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1996 McGUIRE-SYSTEMATICS OF CROTAPHYTID LIZARDS 9 debate concerning crotaphytids has been whether or not Garnbelia should be synonymized with Crota- phytus (Smith, 1946; Robison and Tanner, 1962; Weiner and Smith, 1965; Montanucci, 1969, 1978; Montanucci et al., 1975; Tanner and Banta, 1977). Nevertheless, very little character evidence has been presented supporting the monophyly of Crotaphy- tidae. Etheridge and de Queiroz (1988) recognized crotaphytids as a monophyletic group on the basis of a unique combination of derived, yet highly ho- moplastic features: the presence of posterior coracoid fenestrae and female gravid coloration, and the absence of postfrontal bones and a middorsal scale row. Frost and Etheridge (1 989) considered crotaphytids to be monophyletic on the basis of three reversals: presence of palatine teeth, posterior coracoid fenestrae, and ribs on the third cervical ver- tebra (the last ofwhich is only infrequently observed in crotaphytids). In each of these analyses, character support for Crotaphytidae was dependent upon its placement within the ingroup topology. The follow- ing is a list of synapomorphies of Crotaphytidae recognized in this study: presence of black oral pig- mentation (reversed within Crotaphytus), presence of a posterolaterally projecting jugal-ectopterygoid tubercle immediately posterior to the maxillary tooth row, presence of posterior coracoid fenestrae, the tympanic crest of the retroarticular process of the mandible curves posterodorsally, the parietal and frontal strongly overlap the medial process of the postorbital, the supratemporal lies in a groove along the ventral or ventrolateral border of the supratemporal process of the parietal (reversed in most G. silus or convergent in Crotaphytus and other Garn- belia), presence of palatine teeth, and contact of the prefrontal and jugal in the anterolateral border of the orbit. The terminal taxa utilized in this study include the currently recognized species of Crotaphyrus (C. antiquus, C. bicinctores, C. collaris, C. dickersonae, C. grismeri, C. insularis, C. reticulatus, and C. ves- tigium) and Garnbelia (G. coronat, G. silus, and G. wislizenii). Over the course of this study, it was determined that at least one and probably two addi- tional species should be recognized. These include two taxa currently recognized as subspecies, C. c. nebrius and G. bv. copei (see taxonomic accounts for data supporting the elevation of these taxa to full species). These species were also included in the analysis. Another population of Gambelia that may even- tually prove to be a full species is the population of G. ivislizenii on Isla Tiburon in the Gulf of Califor- nia. The four osteological specimens examined in this study lacked autotomic fracture planes in the caudal vertebrae. Fracture planes are present in all other G. \clislizenii (n = 19) and G. silus (n = 5) specimens examined, although they appeared to be fused in three of ten G. copei. Unfortunately, no osteological specimens were available from adjacent Sonora and it could not be determined ifthe absence of fracture planes is confined to this insular population. If this population proves to be a separate species, it may be the only endemic reptile or am- phibian on Isla Tiburon, a land-bridge island that supports an extensive herpetofauna. The remaining subspecies of Crotaphytus collaris and Gambelia nislizenii were not treated as separate terminal taxa because no evidence has been presented, nor has any been discovered over the course of this investigation, suggesting that these forms are discrete evolutionary entities. Rather, they are pat- tern or convenience classes (Frost et al., 1992), color morphs largely consistent over an extensive area, but grading smoothly into other color morphs at their boundaries. Etheridge and de Queiroz (1988) and Frost and Etheridge (1 989) provided evidence for the monophyly of nine suprageneric groups (elevated to families in the latter study) within Iguania. Interfamilial resolution was elusive and their strict consensus tree (at the familial level) was an unresolved polytomy. However, they were able to substantially reduce the number of equally parsimonious interfamilial to- pologies as depicted in their 12 unrooted trees with rooting points (Fig. I). Thus, despite the continuing lack of unambiguous interfamilial resolution, the outgroup situation has improved considerably. In this analysis, characters were considered to be po- larized only when the polarity assessment was consistent with all 12 unrooted trees. For each of the eight remaining iguanian families, exemplars were examined for the purpose of character polarization. The choice of exemplars was based whenever possible on the results of recent intrafamilial phylogenetic analyses. Thus, basal lin- eages have been proposed for clades within the families Phrynosomatidae (Presch, 1969; Montanucci, 1987; Etheridge and de Queiroz, 1 988; de Queiroz, 1989, 1992; Wiens, 1993a, 1993b), Tropiduridae
BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32 Fig. 1.-The 12 unrooted trees discovered by Frost and Etheridge (1989) in their phylogenetic analysis of iguanian lizards. The open circles represent the discovered rooting points for these unrooted trees. Ch = Chamaeleonidae, Co - Corytophanidae, Cr = Crotaphytidae, Ho = Hoplocercidae, Ig = Iguanidae, Op = Opluridae, Ph = Phrynosomatidae, Po - Polychrotidae, TI = Tropiduridae. (Etheridge and de Queiroz, 1988; Frost and Ether- idge, 1989; Frost, 1992; Pregill, 1992; Etheridge, 1999, Corytophanidae (Etheridge and de Queiroz, 1988; Lang, 1989), Hoplocercidae (Etheridge and de Queiroz, 1988), Iguanidae (de Queiroz, 1987; Norell and de Queiroz, 199 l), Polychrotidae (Guyer and Savage, 1986, 1992; Etheridge and de Queiroz, 1988; Cannatella and de Queiroz, 1989), and Cha- maeleonidae (Moody, 1980, 1987; Klaver, 198 1 ; Klaver and Bohme, 1986; Hillenius, 1986, 1988; Rieppel, 1987; Frost and Etheridge, 1989). For the remaining family (Opluridae), only the phenetic analysis of Blanc et al. (1983) was available. For this lineage, I examined Chalaradon and as many species of Oplurus as possible. A list of outgroup taxa examined for this study is provided in Appendix 1 and a data matrix documenting the character states observed in these taxa is provided in Appendix 3.
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