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McGUIRE-SYSTEMATICS OF CROTAPHYTID LIZARDS 4 7 are deposited in the oral mucosa and at least some of the underlying fascia of the M. adductor mandibulae complex. There is interspecific variation in the extent of the pigmentation as well. In Garnbelia, C. collaris, C. rtebrirts, and C. rcticctlarrts, the coverage and density of the oral melanin is extensive. The pigments are present on the floor of the buccal cavity as well as on the fauces of the roof of the cavity. In C. a~ttiqutts and C. dickersortae, black oral melanin is present but it is less extensive in both coverage and density. Stebbins (1954) noted that G. )vislizenii from the Painted Desert region of Arizona may lack this coloration. However, this observation has not been confirmed in the present study and Stebbins himself (personal communication, 199 1) does not recall where he obtained this information. A black oral mucosa appears to bc absent from all basal outgroup taxa outside of the family Polychrotidae (the throat lining is deep violet in Tropidtinis urztbra, Etheridge, 1970). Within Polychrotidae, black oral melanin is present in some Polychrus (P. ntar~~torattts, P. actttirostris), Pristidactyltts volcanensis, Leiosartrcts catarnarcensis, Lb-ostrophtis vaautieri (Etheridge and Williams, 199 I), Anisolepis Fig. 33.-Ventral view of a scries of Cromaphytus collaris. grilli (Etheridge and Williams, 199 1 ), I'lccwacosattri4s heteroderrnis, and all three species of Cltantaeleolis (Schwanz and Henderson, 199 1). It is variably present in Pristidactj~lrts torqrtatts. The absence of black oral melanin has been verified in Pol~!chrci.s liogasrer, P. prttaroscts. Pristidactj~lus acltaleruis, P. scapt4lattts, P. casuhatiensis. Leiosairrrrs beNi, U. gallardoi, Enyaliits bilirteatus, E. braziliertesis. E. catertatlis, E. iheringii, E. perdittis, and E. picttts. Although black oral melanin may prove to be the ancestral condition for Polychrotidae, the family does not appear to be the sister taxon of Crotaphytidae (Frost and Etheridge, 1 989). Therefore, the presence of black oral melanin is treated as the derived state. Collars (Characters 72-75; Fig. 30-36). -A]though all Crotaphj~trts species are characterized by the presence of at least one collar, there is considerable interspecific variation. Most species have two well-developed black collars, with relatively thick white borders that encircle or partially encircle them. The collar configurations of Crotaphj~tcts reticttlattcs and C. antiqults suggest that the transversely arranged series of black spots (each of which is bordered with white) present in these species may have
4 8 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32 been the precursor to the black collars outlined in white that are found in all Crotaphytus species. This is especially evident in the posterior collar markings, which in C. reticulatus are usually little more than a few closely approximating black spots with white borders. Furthermore, in most individuals there are dark pigments bleeding into the intervening areas between the black spots. A similar situation is sometimes present in the anterior collar as well. With respect to the outgroup taxa, it is unlikely that a white-bordered collar or pair of collars is the ancestral state in all but Opluridae (collars present in 0. cuvieri and 0. cyclurus). Therefore, the presence of white-bordered collars is treated as the derived state. Additional variation occurs in C. bicinctores, C anfiquus, C. collaris, C. dickersonae, C'. grismeri, and C. nebrius, where the posterior collars are either in contact or only narrowly separated at their medial margins (Fig. 30-32, 36). In C. irtsularis and C. vesfigiurn, the posterior collars are broadly separated (Fig. 32C, D). The condition in C. reticirlarrrsis more difficult to interpret because of the weak development of the posterior collar and it is tentatively coded as widely separated. Because Gambelia, the nearest outgroup to Crotaphytus, lacks collars, this character was left unpolarized. Fig. 34.-Ventral view of an adult male Crotaphytus nehrircs. In all adult male Crotaphytus except C. collaris, the anterior collar is complete ventrally by way of dark brown or black pigmentation within the trans- verse gular fold (Fig. 33-35). Because the nearest outgroup taxa lack collars, this character was left unpolarized. In Crotaphytus collaris and C. nebrius, the posterior collar passes through the antehumeral fold before reaching the proximal dorsal surface of the brachium. A less developed condilion usually occurs in C. reticulatus, where the collar passes through the extensive antehumeral mite pocket and isolated black patches may extend a short distance onto the proximal dorsal surface of the brachium. In C. an- tiqtrus, the posterior collar marking of males either terminates at the forelimb insertion or melanic spots extend onto the brachium, while in females, the collar marking generally terminates before entering the antehumeral fold (although in one individual [MZFC 67551, the marking seems to continue through much of the underlying mite pocket). In C. dickersonae, the posterior collar just reaches the dorsal surface of the forelimb insertion and may extend slightly onto the brachium as in C. reticulatus. How- ever, the collar marking does not pass through the antehumeral fold in this species because the antehumeral fold terminates anterior to the posterior
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