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Biological Journal of the Linnean Society. 101, 288-322 Biological Journal of the Linnean Society, 2010, 101, 288–322. With 9 figures Molecular systematics of Selenops spiders (Araneae: Selenopidae) from North and Central America: implications for Caribbean biogeography SARAH C. CREWS1,2* and ROSEMARY G. GILLESPIE1 1University of California Berkeley, Department of Environmental Sciences Policy and Management, 137 Mulford Hall, Berkeley, CA 94720-3114, USA 2Berkeley City College, Department of Science and Biotechnology, 2050 Center Street, Berkeley, CA 94704, USA Received 16 February 2010; revised 3 May 2010; accepted for publication 3 May 2010bij_1494 288..322 The Caribbean region includes a geologically complex mix of islands, which have served as a backdrop for some significant studies of biogeography, mostly with vertebrates. Here, we use the tropical/subtropical spider genus Selenops (Selenopidae) to obtain a finer resolution of the role of geology in shaping patterns of species diversity. We obtained a broad geographic sample from over 200 localities from both the islands and American mainland. DNA sequence data were generated for three mitochondrial genes and one nuclear gene for eleven outgroup taxa and nearly 60 selenopid species. Phylogenetic analysis of the data revealed several biogeographic patterns common to other lineages that have diversified in the region, the most significant being: (1) a distinct biogeographic break between Northern and Southern Lesser Antilles, although with a slight shift in the location of the disjunction; (2) diversification within the islands of Jamaica and Hispaniola; (3) higher diversity of species in the Greater Antilles relative to the Lesser Antilles. However, a strikingly unique pattern in Caribbean Selenops is that Cuban species are not basal in the Caribbean clade. Analyses to test competing hypotheses of vicariance and dispersal support colonization through GAARlandia, an Eocene–Oligocene land span extending from South America to the Greater Antilles, rather than over-water dispersal. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322. ADDITIONAL KEYWORDS: Bayesian phylogenetics – island biogeography – likelihood analysis of geographic range evolution. INTRODUCTION insights into the complex interaction between coloni- zation and diversification. In particular, the Carib- Remote islands form the basis for many biological bean has served as the setting for the establishment studies because of their ability to act as a laboratory, of most of the central tenets in the equilibrium theory with repeated sets of ecological and/or evolutionary of island biogeography (Munroe, 1948), the argu- experiments occurring within a circumscribed time ments being formulated independently by MacArthur frame (Cronk, 1997; Losos et al., 1998; Gillespie & and Wilson (1963, 1967) much later (Lomolino & Roderick, 2002; Gillespie, 2004; Ricklefs & Berming- Brown, 2009). More recent research on the islands ham, 2008). While the Hawaiian Islands have served has allowed an understanding of the interplay as a model system for processes of in situ diversifica- between ecological and evolutionary processes in tion, the long history of studies on the biota of the shaping species diversity (Losos & Schluter, 2000; Caribbean has provided some of the most important Schoener, Spiller & Losos, 2001). The primary feature of the Caribbean region that makes it particularly useful for examining the inter- *Corresponding author. E-mail: [email protected] action between colonization and diversification is its 288 © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322 SYSTEMATICS AND BIOGEOGRAPHY OF SELENOPS 289 long and complex geological history. The Caribbean South America to the Greater Antilles during the Basin began forming nearly 140 Mya. Islands in the Eocene–Oligocene transition 35–33 Mya. The land basin consist of four different types: (1) land-bridge span, although probably short-lived, may have pro- islands which were connected to each other or to the vided an avenue for terrestrial organisms to colonize mainland at times of lower sea level; (2) continental the Greater Antilles from South America. Among islands which broke off from the mainland through mammals, molecular phylogenies of primates and tectonic displacement; (3) uplifted limestone islands; hystricognath rodents are consistent with the model, and (4) volcanic islands (MacPhee & Iturralde-Vinent, while sloths and insectivorans are not (Dávalos, 2005; Robertson, 2009). Despite their limited isola- 2004). The pattern in plants is similarly mixed. tion, the age and geologic complexity of the area have Molecular phylogenetic data from the genera Croton provided ‘well-defined paths of entry by which immi- (Euphorbiaceae) (van Ee et al., 2008) and Styrax (Sty- grants may reach’ the islands (Munroe, 1948). More- racaceae) (Fritsch, 2003) show that the timing of over, the islands have served as the setting for divergence of lineages is consistent with the GAAR- adaptive radiation among lineages with limited dis- landia hypothesis. However, similar data from persal ability, in particular lizards of the genus Anolis endemic legume radiations in the Greater Antilles, (Losos, 1992, 1994, 2009), frogs of the genus Eleuth- although initially thought to indicate ancient splitting erodactylus (Hedges, 1989; Heinicke, Duellman & between lineages consistent with the GAARlandia Hedges, 2007), some lineages of insects [e.g. beetles hypothesis (Lavin et al., 2001), show more recent (Liebherr, 1988b), flies (Wilder & Hollocher, 2003)] diversification (Lavin & Beyra-Matos, 2008), which is and plants [e.g. lineages within the Melastomaceae likely to hold also for lineages of Asteraceae (Michelangeli et al., 2008) and Asteraceae (Francisco- (Francisco-Ortega et al., 2008). Ortega et al., 2008)]. Although studies to date have Clearly, the timing and frequency of dispersal and provided insights into how the individual lineages vicariance, and the interplay between the two, varies have colonized and subsequently diversified within across biotic assemblages. The challenge, then, is to the island system, notable controversies remain, understand the circumstances dictating the relative including the source of colonists and the means by roles of each and how they interact. Arthropods, which they colonized the islands, biogeographic pat- because they can provide a fine-scale resolution of terns within lineages and whether these patterns biogeographic patterns (Ferrier et al, 2004), are ideal might be expected to be shared across multiple lin- candidates for elucidating the nature of these rela- eages (Guyer & Savage, 1986; Williams, 1989; tionships. Although the biogeography of terrestrial Hedges, Hass & Maxon, 1992; Crother & Guyer, 1996; invertebrates in the Caribbean has been examined in Hedges, 1996a,b). some detail (see Liebherr, 1988a and chapters A particular focus of debate has been the role of therein), few recent studies have been attempted, vicariance vs. dispersal in shaping the Caribbean with little molecular information on the timing and biota. Hedges and colleagues (Hedges et al., 1992; nature of the interplay between colonization and Hedges, 1996a,b; Hedges & Heinicke, 2007; Heinicke diversification. However, there are some notable et al., 2007), working with herpetofauna, have sug- exceptions (Davies & Bermingham, 2002; Wilder & gested that the absence of lineages older than the Hollocher, 2003; Brisson, Wilder & Hollocher, 2006). break-up of the proto-Antilles (a contiguous land In particular, recent studies on spiders (Sicariidae: mass between North and South America) precludes a Loxosceles) support the GAARlandia hypothesis in the vicariant origin and they argue for the initial coloni- colonization of the lineage of North from South zation of most taxa via over-water dispersal on America (Binford et al., 2008), while crickets show a flotsam. A similarly dominant role for dispersal has more mixed pattern of both vicariance and dispersal, been suggested for multiple lineages of plants, such coupled with intra-island diversification (Oneal, as Miconieae (Michelangeli et al., 2008). In contrast, 2009). other studies have suggested that vicariance has In this study, we combine molecular and morpho- played a larger role than dispersal in the initial logical methods to examine the phylogenetic relation- colonization of the Caribbean; for example, in lizards ships and biogeographic history of the cursorial and (Crother & Guyer, 1996; Iturralde-Vinent & MacPhee, dispersal-limited spider genus Selenops (Araneae: 1999; MacPhee & Iturralde-Vinent, 2005) and some Selenopidae) in the Caribbean. These primarily tropi- plants [e.g. Euphorbiaceae (van Ee et al., 2008)]. cal and subtropical spiders (Muma, 1953; Corronca, A related controversy focuses on the hypothesis of 1998; Alayón, 2005) are distinctive in that they are GAARlandia (Greater Antilles + Aves Ridge), first extremely dorsoventrally flattened and exceedingly proposed by Iturralde-Vinent & MacPhee (1999), who fast. They are found in a variety of habitats and used geological data and fossil evidence to demon- microhabitats (Crews, Wienskoski & Gillespie, 2008). strate the likely existence of a land span connecting Although the genera and species groups have © 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
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