: Gros Islet, Pigeon Island, on top ofhill, 14°05′31.3″N, 60°57′03.8″W, 12.III.2007,SCC07_044

sel_811, sel_812, sel_813, sel_814


St Lucia: Gros Islet, Beausejour, past cricketsponsor’s office, 14°04′43.1″N, 60°56′31.1″W,12.III.2007, SCC07_045



Martinique: Le Diamant, Grand Anse duDiamant, 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

sel_800, sel_801, sel_802, sel_803


Dominica: Roseau: Botanical Park, top of hillnear shrine, 15°17.998′N, 61°22.754′W,1.XI.2004, SCC04_059



Dominica: Jimmit-Warner, on top of hill withcellular phone antenna, 15°22.690′N,061°24.003′W, 2.XI.2004, SCC04_061

sel_102, sel_103, sel_104, sel_105


Dominica: Cabrits National Park, near trail tofort, 15°35.049′N, 61°28.371′W, 2.XI.2004,SCC04_060


32 Selenops n. sp. 7 Guadeloupe: Les Saintes, top of Le Chameau,15°51′28.1″N, 61°35′39.8″W, 6.III.2007,SCC07_039

sel_783, sel_784, sel_785

33 Selenops n. sp. 7 Guadeloupe: Basse-Terre, Parc Archélogiquedes Roches Gravées, near Trois-Rivieres,15°58.394′N, 61°38.347′W, 10.XI.2004,SCC04_064

sel_114



APPENDIX Continued


34 Selenops n. sp. 7 Guadeloupe: Basse-Terre, Trois Rivieres, end ofTrail Sentier de l′Acomat off Rue NelsonMandela, 15°58′03.0″N, 61°37′50.1″W,5.III.2007, SCC04_038

sel_778, sel_779, sel_780, sel_781

35 Selenops n. sp. 7 Guadeloupe: Basse-Terre, near Vieux Fort onthe D6, along road at Forêt Domaniale duLitoral, 15°57.943′N, 061°42.517′W,11.XI.2004, SCC04_065

sel_115

36 Selenops n. sp. 7 Guadeloupe: Gran-Terre, Pointe du Chateaux,16°14′51.6″N, 61°11′02.6″W, 7.III.2007,SCC07_040


37 Selenops n. sp. 7 Montserrat: Jack Boy Hill, 17°46′02.1″N,62°10′17.0″W, 2.III.2007, SCC07_035

sel_758, sel_759, sel_760, sel_761,sel_762, sel_763, sel_764,sel_765, sel_766, sel_767, sel_768

38 Selenops n. sp. 7 Montserrat: Sweet Water Ghaut, 16°47′07.2″N,62°10′59.8″W, 2.III.2007, SCC07_036

sel_769

39 Selenops n. sp. 7 Montserrat: Silver Hills, north side of SilverHill, 16°48′41.3″N, 62°11′28.7″W, 3.III.2007,SCC07_037


40 Selenops n. sp. 7 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

sel_754, sel_755, sel_756, sel_757

41 Selenops n. sp. 7 Antigua: Indian Town, east of Veranda Resort,17°05′50.2″N, 61°40′53.0″W, 27.II.2007,SCC07_033

sel_753

42 Selenopslindborgi

St Kitts and Nevis: Nevis, Tamarind Bay,Galliput Restaurant, 17°09′48.53″N,62°37′50.02″W, 23.II.2007, SCC07_030

sel_740, sel_741, sel_742


St Kitts and Nevis: Nevis, Round Hill entranceto 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

sel_750, sel_751, sel_752


St Kitts and Nevis: St Kitts, Sand Bank Bay,17°14′59.1″N, 62°38′40.8″W, 24.II.2007,SCC07_031


46 Selenops n. sp. 8 Saba: Giles Quarter Trail, 17°36′54.36″N,63°14′35.52″W, 12.III.2008

sel_1021, sel_1022

47 Selenops n. sp. 8 St Maarten: Mullet Bay, abandoned MulletBay Resort, 18°02′48.0″N, 63°027′29.7″W,21.III.2007, SCC07_027

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

48 Selenops n. sp. 8 St Maarten: Emilio Wilson Estate and Park,18°02′32.7″N, 63°03′53.1″W, 20.II.2007,SCC07_026


49 Selenops n. sp. 8 St Maarten: Upper Princess Quarter,18°01′48.0″N, 63°02′08.0″W, 21.III.2007,SCC07_028

sel_731, sel_732



APPENDIX Continued


50 Selenops n. sp. 8 St Maarten: south-east side of island, trailfrom 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


51 Selenopsmexicanus

St Maarten: Philipsburg, Front Street, nearentrance to cruise ship dock, 18°00.906′N,63°02.587′W, 12.XI.2004, SCC04_067

sel_117, sel_118

52 Selenops n. sp. 8 Anguilla: Shoal Bay West, 18°09′52.8″N,63°09′21.3″W, 13.II.2007, SCC07_016

sel_697

53 Selenops n. sp. 8 Anguilla: The Cove, 18°10′14.1″N,63°07′52.6″W, 13.II.2007, SCC07_017

sel_703, sel_704

54 Selenops n. sp. 8 Anguilla: Long Bay, Long Bay Beach,18°11′29.3″N, 63°07′49.7″W, 13.II.2007,SCC07_015


55 Selenops n. sp. 8 Anguilla: Blowing Point, 18°10′18.0″N,63°05′28.7″W, 13.II.2007, SCC07_018

sel_705, sel_706, sel_707

56 Selenops n. sp. 8 Anguilla: Windward Point, 18°16′18.2″N,62°58′05.3″W, 12.II.2007, SCC07_014

sel_696


USVI: St Croix: Fredericksted, Sprat HallBeach, Rte.63, 17°44′09.8″N, 64°53′24.0″W,14.VI.2006, SCC06_056

sel_497, sel_498, sel_499, sel_500


USVI: St Croix: Sprat Hall Hill off West ShoreRoad, first right after subtracking station,17°44′38.4″N, 64°53′22.3″W,14.VI.2006SCC06_055

sel_494, sel_495, sel_496


USVI: St Croix: Butler Bay, West Shore Road,17°45′49.7″N, 64°52′58.8″W, 14.VI.2006,SCC06_054

sel_490, sel_491, sel_492, sel_493


USVI: St Croix: intersecting road from CrequeDam to Mahogany Road (intersects MountVictory Camp) 17°44′27.1″N, 64°51′25.4″W,14.VI.2006, SCC06_057

sel_501


USVI: St Croix: Radio Telescope Station, eastisland, 17°45.398′N, 64°35.045′W, 18.XI.2004,SCC04_072

sel_527

62 SelenopslindborgiSelenopsinsularis

Puerto Rico: Vieques: Laguna Kiani,18°07′02.2″N, 65°33′41.4″W, 19.VI.2006,SCC06_065



Puerto Rico: Vieques: Ruinas Central PlayaGrande, 18°05′43.2″N, 65°31′13.2″W,19.VI.2006, SCC06_064



Puerto Rico: Vieques: Cano Hondo, Cerca dePuerto Mosquito, 18°06′11.0″N, 65°27′05.5″W,19.VI.2006, SCC06_061



Puerto Rico: Vieques: Refugio Nacional de VidaSilvestre, road to Playa Caracas, LagunaPuerto Ferro, 18°06′24″N, 65°25′25.8″W,19.VI.2006, SCC06_063

sel_513, sel_514, sel_515



APPENDIX Continued



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

sel_486, sel_487, sel_488, sel_489


USVI: St Thomas, Estate Perserverance,Perserverance Bay Trail, 18°21.463′N,64°59.753′W, 22.X.2004, SCC04_050

sel_076


USVI: St Thomas, St Peter, house of R.Platenberg, 18°21′22.17″N, 64°56′49.53″W,23.X.2004, SCC04_075

sel_131


USVI: St Thomas, Magen’s Bay Trail,18°21.350′N, 64°55.231′W, 22.X.2004 and9.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

sel_123, sel_132, sel_133, sel_134


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

sel_140, sel_141, sel_142, sel_157


USVI: St John, Cinnamon Bay Loop Trail,18°21.226′N, 64°45.259′W, 16-17.XI.2004,SCC04_070

sel_126, sel_127


USVI: St John, Leinster Bay Trail,18°21.825′N, 64°43.743′W, 16.XI.2004,SCC04_069

sel_136, sel_137, sel_138, sel_139


BVI: Tortola: vic. Sage Mountain,18°24′46.44″N, 64°39′18.43″W, 20.X.2004,SCC04_049

sel_078


BVI: Guana Island: north side near beachhouse, 18°28.793′N, 64°34.473′W, 18.X.2004,SCC04_047

sel_071, sel_081, sel_085, sel_089


BVI: Guana Island: south side near salt pond,18°28.619′N, 64°34.475′W, 18.X.2004,SCC04_046

sel_079, sel_084, sel_086, sel_087


BVI: Virgin Gorda, lower trail up Gorda Peak,18°28.774′N, 64°24.210′W, 19.X.2004,SCC04_048


80 Selenopssubmaculosus

Cuba: Sierra de Mesa, Pinar del Rio sel_276

81 Selenops simius Cayman Islands: Grand Cayman, QueenElizabeth II Botanic gardens, storage facilitysouth of main building, 19°19.055′N,81°09.527′W, 30.IX.2004, SCC04_021

sel_046, sel_066, sel_067

82 Selenops simius Cayman Islands: Grand Cayman, QueenElizabeth II Botanic gardens, tree trailbehind iguanas, 19°19.042′N, 81°10.081′W,2.X.2004, SCC04_022

sel_080



APPENDIX Continued


83 Selenops simius Cayman Islands: Little Cayman, on roadacross street from Pirate’s Point Resort,19°39.754′N, 80°06.032′W, 3.X.2004,SCC04_023

sel_022

84 Selenops simius Cayman Islands: Cayman Brac, National TrustHouse off West End Road, 19°42.019′N,79°52.084′W, 3.X.2004, SCC04_025


85 Selenopscandidus

Jamaica: Westmoreland Paris, near New Hopeon road toward Savanna-la-Mar,18°14′55.4″N, 78°14′41.0″W, 29.V.2006,SCC06_027

sel_362

86 Selenopscandidus

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

sel_357, sel_358, sel_359, sel_360

87 Selenopscandidus

Jamaica: Clarendon Parish, off road toLluidasvale, 18°07′50.8″N, 77°10′05.0″W, 31May 2006, SCC06_028

sel_363, sel_364

88 Selenops n. sp.17

Jamaica: St Catherine Parish, Hellshire Hills,A2 Depression, 17°51′59.3″N, 76°57′54.0″W,3.VI.2006, SCC06_031


89 Selenopscandidus

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


90 Selenopscandidus

Jamaica: St Andrew Parish, Castleton BotanicGardens, 18°10′20.3″N, 76°49′27.6″W,27.V.2006, SCC06_022

sel_350, sel_351

91 Selenopscandidus

Jamaica: St Andrew Parish, Hermitage DamRoad, 2–6 km from junction with Stony Hill,18°04′25.4″N, 76°47′01.3″W, 5.VI.2006,SCC06_033

sel_385

92 Selenopspetrunkevitchi

Jamaica: St Thomas Parish, Blue MountainsNational Park, Whitfield Hall, 18°02′54.8″N,76°37′03.7″W, 1.VI.2006, SCC06_029

sel_365, sel_366, sel_367, sel_368,sel_369, sel_370, sel_371,sel_372, sel_373, sel_374,sel_375, sel_376


Jamaica: St Thomas Parish, near 12 mile BullBay, on left side of road heading east,17°55′32.5″N, 76°38′31.0″W, 5.VI.2006,SCC06_034


94 SelenopsinsularisSelenopspensilis

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 RioMulito, 18°13.895′N, 71°45.190′W,25.XI.2004, SCC04_082

sel_156

96 Selenopsmarcanoi

Hispaniola: Dominican Republic: Prov.Pedernales, Rio Mulito (El Banano),18°09.165′N, 071°45.388′W, 25.XI.2004,SCC04_081




APPENDIX Continued


97 Selenops phaselus Hispaniola: Dominican Republic: Prov.Pedernales, Parque Nacional Sierra deBaoruco, Las Abejas, 18°08.804′N,71°37.164′W, 24-Nov-04 and 10-Oct-06,SCC04_077 and SCC06_072


98 Selenops phaselus Hispaniola: Dominican Republic: Prov.Pedernales, 26 km north of Cabo Rojo, Sierrade Baoruco, 18°06.490′N, 71°37.316′W,24.XI.2004, SCC04_078

sel_148, sel_159, sel_215


Hispaniola: Dominican Republic: Prov.Pedernales, 13–14 km north of Cabo Rojo onCarretera ALCOA, 18°01.962′N, 71°38.748′W,24.XI.2004, SCC04_079


100 Selenopsinsularis

Hispaniola: Dominican Republic: Prov.Pedernales, Parque Jaragua, VII.2006

sel_586, sel_589

101 Selenops n. sp.14Selenops n. sp.15

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 MamaCocoño, 25.X.2003, SCC03_021

sel_018


Hispaniola: Dominican Republic, Prov.Pedernales, Laguna N. sp. 15, El Cajuil,9.X.2006, SCC06_070


104 Selenopsinsularis s. n.sp. 1

Hispaniola: Dominican Republic: Prov.Barahona, Carretera Higuero-Polo,26.XI.2004, SCC04_083


105 Selenops phaselusSelenops n. sp.13

Hispaniola: Dominican Republic: Prov.Barahona, Polo Coffee Plantation, July 2006



Hispaniola: Dominican Republic: Prov.Barahona, Barahona, CoralSol Resort, SanRafael 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


Hispaniola: Dominican Republic: Prov. Peravia,Bani, road from Bani to Manaclar, past LaLaguna, 18°21.343′N, 70°21.077′W,27.XI.2004, SCC04_086

sel_175


Hispaniola: Dominican Republic: Prov. Peravia,Bani, Rio Nizao, 18°16.915′N, 70°12.101′W,27.XI.2004, SCC04_087

sel_176, sel_177, sel_178, sel_179


Hispaniola: Dominican Republic: Prov. SanCristóbal, Engombe, farm of autonomousuniversity, 18°27.360′N, 70°00.306′W,27.X.2003, SCC03_023



Hispaniola: Dominican Republic: Prov. LaAltagracia, Parque del Este, Guaraguao,18°19.968′N, 68°48.709′W, 30.XI.2004,SCC04_090




APPENDIX Continued



Hispaniola: Dominican Republic: Prov. LaAltagracia, Parque del Este, Boca de Yuma,18°21.875′N, 68°37.080′W, 29-30.XI.2004,SCC04_089


112 SelenopslindborgiSelenopsinsularisSelenops n. sp.9

Hispaniola: Domnican Republic: Prov. LaAltagracia, Punta Cana Resort,18°30′55.53″N, 68°22′28.73″W, 5-9.VII.2006,SCC04_066



Hispaniola: Dominican Republic: Prov: HatoMajor, Los Haitises, 1 km south of El Valleynear Hato Mayor, 18°58′24.01″N,69°22′34.58″W, VI.2006

sel_555


Hispaniola: Dominican Republic: Prov.Samaná, Las Terrenas, 19°19′28.88″N,69°32′50.77″W, VI.2006


115 Selenops n. sp. 2 Hispaniola: Dominican Republic: Prov. LaVega, Constanza, Alto Cerro Hotel,18°54′14.16″N, 70°44′41.14″W, VI.2006

sel_546


Hispaniola: Dominican Republic: Prov. PuertoPlata, Sosua, VI.2006

sel_578

117 SelenopsinsularisSelenops n. sp.12

Hispaniola: Dominican Republic: Prov. PuertoPlata, 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, rangerstation at trail to Loma del Oro,19°12′05.2″N, 71°00′04.8″W, 13.X.2006,SCC06_074

sel_639

120 SelenopsinsularisSelenops n. sp.10

Hispaniola: Dominican Republic, Prov. MontiCristi, Monte Cristi, El Morro,19°53′42.44″N, 71°39′14.48″W, 8.X.2006,SCC06_069


121 SelenopsinsularisSelenopspensilisSelenops bani

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

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

122 Selenops phaselus Hispaniola: Haiti: Kenskoff, Belot-Montcel,18°27′11.3″N, 72°21′06.4″W, 20-21.X.2006,SCC06_076


123 Selenops n. sp. 3 Puerto Rico: Isla Mona: Bajura de Empalme,18°06′25.07″N, 67°53′10.41″W

sel_846, sel_847


Puerto Rico: Isla Mona: Sardinera,18°05′46.76″N, 67°56′12.35″W

sel_838, sel_839, sel_840, sel_841,sel _842, sel_843, sel_844,sel_845



APPENDIX Continued


125 Selenops n. sp. 3 Puerto Rico: Isla Mona: Camino de los Cobros,18°04′02.68″N, 67°52′45.75″W, VII.2006

sel_502

126 Selenops n. sp. 3SelenopsinsularisSelenopslindborgi

Puerto Rico: Maricao, Bosque Estatl deMaricao, 18°08′51.2″N, 66°59′35.0″W,10.VI.2006, SCC06_045



Puerto Rico: Susua State Forest, SabanaGrande, 18°04′15.0″N, 66°54′31.6″W,10.VI.2006, SCC06_046



Puerto Rico: Quebradillas, Merendero deGuajataca, 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, Carretera657, km 1.9, 18°25′15.9″N, 66°40′35.2″W,7.VI.2006, SCC06_035



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, Carretera146, km 16.3 Interior Camino MaximoNuñez, Sector Los Gonzalez, 18°18′33.8″N,66°32′42.8″W, 15.VI.2006, SCC06_060

sel_503


Puerto Rico: Coamo, Baños de Coamo,18°02′19.4″N, 66°22′27.0″W, 11.VI.2006,SCC06_048

sel_462, sel_463


Puerto Rico: Salina, Reserva Jobos, ParqueJagueys, 17°57′13.9″N, 66°15′03.5″W,11.VI.2006, SCC06_049

sel_464

134 Selenopsinsularus

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

sel_461


Puerto Rico: Manuabo, Mariani Creek,18°00′29.7″N, 65°52′17.0″W, 11.VI.2006,SCC06_050

sel_465, sel_466, sel_467, sel_468


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_416, sel_417, sel_418



APPENDIX Continued



Puerto Rico: Fajardo, Seven Seas Public Beach,18°22′03.7″N, 65°38′04.9″W, 8.VI.2006,SCC06_039



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 atMorgan’s Bluff, 25°10′30.1″N, 78°01′26.2″W,13.V.2006, SCC06_003

sel_309, sel_310, sel_311, sel_666


Bahamas: Andros Island: International FieldStation, 24°53′51.1″N, 77°55′50.1″W,12.V.2006, SCC06_001

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


Bahamas: Pigeon Cay, near IFS on AndrosIsland, 24°52′54.4″N, 77°53′53.5″W,13.V.2006, SCC06_004

sel_312


Bahamas: Andros Island: Cargill Creek,24°30′00.37″N, 77°43′15.61″W, 13.V.2006

sel_668

145 Selenops aissus Bahamas: Abaco: Ralph’s Chimney off Queen’s(Abaco) Highway, 26°14′58.2N, 77°11′25.4″W,14.V.2006, SCC06_006

sel_315

146 Selenops aissus Bahamas: Abaco: Abaco National Park,26°03′44.0″N, 77°12′46.2″W, 14.V.2006,SCC06_005

sel_313


Bahamas: Great Exuma: Bahamas SoundSubdivision near old airport, 23°27′56.0″N,75°46′24.8″W, 18.V.2006, SCC06_011

sel_332

148 Selenops aissus Bahamas: Great Exuma: Regatta Point,23°30′24.7″N, 75°45′58.0″W, 18.V.2006,SCC06_009


149 Selenops aissus Bahamas: Stocking Island, near Great Exuma,23°32′08.9″N, 75°46′29.6″W, 18.V.2006,SCC06_010


150 Selenops aissus 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

sel_316


Bahamas: Great Inagua: Old Aerostat Base,21°06.06.7″N, 73°39′01.9″W, 16.V.2006,SCC06_228

sel_317, sel_318

153 Selenops n. sp. 11 Turks and Caicos: Providenciales, North-WestPoint Pond Nature Reserve, 21°50′32.1″N,72°19′43.7″W, 8.II.2007, SCC07_010 andSCC07_011

sel_689, sel_690, sel_691, sel_692



APPENDIX Continued


154 Selenops n. sp. 11 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


155 Selenops n. sp. 11 Turks and Caicos: Providenciales, The Bight,21°47′00.6″N, 72°13′06.4″W, 22.V.2006,SCC06_015

sel_344, sel_345

156 Selenops n. sp. 11 Turks and Caicos: North Caicos, Wade’s GreenPlantation, 21°55′13.36″N, 72°01′12.45″W,2–3.II.2007, SCC07_001 and SCC07_005


157 Selenops n. sp. 11 Turks and Caicos: Middle Caicos, Garden PondField Road, 21°48′24.76″N, 71°45′42.78″W,3.II.2007, SCC07_004

sel_676, sel_677, sel_678, sel_679

158 Selenops n. sp. 11 Turks and Caicos: Providenciales, South ViewDrive off of Leeward Highway, 21°46′45.7″N,72°13′45.4″W, 10.II.2007, SCC07_012

sel_693

159 Selenops debilsgp. species 1

USA: California, San Diego Co., Upper OtayRiver Valley, 32°59′50.33″N, 116°19′09.27″W,19.V.2003

sel_002


USA: California, San Diego Co., Jamul, LyonsValley, north of Lyons Peak, 32°44′03.93″N,116°53′51.23″W, 29.VII.2007

sel_837


USA: California, San Diego Co., Anza BorregoDesert State Park, Carrizo Palm Grove,32°44′31.29″N, 116°12′51.11″W

sel_021, sel_214


Mexico: Baja California Sur, 2 km east ofBallena between San Ignacio and SanJuanico, 26°27′10.60″N, 111°34′53.39″W

sel_210, sel_213


Mexico: Baja California Sur, 3 km west of VillaInsurgentes on road to San Miguel deComundu, 25°16′50.46″N, 111°50′03.75″W

sel_212


Mexico: Baja California Sur, Cuevas Pintas,26°01′38.63″N, 111°30′24.45’W

sel_009


USA: Arizona, Coconino Co., Monument Trail,flat near archaeological site, 36°25.309′N,112°27.483′W, 13.VIII.2004, SCC04_013

sel_211


USA: Arizona: Santa Cruz Co., MaderaCanyon, Mt Wrightson Trail, 31°43′06.86’N,110°52′22.45’W

sel_264, sel_270, sel_271, sel_272


Mexico: Sonora, Sonoran Highway on road toYecora, 28°23′15.01’N, 108°55′41.42’W

sel_851


USA: Texas: Val Verde Co., Seminole Canyon,Highway 90, under bridge, 29°42′21.12’N,101°18′28.48’W

sel_208


Mexico: Hidalgo, Villa Flores, Ejido ′El Rayo’,24°14′1.00’N, 99°1′4.50’W, 2.XI.2007

sel_1008

170 Selenops abyssus Mexico: Colima, Manzanillo, MunicipioManzanillo, 1.2 to 1.4 km east La Central,19°8′56.4’N, 104°25′35.3’W, X.2005

sel_1004

171 Selenops abyssus Mexico: Colima, Municipio Ixtlahuacan,Tamala, 19°5′2.13’N, 103°47′26.21″

sel_1013



APPENDIX Continued


172 Selenops abyssus Mexico: Michoacan, Municipio Coalcomán,Coalcomán, 18°24′12.81″N, 103°07′58.82″W,X.1005

sel_1006, sel_1012

173 Selenops n. sp. 4 Mexico: Morelos, Cuernavaca, ColoniaChamilpa, Instituto de Biotecnología,UNAM, 18°55′51.86″N, 99°14′16.60″W

sel_1010

174 Selenops gracilis Mexico: Guerrero, Arcelia, Campo Morado,17°34′60.00″N, 100°4′60.00″W

sel_1014, sel_1015

175 Selenopsnigromaculatus?This is animmaturespecimen, butis found nearthe typelocality of S.nigromaculatus,hence the ‘?’

Mexico: Guerrero, Omiltemi, Chilpancingo,Camino al Omiltemi, 17°33′3.76″N,99°30′21.83″W

sel_1007

176 Selenops n. sp. 4 Mexico: Puebla, Municipio Zapotitlan de lasSalinas, Cerro el Pajarito, 18°22′48.3″N,97°30′26.9″W, V.2005

sel_1002, sel_1005


Mexico: Chiapas, Tuxtla-Gutierrez, CañondeSumidero, 16°49′41.46″N, 93°6′22.22″W

1016, 1017


Mexico: Chiapas, Berriozabal, dirt road fromEfrain A. Gutierrez, approx. 8 km northBerriozabal, 16°52′27.45″N, 93°17′28.20″W

sel_848


Mexico: Chiapas, Pueblo Nueva Solistahuacan,17°11.550′N, 92°54.875′W, 17.IX.2004,SCC04_018

sel_043


Mexico: Chiapas, La Reforma, Municipio LaConcordia, 15°54.212′N, 92°40.157′W,18.IX.2004, SCC04_018b

sel_044

181 SelenopsmexicanusSelenops n. sp.19

Mexico: Chiapas, Huixtla, Las Golindrinas,15°25.747′N, 92°39.270′W, 23.IX.2004,SCC04_020

sel_031, sel_034, sel_035, sel_036,sel_037, sel_038, sel_039,sel_040, sel_041, sel_045,sel_1011


Mexico: Chiapas, Municipio Motozintla deMendoza, 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

sel_849


Mexico: Veracruz, Municipio Tamalin, ElMamey, 21°31′13.63″N, 97°38′31.75″W

sel_1018, sel_1019, sel_1020


Guatemala: Petén, Sta Elena de la Cruz,Colonia del Bosque, near Flores, CuevaActun Kan, 16°54′10.9″N, 89°53′44.3″W,1.I.2008, SCC08_001

sel_865, sel_866, sel_867, sel_868

186 Selenopsbifurcatus

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




APPENDIX Continued



El Salvador: Dep. Chaletenango, Chaletenangooff Carretera Troncal del Norte, Hotel Maya,14°16′29.1″N, 89°08′32.7″W, 5-6.I.2008,SCC08_009

sel_927


El Salvador: Dep. Chaletenango, Mun.Chaletenango, La Cueva del Corridor,6.I.2008, SCC08_010


189 SelenopsbifurcatusSelenopsmexicanus

El Salvador: Dep. Chaletenango, Chaletenango,Municipio Tejutto, Rest. Eucalyptos,14°12′20.4″N, 89°06′43.9″W, 5.I.2008,SCC08_008

sel_921, sel_922, sel_923, sel_924


El Salvador: San Salvador, Museo Nacional deHistoria Natural grounds, 13°40′23.4″N,89°11′53.6″W, 3.I.2008, SCC08_003



El Salvador: near San Vicente, Mun. Tepetitánvic. Finca El Carmen, 13°37′53.0″N,88°50′19.5″W, 4.I.2008, SCC08_005



El Salvador: Dep. San Vicente, vic. SanVicente, road to Zacatecoluca, behind stripclub ‘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

sel_912, sel_913, sel_914, sel_915


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

sel_930


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, MinaEl 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, CaminoSanta Rosa, Puente La Guayabita,12°44′31.2″N, 86°22′44.6″W, SCC08_017,SCC08_017

sel_961, sel_962, sel_963, sel_964,sel_965, sel_966, sel_967,sel_968, sel_969, sel_970


Nicaragua: Dept Matagalpa, Alc. San Ramon,Mata Palo, 12°56′16.5″N, 85°51′12.2″W,14.I.2008, SCC08_018

sel_971, sel_972, sel_973, sel_974


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_963, sel_975, sel_976, sel_977,sel_978, sel_979, sel_980,sel_981, sel_982, sel_983,sel_985, sel_986



APPENDIX Continued



Nicaragua: Lago Nicaragua, Isla Ometepe,Volcán Concepción, Charco Verde, HotelFinca Vincenzia and up hill across thestreet, 11°28′42.6″N, 35°38′20.6″W and11°29′31.2″N, 85°38′14.1″W, 12-13.I.2008,SCC08_014, SCC08_015

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


Costa Rica: Guanacaste, Palo Verde NationalPark, Cueva Las Tigres, 10°21′58.9″N,85°21′14.2″W, 17.I.2008, SCC08_022

sel_989, sel_990


Costa Rica: Guanacaste: Palo Verde FieldStation, hill behind OTES office,10°20′42.5″N, 85°20′19.1″W, 17.I.2008,SCC08_021

sel_987, sel_988


Costa Rica: Guanacaste: Nicoya Peninsula,near Loma Bonita, 10°15′04.0″N,85°17′30.5″W, 18.I.2008, SCC08_023

sel_991, sel_992, sel_993, sel_994

205 Selenops banksi Panamá: Barro Colorado Island, I.2008 sel_1000, sel_1001206 Selenops

mexicanusPanamá: STRI, Galeta, Plot F, 2004 sel_265, sel_266, sel_267, sel_268

207 Selenopsbursarius

Japan: Kyoto, Shugakuin, 23.VIII.2007 sel_861, sel_862, sel_863

208 Selenops radiatus Namibia: 12.VI.2006 sel_998, 999209 Selenops radiatus Tanzania: Iringa, Lutheran House Hostel sel_997210 Anyphops

barnardiSouth Africa: Guateng, Roodeport, Ruimsig

Butterfly Reservesel_547, sel_548

211 Anyphopsparvulus

South Africa: Tsitsikamma National Park,78 km E Knysna

sel_549

212 Anyphopstugelanus

South Africa: Grahmstown Municipal. CaravanPark

sel_550

213 Anyphopsstauntoni

South Africa: Eastern Cape, Kai Mouth, 58 kmNE East London

sel_551

214 Anyphops kraussi South Africa: Table Mountain National Park,Newland’s Forest

sel_552

215 Garcoropsmadagascarensis

Madagascar: Reserve Nat. Integrale de Lokobe,3.61 km ESE Hellville

sel_553

216 Hovops sp. Madagascar: Park National Montagne d’Ambremontane rainforest

sel_275

217 New genus sp. 1Australia

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

T80881, T80996


Australia: Western Australia: Two Peoples BayNature Reserve, granite outcrop, site 6,34°59′18″S, 118°44″E, 14.X.2006, undergranite rock, ML Moir, JM Waldock

T78485, T78489


Australia: Western Australia: Fiztgerald RiverNational Park, East Mt Barren, site 7,33°55′28″S, 120°01′13″E, 25.XI.2006, underrock ML Moir, KEC Brennan

T78500, T78498



Documents

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