Relationships of West Indian Anolis (Sauria: Iguanidae): An Approach Using Slow-Evolving Protein Loci

– Protein variation in 49 West Indian species of the iguanid lizard genus Anolis was examined by sequential electrophoresis at 12 slow-evolving loci. The use of this technique nearly doubled the total number of alleles detected (121 before, 233 after). Genetic distance and parsimony analyses identified intra-island radiations on Cuba, Jamaica, Hispaniola, and Puerto Rico, and found little evidence of close inter-island relationships. In most cases, these island radiations (series) defined by protein data were supported by other data (morphology, immunology, chromosomes). No support was obtained for previously defined higher-level groupings (sections and subsections) within the genus. A revised classification is proposed that recognizes 21 series of West Indian Anolis (131 spp.) with distributions centering on the following islands or island groups: Cuba (alutaceus, argillaceous, carolinensis, equestris, lucius, and sagrei), Jamaica (grahami), Hispaniola (chlorocyanus, christophei, cuvieri, cybotes, darlingtoni, distichus, hendersoni, monticola, semilineatus, and sheplani), the Puerto Rican Bank (cristatellus, occultus), the northern Lesser Antilles (bimaculatus), and the southern Lesser Antilles (roquet). No categories above the level of series are recognized due to conflicting evidence for higher-level relationships. Although the existence of Anolis on the North Island (Hispaniola) in the Eocene or Oligocene indicates an early arrival of the genus in the West Indies, molecular dating suggests that mid-Tertiary dispersal and not early-Tertiary vicariance best explains the present distribution of the group. Extensive intra-island radiation occurred during the late-Tertiary (Miocene-Present) with relatively little inter-island dispersal among the Greater Antilles. The iguanid lizard genus Anolis (sensu Williams, 1976a, b) comprises over 300 described species. In addition to being the largest amniote genus, it has figured prominently in the ecological, ethnological, and systematic literature. However, the phylogeny of this group has been a continuous challenge to systematists since the initial work done by Etheridge (1960). Williams (1976a, b) published the first comprehensive taxonomy of these animals. His classification relied primarily on osteology and divided the genus into two sections, alpha and beta. These sections originally were established by Etheridge (1960) and are defined osteologically by the absence (alpha) or presence (beta) of transverse processes on posterior caudal vertebrae. Williams further divided the alpha section into two subsections, also osteologically defined. 1 Current address: Department of Biological Sciences, University of California, Santa Barbara, California 93106, USA (KLB); and Department of Biology, 208 Mueller Lab, Penn State University, University Park, Pennsylvania 16802, USA (SBH). The subsections are based on the shape of the interclavicle, which is arrow-shaped (punctatus subsection) or T-shaped (carolinensis subsection). Subsections contain series that are defined, again, by osteological characters as well as by karyological and scale characters. Series are further broken in to subse r i e s , s p e c i e s g r o u p s a n d subgroups. Most of the systematic work on Anolis has involved West Indian taxa (131 spp.). Albumin immunological data presented by Wyles and Gorman (1980a) and Shochat and Dessauer (1981) did not support the alpha-beta dichotomy. They found that members of the grahami series (beta section) and cristatellus series (alpha section) clustered more closely with each other than either did with any other series within their own sections. Shochat and Dessauer erected the “Central Caribbean series complex” to include series of alpha and beta anoles that appeared to form a monophyletic group. Guyer and Savage (1986) proposed a revised classification of Anolis after reana-