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McGUIRE-SYSTEMATICS OF CROTAPHYTID LIZARDS 37 Fig. 25.-Ventral view of the neck folds of Garnbelia wislizenii (SDSNH 68663). Fold terminology follows Frost (1992). AGF = antegular fold, GF = gular fold. folds may occur; however, they are shorter and appear near the lateral borders of the gular fold. As a result, the area of reduced squamation seen in Gambelia takes the form of a uniform band that extends across the width of the gular fold. Those outgroup taxa with gular folds examined here display both conditions of the fold with phrynosomatids, the oplurid Chalaradon madagascariensis, chamaeleonids (except Hydrosaurus pusfulatus), the corytophanids Basiliscus vittatus and Laemanctus, the hoplocercid Enyalioides lariceps (four of five), and polychrotids displaying the Gambelia form, and the hoplocercids Enyalioides praestabilis and E. oshaugnessyi, the corytophanid Basiliscur plumifons, the oplurid genus Oplum (0. firinensis and 0. saxicolus variable), and the iguanid Dipsosaurus displaying the Crotaphytus form. Most tropidurids have incomplete gular folds or lack them altogether; thus, the evaluation of this character for Tropiduridae is difficult. Uranoscodon superciliosus, which has a complete gular fold, displays the Gambelia form. The only other species within Tropidurini with Longltudlnal neck Postauricular I Supra-auricular Oblique neck --" Fig. 26.- Lateral view of the neck folds of Cro~aphy~usre~iculatur (EL 3250). Fold terminology follows Frost (1992). complete gular folds are Tropidurus azureum. T. flaviceps, and T. plica (Frost, 1992), species far re- moved from the basal lineages of the clade, and, thus, unable to shed light on this polarity decision. Because ofambiguity in the outgroup taxa, this character was left unpolarized. Supra-auricular Fold (Character 56; Fig. 26,27). - Frost (1992) defined the supra-auricular fold as a continuation of the dorsolateral fold that passes above the tympanum. In crotaphytids, a similar fold is present; however, it originates from the postauricular fold at a point roughly midway between the dorsal and ventral borders of the external auditory meatus. Without strong evidence to the contrary, I treat the crotaphytid fold as homologous with that described by Frost (1992) and therefore apply his standardized nomenclature. The condition of the supra-auricular fold, in which it originates midway between dorsal and ventral borders of the external auditory meatus, is present in many iguanian taxa Fig. 27.-Lateral view of the neck folds of Gambelia wis1i:enii (SDSNH 68663). Fold terminology follows Frost (1 992).
BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY Fig. 28.-An antehumeral mite pocket in a juvenile Crotaphyttcr grismeri. and, therefore, could not be included in the phy- logenetic analysis. The supra-auricular fold differs between Crota- phytus and Gambelia. In Crotaphytus, the fold extends posterodorsally at an angle of roughly 45 de- grees. In Gambelia, the fold extends posteriorly along a horizontal plane. In most of the outgroup taxa that have a supra-auricular fold, the fold either projects posteriorly along a horizontal axis, or occasionally, posteroventrally. However, some taxa may have a Crotaphytus-like supra-auricular fold (often vari- ably), including the phrynosomatids Petrosaurus re- pens. Uta stansburiana. U. squamata, U. palmeri. Urosaurus auriculattcs, and Phrynosoma coronatum, the tropidurids Leiocephalus schreibersi, L. melan- ochlorus, and L. psammodromus, the hoplocercid Enyalioides oshaughnessyi, and the chamaeleonids Urotnastyx acanthinurus and U. philbyi. Because of this variation and because many outgroup taxa can- not be scored for this feature, this character was left unpolarized. Antehutneral Fold (Fig. 26-28).-The antehumeral fold of crotaphytids is strongly developed, curving posteriorly over the forelimb insertion. The NO. 32 deepest portion of the fold is directly dorsal to the forelimb, a condition rarely observed in the outgroups. Furthermore, the antehumeral fold often extends posteriorly beyond the forelimb insertion, then continues posteroventrally or ventrally forming a complete arc. This condition is again uncommon in the outgroups. However, there is sufficient variation within Iguania that I have chosen not to code this as a character. The antehumeral fold of Crolaphytus dickersonae is unique among crotaphytids in ter- minating anterior to the forelimb insertion. Al- though this condition is probably derived within Crotaphytidae, another character, presence or absence of an antehumeral mite pocket, is certainly not independent. Therefore, this character is con- sidered under the section dealing with the antehumeral mite pocket. tlntehutneral mile Pocket (Character 57; Fig. 28).-In all Crotaphytus except C. dickersonae, the antehumeral fold is well developed (deep), with an area of reduced squamation dorsal to the forelimb insertion. The pocket almost always is inhabited by large numbers of trombiculid mite larvae. The presence of a mite pocket in this portion of the ante-
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