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1996 McGUIRE-SYSTEMATICS OF CROTAPHYTID LIZARDS 7 1 midway between forelimb and hindlimb insertions. Tail long, cylindrical in both sexes and all age groups. Paired, median row of subcaudals may or may not be larger than adjacent subcaudals and lateral cau- dals. Enlarged or slightly enlarged postanal scales present in males. Deep postfemoral dermal mite pocket present at hindlimb insertion. Femoral pores 16 to 20, femoral pores do not extend beyond angle of knee, separated medially by 19 to 25 granular scales. Subdigital la- mellae on fourth toe 18 to 22. Coloration in Life.-Males of this species are characterized by a dorsal color pattern consisting of a thick white reticulum on a dark brown field. The reticulations differ from those of C. reticulatus in that they are thicker, and all, or nearly all, of the dorsal body reticulations enclose black pigments. A few of the forelimb and hindlimb reticulations may also enclose black pigments. As in C. reticulatus, the reticulum is present on nearly the entire dorsal surface including the body, the anterior half of the tail, all four limbs, the lateral surface of the head, and the superficial mandibular area. The anterior and posterior collar markings are better developed than those of C. reticulatus and the anterior collar is complete ventrally. Black pigmentation is present in the central gular region, as in all other adult male Cro- taphytus except C. collaris and C. nebrius. The dorsal surface of the head is patternless, but it is not of paler coloration than the remaining dorsal surfaces, as is usually the case with C. dickersonae, C. bicinctores, C. grismeri. C. vesrigiurn, and C. insularis. Small inguinal patches largely confined to the proximal ventral surface of the thigh are present in all adult males. The femoral pores are jet black. The coloration of females is less vibrant than that of males. The dorsal base color is grayish brown, the white reticulum is not as bright, the dorsal reticulum encloses dark gray pigments rather than black, the femoral pore exudate is gray, and the melanic inguinal patches and black pigments of the gular fold and central gular region are absent. Fe- males develop orange gravid coloration during the reproductive period. The one subadult female that I have examined in life had a bright yellow tail and hindlimbs. Distribution (Fig. 43). -Known to occur in the Sierras de San Lorenzo, Texas, and Solis of extreme southwestern Coahuila, Mexico. Fossil Record. -None. Natural History. -The following natural history observations were made on 23 and 25 June 1994. As are all Crotaphytus except C. reticulatus, C, an- Texae Fig. 43.-Geographic distribution of Crotaphytus anriquus. The asterisk indicates the location of the Sierras dc San Lorenzo, Texas, and Solis in southwestern Coahuila, Mexico. tiquus is strongly saxicolous and usually is observed basking on large limestone rocks and outcrops. When alarmed, they generally take refuge beneath a nearby rock or under the rock upon which they were perched. The habitat at the type locality is fairly typical Chi- huahuan Desert scrub with the dominant plant species being Larrea divaricata, Jarropha dioica, Fou- qiiieria splendens, Agave lechuguilla, Lippia grav- eolens, Opuntia cholla, two unidentified species of Opuntia (one resembling prickly pear, the other similar in habitus to pencil cholla), and (possibly) Echi- nocactus sp. Additional reptile and amphibian species observed at the type locality include Cnemi- dophorus inornarus, C. septem vittarus, Coleonyx brevis, Cophosaurus rexanus, Phrynosoma modes- turn. Ura stansburiana, Scaphiopus couchii, and an undescribed species of Sceloporus similar to S. jar- rovii cyanostictus. A third species of Cnernidophonts (possibly C. marmoratus) is also present. Crotaphytus antiquus are abundant and I observed more than 25 individuals in an area of about 1.5 km in length and roughly 200 m in width. This species usually runs quadrupedally, but was observed to use bipedal locomotion on occasion. They
72 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32 are able to take off bipedally from a standing start, as are all other Crotaphytus species. This species appears to be territorial, which is the case for all other Crotaphytus that have been studied (Fitch, 1956; Moehn, 1976; Montanucci, 197 1; Sanborn and Loomis, 1979; Yedlin and Ferguson, 1973; plus numerous additional references). On 25 June 1994, I witnessed apparent temtorial behavior when an adult male chased another adult male over approx- imately 10 m after the first male ventured into the area occupied by the second male. During the in- teraction, the pursuing male appeared to have its gular pouch fully depressed, a behavior that appears to be associated with aggression in all Crotaphytus species (Fitch, 1956; Sanborn and Loomis, 1979; personal observation). Very little is known about the reproductive behavior of this species. However, since all but one of the females observed displayed orange gravid coloration in various stages of intensity, it is clear that the reproductive cycle includes late June. One of the females bearing gravid coloration appeared emaci- ated, as if she had just oviposited. No juveniles were observed, suggesting that the year's early clutches had not yet hatched. Some individuals (TNHC 53 154, 53 159) contained yolked ovarian follicles together with corpora lutea and distended, vascu- larized oviducts, suggesting that this species can pro- duce at least two clutches in a single reproductive season. One large female (SVL = 89 mm) contained four shelled eggs, another (SVL = 89 mm) contained three shelled eggs, and four additional females contained between one and four yolked ovarian follicles, suggesting that the species has a relatively small clutch size. The only observation made regarding feeding habits is that one adult male that was prepared as a skeleton contained the remains of an unidentified coleopteran insect. Illustrations. -Black-and-white photographs of males and females were provided by Axtell and Webb (1 995). Crotaphytus bicinctores Smith and Tanner (Fig. 32A) Croraphyrus collaris bicinctores Smith and Tanner, 1972:27; fig. 1.2. Type Iacality: "Mercury Pass, Nevada Test Site, Nye Co., Nevada" (holotype: BYU 23883). Croraphyrus insularis bicinctores- Axtell, 1972:72 1; fig. 2, S b , 6. Crotaphyrus bicincrores-Sanborn and Loomis, 1979: 105. Etymology.-From the Latin bi. two, and cinct, banded or girdled, in reference to "the divided banding (presumably of the collar) in the Great Basin populations" (fide Tanner, personal communication, 1993). Diagnosis. - Crotaphytus bicinctores can be distinguished from C. reticulatus. C. collaris, C. ne- brim, and C. dickersonae by the absence of black oral melanin. It can be further distinguished from C. reticulat us, C. collaris, and C. nebrius by the presence in adult males of a strongly laterally com- pressed tail with a pale white dorsal caudal stripe, enlarged dark brown or black inguinal patches that extend between one-third and two-thirds of the dis- tance between the hindlimb and forelimb insertions, and a pale tan or off-white patternless region on the dorsal surface of the head. It may be further distinguished from C. reticulatus as well as C. antiquus by a dorsal body pattern of white spots and dashes on a brown field rather than white reticulations on a gold, tan, or brown field. It may be further distinguished from C. nebrius by its brown dorsal coloration rather than pale tan. It may be further distinguished from C. collaris by the presence of dark brown or black pigmentation in the gular fold (= ventrally complete anterior collar). It may be distinguished from C. grismeri, C. vestigium, and C. insularis by the presence of broad tan or buff transverse dorsal body bands. It may be further distinguished from C. grismeri by the absence ofa green- ish tint to the white bar that separates the collars, by a pattern of white reticulations on a brown field on the forelimbsand hindlimbs rather than a pattern of yellow forelimbs with minute brown spotting on the proximal dorsal surface of the brachium and a hindlimb coloration that is nearly patternless yellow with scattered minute brown spots from the distal thigh to the distal terminus of the limb, by the absence of a pale orange tail coloration in subadult females, and by the absence of a well-defined pale tan dorsal caudal stripe in juveniles of both sexes. It may be further distinguished from C. insularis and C. vestigium by the presence of a dorsally complete or narrowly separated posterior collar rather than a posterior collar that is broadly separated dorsally or completely absent. It can be further distinguished from C. insularis by the presence of a relatively broad nasal process of the premaxilla, the absence of olive green ventrolateral coloration in adult males, the presence ofa pattern of small white spots and dashes (occasionally transverse bands), rather than a pattern ofthicker, elongate white dash-
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