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McGUIRE- SYSTEMATICS OF CROTAPHYTID LIZARDS E~ytnoIogy. -From the Latin vestigium. a footprint, a track, a tncc. In referencc to the reduced collars of this species (Tanner, personal communication, 1993). Diagnosis. - Crotaphyf us sestigium can be distinguished from all other Crotaphytus except C. insularis and C. rericulatus by the presence of widely separated posterior collars. It can be distinguished from all other species of Crotaphytus by the presence of slender, white transverse dorsal body bars. It can be further distinguished from C. reficulatus. C. collaris, C. nebrius, and C. dickersonae by the absence of black oral melanin. It can be further distinguished from C. reticulatus, C. collaris, and C. nebrius by the presence in adult males of a strongly laterally compressed tail, a white or off-white dorsal caudal stripe, a pale tan or white patternless region on the dorsal surface of the head, and enlarged dark brown or black inguinal patches (rather than the small inguinal patches of C. nebrilrs and some C. collaris). It can be distinguished from C. antiqutts and further distinguished from C. retic~rlatus in the absence of a dorsal pattern composed ofa white reticulum with some or all of the reticulations enclosing black pigmentation. It can be further distinguished from C. grismeri by the absence ofa greenish tint to the white bar that separates the anterior and posterior collars, by the hindlimb pattern consisting of white reticulations or spots on a brown field (field occasionally yellowish distal to the knee), by the presence ofolive green or burnt orange ventrolateral coloration, and by its much larger maximum adult SVL. It can be distinguished from C. insularis by its broader nasal process of the premaxilla and its more strongly developed posterior collar. Ifariation (n = 28).-Rostra1 approximately four times wider than high, usually rectangular in shape. Rostral bordered by two to five postrostrals. Remaining snout scales irregularly arranged, an enlarged middorsal series may be present. Nasals separated by three to five internasals. Frontonasals occasionally enlarged. Canthals three; five to seven scales separate canthals of left and right sides. Supraorbital semicircles present, median scales rarely fuse to form an azygous frontal. Supraoculars flat or convex, smooth, becoming progressively larger medially such that medial scales are two to four times larger than lateral ones. Circumorbitals present, not well differentiated from supraoculars. Superciliaries nine to 12, extremely elongate medial scale occasionally present. Palpebrals ovoid, slightly convex, interspersed with numerous interstitial granules. Preoculars, suboculars, and postoculars form an arc of six to 1 1 rectangular scales, second, third, or fourth scale not elongate. Supralabials ten to 18, usually slightly longer than high except anteriormost scale, which is square or pentagonal. Lorilabials in two to three rows, ovoid to rectangular, juxtaposed, sepa- rating supralabiais from suboculars and nasals. Ap- erture of external auditory meatus rectangular or ovoid, often constricted at or above the midpoint, approximately two to four times higher than wide, with small, strongly convex, somewhat conical au- ricular scales lining anterior margin. Mental pentagonal, one to 1.5 times wider than high, bordered laterally by anterior infralabials and posteriorly by a pair of large postmentals. Postmcntals may or may not be separated from infralabials by one or two sublabials. Chinshields weakly differentiated or un- differentiated. Infralabials 1 1 to 17, square or wider than high, inferior border convex. Gulars granular, strongly convex and beadlike, each scale separated from adjacent scales by numerous asymmetrically arranged interstitial granules. Dorsal scales in approximately 156 to 21 2 rows midway between forelimb and hindlimb insertions. Tail long, cylindrical to oval in females and juve- niles over entire length, anterior one-half strongly compressed laterally in adult males. Paired, median row of subcaudals larger than adjacent subcaudals and lateral caudals. Enlarged postanal scales in males present. Deep postfemoral dermal mite pocket present at hindlimb insertion. Femoral pores 15 to 25, femoral pores do not extend beyond angle of knee, separated medially by 17 to 24 granular scales. Subdigital la- mellae on fourth toe 15 to 25. Coloration in Lijie. -Dorsal body coloration in adult males is brown. The white component of the dorsal pattern is composed of white spots and dashes on the body, a reticulated tail and hindlimbs, and forelimbs that are either reticulated, spotted, or nearly patternless. Slender, transverse body bars are present in both sexes. Reticulations are always present on the superficial mandibular and temporal regions. A broad white or off-white caudal vertebral stripe is present. The dorsal surface of the head is pale-colored, and is conspicuously patternless. Ei- ther olive green or golden orange ventrolateral coloration is present in adult males, with the former color present in individuals north of Bahia de San Luis Gonzaga, Baja California, Mexico, and the lat- ter color present in individuals from Bahia de Los Angeles southward. The ventrolateral coloration of
96 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32 individuals occurring between Bahia de San Luis Gonzaga and Bahia de Los Angeles is not known. The gular coloration in adult males is generally slate gray or gun-barrel blue, with a black central gular component. The peripheral gular pattern is the stan- dard reticulate form. Anterior collar markings are always present and posterior collar marks are only rarely lacking. The posterior markings are widely separated middorsally. The anterior collar markings are complete ventrally in adult males, with black pigments extending through the gular fold. A pair of black nuchal spots are not present middorsally between the anterior collar markings. Enlarged melanic axillary patches immediately posterior to the forelimb insertion are variably present. Large melanic inguinal patches are always present. The femoral pores are generally off-white to gray in color. Paired, melanic keels are always present on the ven- tral surface of the caudal extremity. Females are less vividly marked than males. The dorsal coloration is usually gray or greenish gray. The head and gular markings are less developed and male color pattern characteristics such as the white dorsal caudal stripe, ventrally complete anterior collar markings, and melanic inguinal patches, axillary patches, and central gular spot are lacking. Gravid females develop vivid orange or reddish lateral bars. The tail is not brightly colored in adult or subadult females of this species. Size. -This species exhibits strong sexual dimor- phism with males reaching larger adult size (maximum observed SVL = 125 mm) than females (maximum observed SVL = 98 mm). Distribution (Fig. 48). - Crotaphyrus vestigium in - habits the peninsular ranges and adjacent rocky habitats from the northern slope of the San Jacinto <strong>Mountain</strong>s in southern California to the southern margin of the volcanic Magdalena Plain in Baja Cal- ifornia Sur. In southern California and northern Baja California, C. vestigium is limited to the eastern face of the peninsular ranges. There is a gap in the peninsular ranges between the southern edge of the Si- erra San Pedro Martir and the northern edge of the Sierra La Asamblea and C. vstigiurn occurs on either side of the peninsular ranges from this point southward. Furthermore, its range extends nonh- ward along the western side of the peninsular ranges from this gap to a point at least as far north as the vicinity of Rancho San Jose (Meling's Ranch) and even approaches the Pacific Coast at Mesa San Car- los (Bostic, 1971). The known southern distributional limit of C. vestigiunr is 27.7 km (by road) S of San Jose de Comondu (McGuire, 1991). It is likely that the actual distributional limit is bounded by the volcanic mesas that terminate near this locality. Crotaphyttcs vestigittm apparently does not inhabit the isolated Sierra Santa Clara and Sierra Vizcaino on the Vizcaino Peninsula (Grismer et al., 1994). Fossil Record. -None. Narrtral History. -Very little has been written regarding the natural history of Crotaphytus vestigium. Sanborn and Loomis (1979) discussed the display patterns for this species and noted that it inhabits rocky outcroppings on the more rugged portions of the alluvial fans and mountain slopes at their San Jacinto <strong>Mountain</strong>s study site. Common plant species at this locality included Larrea tridenrara, Encelia Jarinosa, and Ambrosia dumosa. Welsh ( 1 988) collected two individuals, one of which was found on a rocky volcanic slope in central desert scrub and the other on a granitic outcrop in coastal sage scrub. Bostic (1 97 1) observed two individuals on Mesa San Carlos, a table-topped mountain overlooking the Pacific coast of Baja California approximately 350 km south of the United States-Mexico border. One of these individuals was foraging among large basaltic rocks along the edge of the mesa while the other was seen basking on a large basaltic outcropping on top of the mesa proper. Crotaphytus vestigiurn is a denizen of desert hillsides, alluvial fans, canyons, and lava flows, always in association with rocks. They occur in some of the most xeric habitats of Nonh America such as the eastern bases of the Sierra de Juarez and Sierra San Pedro Martir where they may be observed basking during the heat of the day. The rocky habitats in which they occur generally are characterized by scant vegetation. Common plant taxa with which C. vestigiu~n is often associated include Fouquieria splendens, F. diguni, Opunria, Larrea tridenfata, Pachycormus discolor, Bursera, Ferocacrus, Pachycereus pringlei, Prosopis, and numerous additional xerophilic species. When alarmed, this species can move with great speed over complex rocky terrain by bounding bipedally from one stone to the next, often taking refuge beneath a larger rock. The activity season for adult Crotaphyttcs vestigium probably commences in March. Adults have been observed as early as 1 April 1992 at the foot of the Sierra La Asamblea, Baja California, and adult males, gravid females, and subadults have been
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