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of the Tucson Valley in the Santa Catalina Mountains). It is very likely that they occur in the remaining mountain ranges south of the Gila River, although the Baboquivari Mountains may be inhabited by C. collaris (Axtell and Montanucci, 1977). Crotaphytus bicinctores generally skins the northern border of C. nebrius' range on the north side of the Gila River, but crosses the river at the Sentinel Plain. a region uninhabited by C. nebrius, as no Sonoran mountain ranges project northward into this area. In Sonora, Cro~aphyfus nebritls occurs in the transversely oriented foothills that follow the United States-Mexico border along Mexican Highway 2 (the Pinacate Region). The north-south trending ranges of southwestern Arizona project northward from these foothills and probably provide the corridor through which C. nebritcs entered these mountains. They have been collected from several mountain ranges to the south and east in northern Sonora including the Sierra Cubabi, Sierra La Gloria, Sierra El Alamo, and Sierra El Rajon. One specimen is known from either the Sierra Cibuta or Sierra El Pinto (AMNH 73758, 25.6 km S Nogales), a more eastern locality in the northern foothills of the Sierra Madre Occidental. There is a relatively large gap in the known distribution of the species between the Caborca region (Sierra El Rajon) and the Hermosillo region. However, a series of specimens are known from the foothills between Hermosillo and the Guaymas region. Finally, the remaining specimens have been taken from the foothills of the Sierra Madre Occidental, in a series of north-south trending valleys separated by presumably uninhabitable densely vegetated mountain ranges. It is likely that C. nebrius reached these localities by way of major river drainages entering from the south, such as the Rio Sonora and Rio Yaqui, as suitable open habitat appears to be restricted to these drainage systems. Populations of Crofaphytus nebrius are only narrowly separated from those of C. bicincrores at two localities and in both cases the bamer that prevents contact is the Gila River. Crotaphytus nebrius occurs on the northern edge of the Gila Mountains and is separated from a population of C. bicincrores in the Laguna Mountains approximately 0.4 km to the north on the opposite side of the Gila River. Similarly, C. nebrius occurs on the western margin of the Buckeye Hills, while C. bicinctores occurs on the extreme eastern margin of the Gila Bend Mountains only a few hundred meters to the west on the opposite shore of the Gila River. Thus, C. nebrius may be observed on the east side of the Gillespie Bridge OF CROTAPHYTID LIZARDS and C. bicinclores can be observed moments later on the west side. Several questions remain regarding the distribution of C. nebrius. First, C. nebrius occurs as far nonh as 1 1.7 km N Huasabas and 19.5 km N Bacadehuachi in the Sierra Madre Occidental, while C. collaris is known from as far south as the Bavispe Region, approximately 60 km to the nonh. It is unknown whether this gap is real or an artifact of collecting. The habitat in the Huasabas and Bacadehuachi regions appears to be marginal and the presence of higher elevation mountains between this area and the Bavispe region strongly suggests that a contact zone does not exist here. However, this remains to be substantiated with additional field studies. Second, a specimen of C. bicincrores was observed by the author at Black Gap, Maricopa County, Arizona, a narrow pass on the western periphery of the Sauceda Mountains through which Arizona State Highway 85 passes. This observation was extremely surprising given that this area is apparently well isolated from known C. bicincrores populations north of the Gila Bend River and on the Sentinel Plain. If C. bicinctores has an established population at this locality, it is likely that C. nebrius and C. bicinctores contact somewhere in the Sauceda or Maricopa mountains. Several later attempts to find C. bicincrores or C. nebrius at this locality were unsuccessful. Fossil Record. -Van Devender and Mead ( 1 978) referred a maxilla and dentary from late Pleistocene deposits in the Tucson Mountains and Wolcott Peak, Pima County, Arizona, to Crotaphy~us collaris. Van Devender et al. (1991) referred dentary, maxillae, and tooth crown material from late Pleistocene deposits in Organ Pipe Cactus National Monument to either C. collaris or C. insularis. Because the Tucson Mountains and Organ Pipe Cactus National Monument are currently inhabited by C. nebrius, this material probably should be referred to C. nebrius on distributional grounds. NatltralHislory. -Nothing has been published regarding the natural history of this species but I have made the following observations. Croraphj)rics nebrius occurs in a diversity of habitats, although always in association with rocks. In the northern portion of its range it may be found in extremely xeric habitats characterized by granitic outcroppings or volcanic flows. In the southern portion of its range, it is occurs in rocky areas often with relatively dense arid-tropical thornscrub vegetation. In these areas, C. nebritcs may be concentrated in arroyo bottoms
BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32 and less vegetated stream and river valleys. In the Crotaphytus oligocenicttst Holman northwestem portion of its range, the species is often found perched on granitic rocks that lay in sandy washes at the bases of rocky hillsides. No observations have been made with respect to the feeding habits of this species although it is likely ~ ~ ~ r oligoceniclcc o p h ~ Holman, ~ ~ 1972: 16 13. T~~~ locality: "From earlv . Olieocene. - . Cvoress -. Hills Fonnation. north branch - ~ ~ of Calf Creek, in L. S. 4, Sec. 8, twp. 8, range 22, W. 3rd mer., elevation 3600 ft (I 100 m)" (holotype: Saskatchewan Museum of Natural History number 1444). that arthropods and small lizards make up the bulk of the diet as in other Crotaphytus species. Efymology.-Named in reference to the time period during which these lizards lived. The activity period for the species may extend between March and at least late September. Adult and subadult males were observed on 19 March in the Buckeye Hills, Maricopa County, Arizona. and juveniles were observed north of Guaymas, Sonora, Mexico, on 27 March. Between 15 and 19 April 1992, juveniles and subadults that apparently had just emerged from hibernation (they were still encrusted with dirt) were observed in the western foothills of the Sierra Madre Occidental and at Quijotoa, Pima County, Arizona. On the same day that the Quijotoa subadults were observed, adult males were observed just north of Ajo (Pima County) and at Mohawk, Yuma County, Arizona. Adults are active at least as late as 1 I August and recently hatched neonates have been observed as late as 19 Septem- Distribution. -Known only from the type locality. Remarks. - Crotaphytus oligocenicust is an extinct species of Oligocene age known only from six dentaries collected at the type locality. Because of the fragmentary nature of the type material, it cannot be determined whether this species shares any of the crotaphytid synapomorphies presented here. Thus, I agree with Estes (1983) in questioning whether this species is in fact a crotaphytid. However, given that no data were discovered in this analysis either supporting or rejecting the placement of this species within Crotaphytidae, no taxonomic rearrangements are herein suggested. A black-andwhite illustration of the holotype material (a right dentary) is given in Holman (1972). ber. It seems likely that adults extend their activities at least into September and juveniles into October or November. Reproductive behavior appears to be typical of the genus. On 14 June 199 1 mating was observed in the Gila Mountains, Yuma County, Arizona. The Crotaphytus reticulatus Baird (Fig. 30C) Crotaphytus micularus Baird, 1858:253. Type locality: laredo and Ringgold Barracks, Starr County, Texas-(Smith and Taylor, 1950): "Laredo"; (Cachran, 196 1 ) "Ringgold Barracks, Montague County, Texas"; (Montanucci, 1976): "Fort Ringmale was observed to grasp the female by a fold of skin of the neck during coitus. The female offered no resistance and thus appeared to be fully receptive. Interestingly, the female bore fully developed gravid gold Military Reservation (- Ringgold Barracks), Starr County, Texas" (lectotype Montanucci, 1976: USNM 2692A). Croraphyrus (Crotaphyrus) reticularus- Weiner and Smith. 1965: 187. coloration, which is consistent with observations made by Montanucci (1965) that this coloration may not deter copulation in Gambelia silus, at least with females that do not display rejection behavior. It Efymo1ogy.-From the Latin reticulatus, made like a net. In reference to the net-like dorsal and gular pattern of white reticulations present in this species. Diagnosis. - Crotaphytus reticulatus can be distherefore seems likely that mating takes place pri- tinguished from all other species of Crotaphytus exmarily in May or June. Recently emergent neonates cept C. antiquus by the presence of an adult color have been observed on 1 1 August in the Silverbell pattern consisting of white reticulations, some of Mountains, Pima County, Arizona, and on 19 Sep- which enclose black pigmentation, and the presence tember in the Gila Mountains. Neonates collected of jet black femoral pores in males. It can be disin the Silverbell Mountains were as small as 44 mm tinguished from C. antiquw by the dorsal coloration SVL and the individual collected in the Gila Moun- of golden tan rather than dark brown and by the tains was 42 mm SVL and still retained a small presence of black pigments in only a subset of the portion of the umbilicus. Thus, neonates appear to hatch out between July and/or August and Septemdorsal body reticulations rather than in all or nearly all of them. It can be further distinguished from C. ber, at least in the northern portion of the range. collaris by the presence of dark brown or black pig- Illustrations. -A black-and-white photograph ap- mentation in the gular fold (= ventrally complete pears in Axtell and Montanucci (1 977). A color pho- anterior collar) in adult males. It can be further distograph of a gravid female was provided in Sprack- tinguished from C. antiquus, C. nebrius, C. dickerland (1 993). sonae, C. grismeri, C. bicinctores, C. insularis, and ~ ~ - - - -
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