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1996 McGUIRE-SYSTEMATICS OF CROTAPHYTID LIZARDS 25 Table 3.- Tooth count data for Crotaphytus. Crotaphytus: antiquus (n = 4) mean * SD 5.8 r 0.50 16.9 + 0.44 22.0 * 1.20 range (s-6) (1 5-19) (2 1-24) bicinctores (n = 24) mean 2 SO 6.2 + 0.53 16.9 2 0.44 23.0 2 2.25 range (5-7) (15-21) (18-28) collaris (n = 49) mean 2 SO 6.2 + 0.76 17.2 2 1.78 21.5 2 2.48 range (5-8) (14-22) (1 6-26) dickersonae (n = 16) mean rc- SD 7.1 2 0.95 20.3 * 2.60 24.8 * 3.51 range (6-9) (16-25) (19-31) grisrneri (n = 5) mean + SD 6.4 -C 0.89 18.6 2 1.96 23.1 + 2.23 range (6-8) (16-21) (I 9-26) insularis (n = 5) mean + SD 6.0 + 0.00 18.1 * 1.60 23.8 + 1.99 range (6) ( 1 5-20) (22-28) nebrius (n = 17) mean + SO 6.2 + 2.17 18.3 * 2.17 22.4 2.73 range (5-7) ( 1 5-23) (1 9-30) reticulatus (n = 25) mean 2 SD 6.0 + 0.64 17.7 + 1.69 21.9 * 1.75 range (5-7) (14-21) ( 1 7-25) taestigiunt (n = 28) mean 2 SO 6.2 5 0.39 18.3 + 1.61 23.3 + 2.12 range (6-7) (15-22) (1 9-28) most crotaphytids have an enlarged ridge that may support palatine teeth. This ridge is usually more developed in Garnbelia than Croraphytus. Most Gambelia (G. wislizenii, 39 of 46; G. copei, eight of nine; G. silus, 17 of 3 1 ) have palatine teeth. Within Crotaphyfus, the palatine ridge is almost always present but the teeth are only variably present (C. bicincrores, ten of 24; C. anriquus, three of four; C. Garnbelia: cope; (n = 9) mean * SD range silw (n = 31) mean * SD range tc~is1i:enii (n - 45) mcan + SD range Table 4.-Tooth count data for Gambelia. collaris, 1 9 of 45; C. dickersonae, 1 2 of 1 6; C. gris- meri, two of five; C. nebrjus. I 1 of 15; C. reticulatus, 17 of 26; C. vesligiurn, ten of 25), although only C. insularis (zero of five) always lacked palatine den- tition. Among the outgroup taxa examined, palatine teeth are present only in some Oplurus (0. quadri- maculaf us) and most pol ychrotids (all but Poly- chrus, although palatine teeth are also absent in all
26 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32 anoles except Chamaeleolis). Because Frost and Etheridge (1989) found Polychncs to be the sister taxon of the anoles, the presence of palatine teeth is considered as the ancestral state for Polychroti- dae. Therefore, if palatine teeth are to be considered apomorphic for Crotaphytidae, it must be assumed that Crotaphytidae and Polychrotidae are not sister taxa. Such a relationship was not supported in the analysis of Frost and Etheridge (1989) as depicted in their 12 equally parsimonious unrooted trees. Therefore, palatine teeth are tentatively considered to be apomorphic for Crotaphytidae. All crotaphytids possess pterygoid teeth on the posteromedial border of the palatine process (Fig. 11). These teeth may form a single row or, late in ontogeny, exist as a patch. During ontogeny, the number of pterygoid teeth clearly increases, although there is not a perfect correlation between number of teeth and SVL and some very large individuals have relatively few teeth. Additional teeth are usually added to the posterior portion of the patch, and in larger individuals, the majority of the teeth are found posteriorly. In some juvenile and most adult Crotaphytus, the posterior aspect of the pterygoid tooth row curves laterally away from the interpterygoid vacuity (Fig. 1 I), while in Gambelia the tooth row follows the margin of the vacuity. Polarization of this character is complicated by the absence of pterygoid teeth in the families Phryno- somatidae and Chamaeleonidae and in some Phy- marurus and Leiocephalus. Furthermore, pterygoid teeth are often intraspecifically variable and limited sample sizes for certain outgroup species probably did not allow them to be coded adequately for this character. However, in the remaining outgroup taxa examined, the pterygoid tooth patch was observed to curve posterolaterally only in Uranoscodon su- perciliosus, Corylophanes percarinatus, some C. cristatus, some Laemancttrs serratus. Brachylophus fasciaius, and Pristidactylus casuhatiensis (see de Queiroz, 1987, for additional iguanid taxa with posterolaterally curved pterygoid tooth patches). There- fore, the posterolateral curving of the pterygoid tooth patch was considered to be the derived state within Crotaphytidae. Scleral Ossicles. -The scleral ossicles are thin, overlapping platelets of bone that form a supportive ring within the anterior portion of the sclera of the eye. De Queiroz (1982) found that most iguanian taxa are characterized by a standard pattern con- sisting of 14 ossicles, with numbers one, six, and eight positive (overlapping both of the iyijacent ossicles), numbers four, seven, and ten negative (overlapped by both of the adjacent ossicles), and the remaining ossicles imbricating (overlapping one of the adjacent ossicles, but itself overlapped by the other). He noted that this pattern is present in Cro- taphyrus collaris, C. vestigiutn, and Gambelia wislizenii. I have verified his observations for these species, and report further that the remaining crotaphytid taxa are also characterized by this appar- ently ancestral iguanian condition. A list of speci- mens for which the scleral ossicles have been examined is provided in Appendix 7. Hyoid Appararits (Characters 34-36; Fig. 16). -A number of differences in the morphology of the hyoid apparatus exist between Crotaphytus and Gam- belia. In Crotaphyrus, the ceratohyals may be greatly expanded proximally, such that a large hook or process is present (processes absent in one of four C. antiquus). Their development is subject to ontoge- netic variation and subadults did not have the hook; therefore, the character was scored only from adults. In Gatnbelia, the proximal portion of the ceratohyal may be somewhat compressed; however, well-developed hooks are absent. This character varies ex- tensively in the outgroups and was therefore left unpolarized. In Gatnbelia, the second ceratobranchials are short, extending posteriorly for about half the length of the ceratohyals and first ceratobranchials, while in Crotaphytus they are longer, extending more than two-thirds the length of the ceratohyals and first ceratobranchials (Robison and Tanner, 1962; Fig. 16). The second ceratobranchials of C. dickersonae are often particularly long and in adult males usually extend as far posteriorly as do the ceratohyals and first ceratobranchials. However, this was not treated as a separate character state because of continuous variation between the extreme C. dickersonae condition and that present in other Crotaphytus, particularly in C. collaris. The longer second ceratobranchials of Croraphytus may function in the de- pression of their more strongly developed gular pouch. The outgroups vary continuously in the length of the second ceratobranchials ran&ng in relative length from very short in Phymarurus to extremely elongate in Polychrus, the anoles, and Brachylophus. Therefore, this character was left unpolarized. In Crotaphytus. the second ceratobranchials are in closecontact, although they are not actually fused, whereas in Gambelia, they may be widely separated
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