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McGUIRE- SYSTEMATICS OF CROTAPHYTID LIZARDS circles absent. Supraoculars small, flat or convex, smooth, becoming progressively larger medially such that medial scales are two to four times larger than lateral ones. Circumorbitals absent. Superciliaries eight to 12, extremely elongate medial scale present. Palpebrals ovoid, slightly convex, may be inter- spersed with numerous interstitial granules. Preo- culars, suboculars, and postoculars form an arc of four to seven rectangular scales, second, third, or fourth scale elongate. Supralabials 13 to 17, usually slightly longer than high except anteriormost scale, which is square. Lorilabials in two to four rows, ovoid to rectangular, juxtaposed, separating supralabials from suboculars and nasals. Aperture 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, somewhal conical auricular 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 postmentals that may be enlarged. Postmentals almost always separated from infralabials by sublabials on at least one side. Chinshields weakly differentiated or undifferentiated. Infralabials 12 to 17, square or wider than high, inferior border convex. Gulan usually flat, but occasionally convex and beadlike; each scale may be separated from adjacent scales by numerous asymmetrically arranged interstitial granules. Dorsal scales in approximately 160 to 200 rows midway between forelimb and hindlimb insertions. Tail long, cylindrical in both sexes and all age groups. Paired, median row of subcaudals not larger than adjacent subcaudals and lateral caudals. Enlarged postanal scales present in males. Deep postfemoral dermal mite pocket present at hindlimb insertion. Femoral pores 20 to 3 1, femoral pores extend beyond angle of knee, separated medially by ten to 18 granular scales. Subdigital la- mellae on fourth toe 20 to 24. Coloration in Life. -Individuals from southern San Diego County, the Sierra de Juarez and Sierra San Pedro Martir, and cismontane northwestern Baja California generally are dark brown in coloration with a pair of large paravertebral spots that are separated by broad, cream-colored transverse bars. There is much lateral flecking; however, lateral spots are lacking. Spots are nearly always absent from the head. In southern populations, such as those in the Vizcaino Desert, the base color of the dorsum is a paler golden tan, the dorsal spots are fragmented, and lateral spots may be present. In some southern individuals, the dorsal spotting may be nearly indistinguishable, with the dorsum peppered with fine pale speckling. This pattern may be more cryptic on the fine aeolian sand characteristic of the Vizcaino Desert (Grismer et al., 1994). The speckled pattern of the southern individuals appears to be an ontogenetic fragmentation of the color pattern characteristic of northern individuals as subadults have been examined with dorsal patterns very similar to those from the northern portion of the peninsula. Gravid coloration in G. copei is similar to that of G. bvislizenii, with orange or red spots often present on the head and/or neck, in two rows of spots on each flank, and on the ventral surface of the tail. Red or orange pigments may be present on the thighs as well. Males lack any form of breeding coloration. A description of the dorsal pattern of G. copei (pattern Cl) was provided in Montanucci (1978). Size. -This species exhibits strong sexual dimorphism with females reaching larger adult size (maximum observed SVL = 126 mm) than males (maximum observed SVL = 120 mm). Distribution (Fig. 50).- Gambelia copei occurs in extreme southcentral California in the vicinities of Cameron Corners (Mahrdt, 1973), Campo, and Potrero Grade southward through all but the San Felipe Desert region of northeastern Baja California to the northern portion of the cape region, Baja California Sur. The species is also found on the Pacific islands of Isla de Cedros, Isla Magdelena, and Isla Santa Margarita off of the west coast of the peninsula. Gat~tbelia copei occurs in the lower Colorado Desert region between El Huerfanito and Bahia de San Luis Gonzaga and is also known from the gulf coast desert region in the vicinities of Bahia de Los Angeles and Punta San Francisquito. However, G. copei apparently does not inhabit the Gulf Coast desert region between Santa Rosalia and the vicinity of Loreto and may be excluded from this region by the intervening Sierra San Pedro and Sierra de La Giganta. This species occurs in high densities on the sandy plains of the Vizcaino Peninsula and its distribution appears to be limited to the western side of the peninsular ranges from this region to a point at least as far south as the southern terminus of the Sierra de La Giganta. It is known from as far south as I km N Rancho Tres Hermanos (N of Todos Santos) in the cape region of Baja California Sur. The only published distribution map specific to Ga))tbelia copei (Banta and Tanner, 1968) is flawed
100 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32 in a number of respects. First, the lower Colorado Desert region between San Felipe and Puertocitos is inhabited by G. wislizenii rather than G. copei. Second, G. copei is not known from the eastern half of the peninsula between Bahia de San Francisquito and La Paz. Finally, G. copei is not known to range throughout the cape region. Gambelia copei is narrowly syntopic with G. wislizenii over a zone of approximately 1.6 km in Paseo de San Matias, Baja California (denoted by a rectangular mark on Figure 50). The two species are separated by a broad, transverse volcanic field that extends from the Sierra San Felipe to the gulf coast between Puertocitos and El Huerfanito, Baja Cali- fornia. This rugged volcanic flow, which is 3 1.5 km in width (by road), appears to act as an effective dispersal bamer for Gambelia along the gulf coast. On the provided dot distribution map (Fig. SO), the question marks represent localities in Baja Cal- ifornia Sur that are questionable because of impre- cise locality data (CAS 18823-San Andreas [San Jorge]; MVZ 37260-Medano Blanco, 37262-sand dunes 12 mi SE Venancio). Fossil Record. -None. Natural History. -Very little has been written re- garding the natural history of Gambelia copei, although it seems likely that it is similar to G. wislizenii in most aspects of its biology. This species is particularly common on the sparsely vegetated aeolian flats of the Vizcaino Peninsula, Baja California Sur, where it is often observed basking on roadside rocks, on the berms adjacent to graded din roads, or moving about in open spaces between clumps of vegetation. Gambelia copei is also found in more xeric creosote scrub habitats in the general vicinities of Paseo de San Matias, Bahia de San Luis Gonzaga, and Bahia de Los Angeles, Baja California, and in coastal sage scrub and oak woodland habitats on the western slopes of the Sierra San Pedro Martir and Sierra de Juarez (Welsh, 1988; personal observation). Where G. copei extends its range into extreme southcentral California, it apparently occurs in relatively densely vegetated chaparral. Gambelia copei shares a number of behavioral similarities with G. wislizenii and G. silus. For ex- ample, all three share a habit of basking on small stones and roadsides berms. Gambelia copei also displays the familiar "freeze" behavior such that when they are threatened, they run to the base of a bush or thicket, flatten themselves to the ground, and remain motionless (Tevis, 1944; personal observation). Like G. wislizenii. this species appears to be a lizard predation specialist as evidenced by the presence of Uta stansburiana and Cailisaurus draconoides in the stomach contents of museum specimens. Banta and Tanner (1 968) observed a Uta stansburiana and a grasshopper in the stomach of an adult female (CAS 8843) from Isla de Cedros. Like its sister taxon G. wislizenii, females attain larger sizes than males. Although rigorous ecological data are lacking, G. copei appears to be nonterritorial as in G. bvislizenii (personal observation). Although little is known of the predators of Gam- belia copei, they are likely to include the coachwhip snake (Masficophisflagellum) and patch-nosed snake (Salvadura hexalepis) as well as other saurophagous snake species, raptors, the Loggerhead Shrike (Lan- ius ludoricianus), the Greater Roadrunner (Geococ- cyx calfirnianus), as well as a number carnivorous mammals such as the coyote (Canis latrans). Only one predation event has been observed by the author, in which a Loggerhead Shrike was observed carrying a nearly full-grown G. copei. The shrike could only fly short distances with the relatively large lizard and, when pursued, was forced to pin the dead lizard on the spine ofa mesquite (Prosopis). Adult Garnbelia copei have been observed as early as 1 April 1993 in the vicinity of Cataviiia, on 9 April 1993 on the Vizcaino Peninsula, and on 10 April 1992 in Paseo de San Matias, indicating that this species emerges from hibernation at a date similar to that of G. wislizenii from southern California (Miller and Stebbins, 1964; Tollestrup, 1979; Mitchell, 1984). Although adult females were observed in April, none were gravid, suggesting that reproductive activity had not yet commenced. The earliest that gravid females have been observed by the author is 3 May 1993 at the Paseo de San Matias locality. Gravid females have also been seen on 27 June 199 1 in the Sierra San Borja and 4 July 199 1 in the Sierra Santa Clara. An emaciated female that appeared to have recently deposited eggs also was observed on 4 July in the Sierra Santa Clara, indicating that mating probably took place in mid to late June. Fitch (1970) examined 90 leopard lizards from Baja California (but did not list localities, so it is possible that some of the specimens were G. wislizenii) and found two of two females collected in March to be gravid, as well as six of nine collected in June, and three of six collected in July. Thus, the reproductive season is more extensive than my ob- servations would indicate.
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