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1996 McGUIRE-SYSTEMATICS OF CROTAPHYTID LIZARDS I I MORPHOLOGY AND CHARACTER DESCRIPTIONS SKULL ROOF Prernaxilla (Characters 1, 2; Fig. 2-5, 7).-The posterodorsally projecting nasal process is long and very slender in Gatnbelia rvislizenii and G. copei (Fig. 4, 5) and broad in most Crotaphytus (Fig. 2, 7) and the single specimen of G. coronat. Gambelia silus (Fig. 3) occasionally has a slender but short nasal process (seven of 30) owing to its truncated snout. In C. insularis, the nasal process is also long and extremely narrow, which may be a consequence of elongation of the snout region. Some variation occurs in C. vestigiutn and C. bicinctores, both with two of 28 specimens having similarly slender nasal processes, and C. grismeri, with one of five having a slender nasal process, although not as extreme as that seen in C. insularis. Among the outgroup taxa, a narrow nasal process was observed only in Peiro- saurus tnearnsi and occasional Uta stansburiana, Dipsosaitrus dorsalis, Phytnaturus pallurna, and Ph. punae (although the condition observed was not as extreme as that observed in Gambelia and C. insularis). Therefore, an elongate, narrow nasal process is considered to be the derived state. In Gambelia, the anteromedial portion of the alveolar shelf at the articulation of the premaxilla and vomers is in the form of a strong vertical ridge. This ridge is rarely present in Crotaphytus (three of 51 C. collaris, one of four C. antiquus). Among the outgroup taxa, a strong vertical ridge was observed only in Corytophanes hernandezi, r\.ficroIophw grayi, two of three Leiocephalus schreibersi, and one of three Phymattrrus patagorticus zapalensis. There- fore, a strong vertical ridge at the alveolar shelf is considered to be the derived state. The premaxillary base is also subject to much variation in crotaphytids. In all Gantbelia, plus many C. antiqiius, C. collaris (primarily those formerly - Fig. 2.-Doml view ofthc skull ofCroraphyturdickersonae(REE Fig. 3.-Dorsal view of the skull of Garnbelia silw (CAS 22713, 2777, adult male, SVL = 116 mm). Scale = 5 mm. adult male, SVL = 101 mm). Scale - 10 mm.
BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32 Fig. 4.-Dona1 view of the skull of Gambelia wislizenii (REE 29 18, adult female, SVL = 1 19 mm). Scale = I0 mm. referred to the subspecies C. c. auriceps and C. c. baileyi), C. grismeri, C. nebrius. and C. reticulatus, the broad, laterally oriented maxillary processes give the base a rectangular shape as opposed to a trapezoidal shape (Fig. 2-4). This condition is either absent or appears rarely in C. bicinctores, C. dickersonae, C. insularis, and C. vestigium. Despite this trend, most Crotaphytus species display continuous variation in this feature with all intermediates between the rectangular and nonrectangular conditions present. Therefore, this character was not included in the phylogene~ic study. Nasals (Character 3; Fig. 24,7).- In Crotaphytus dickersonae (Fig. 2), two of four C. antiquus, and one of 28 C. bicinctores, forward expansion of the nasals results in their overlap of the nasal process Fig. 5.-Anterior portion of the skull of Gambelia wislizenii(REE 29 18, adult female, SVL = 1 19 mm) depicting the saddle-shaped premaxillary-maxillary aniculations. The premaxilla is veriically hatched. Max = maxilla, Nas = nasal. Scale = 5 mm. of the premaxilla well anterior to the posterior bor- der of the external nares (fenestrae exonarina of Oel- rich, 1956). This feature varies ontogenetically in C. dickersonae, with individuals of less than 8 1 mm snout-vent length (SVL) having incomplete contact of the nasals anteriorly (character scored only from adults). The nasals occasionally overlap the nasal process of the premaxilla anterior to the posterior extent of the external nares in Gambelia wislizenii and G. copei. However, this appears to be the result of posterior expansion of the nares rather than an anterior expansion of the nasals and is here considered to be nonhomologous. The nasals only rarely overlap the nasal process of the premaxilla anterior to the posterior extent of the external nares in the outgroup taxa. This condition was observed in Broo- kesia stumpfi and in a number of tropidurid taxa (Ctenoblepharys adspersus, some Phymaturus patagonicus patagonicus, P. p. payuniae, and P. p. so- muncurensis [but not other Phymaturus], many LRi- ocephalus species, and Microlophus grayi). Pregill (1992) considered this feature to be absent from most basal extant Leiocephalus, including L. her- minieri. L. greentvayi. L. punctatus, L. inaguae. L. psammodrornus, and some L. carinatus. Therefore, the conditions observed in Liolaeminae, Leioce- phalinae, and at least one member of Tropidurinae may be convergent. Nevertheless, nasals that overlap the nasal process of the premaxilla may be an- cestral for Liolaeminae and Leiocephalinae, and,
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