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1996 McGUIRE-SYSTEMATICS OF CROTAPHYTID LIZARDS 49 Fig. 35.-Ventral view of a series of adult male Crotaphytus vesrigium. collar. In C. bicinctores and C. grismeri, the posterior collar terminates within the antehumeral fold. In C. vestigium and C. insularis, the posterior collar almost always terminates before reaching the antehumeral fold. The extreme situation exists in C. insularis where, in the few individuals that have a posterior collar, it terminates just before reaching the antehumeral fold. This character is less consis- tent in females, especially with respect to C. reticulatus, in which females either lack collars or have them poorly developed. The four conditions described above were coded as separate character states of an unordered multistate character (state 0 = collar extends well out onto dorsal surface of brachium, state 1 = collar just reaches forelimb insertion, state 2 = collar terminates within antehumeral fold, state 3 = collar terminates before entering antehumeral fold). Again, because the nearest outgroups lack collars, this character was left unpolarized. As stated above, all Crotaphytus species are char- acterized by the presence of at least one collar (but see C. insularis below). In fact, with few exceptions, all Crotaphytus species except C. insularis and fe- male C. reticulatus have two collars. Crotaphytus insularis almost always have only the anterior collar, the posterior collar having apparently been lost (Fig. 32D). The fact that five specimens (CAS 21948, 50879,86754, 148652; SDSNH 53064) have an extremely reduced, but visible, posterior collar is con- sistent with the hypothesis that collar reduction has occurred in this species. Males have a more densely pigmented anterior collar than females, which sometimes have no collar at all. This reduction in both the posterior and anterior collars appears to be derived and hence an autapomorphy for this in- sular species. In C. reticulatus females, the anterior collar marking may be lacking while the posterior collars remain. However, the posterior collar marking in both sexes of this species is often little more than a slightly modified band of black-filled reticulations. This variation was not included in the phy- logenetic analysis because of the potential problem of lack of independence between this state and the wide separation of the posterior collars described above. Dark Nuchal Spots (Character 76; Fig. 36).-A pair of black or dark spots usually occurs between the dorsal extensions of the anterior collar markings in Crotaphytus reticulatus (39 of 5 l), C. antiquus (16 of 16), and C. collaris (58 of 73, and are oc-
BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32 casionally present in C. nebrim (I 2 of 5 I). The spots, which occur between the dorsal extensions of the anterior collar markings, appear to be homologous with the first transversely arranged row of blackfilled hexagonal reticulations seen in C. reticulatus. In C. antiquus, the nuchal spots are always present, but often incompletely separated from the remainder of the anterior collar markings. Black nuchal spots are not present in the outgroup taxa and their presence is coded as the derived state. Ingitinal Patches (Characters 77,78; Fig. 32C, 34, 35).-In several species of Crotaphytus, adult males develop dark brown or black ventral patches in the inguinal region. These patches vary considerably in size with C. bicinctores, C. dickersonae, C. grismeri, C. insularis, and C. vesrigiittn having large patches and C. antiquus, C. nebrius, and C. collaris having smaller ones. All adult male C. bicinctores, C. antiquus, C. dickersonae, C. grismeri, C. insularis, C. nebrius, and C. vesrigiurn develop these patches while only some C. collaris have them. Interestingly, only C. collaris from the western periphery of its range (in the area usually referred to the subspecies C. c. baileyi) are known to have inguinal patches. Thus, there are at least two characters associated with inguinal patches: size of the patches and the frequency with which they occur. Homology of the patches seems likely. Both large and small patches begin development as small ventral spots near the hind limb insertion and the large patches differ only in that they continue to become larger (and probably grow faster). Inguinal patches of the type present in some Crolaphj)rr& are extremely rare in the outgroup taxa. Similar markings are present in Uma exsuland LI. paraphygas (de Queiroz, 1989; although they occur more laterally than in Crotaphyrus), Ufa nolascensis, Uromasryx hardwickii (concentrated on the thigh), and Enyalius iheringii (again, more laterally oriented). This character has been coded two ways: first, as a binary character with the absence of inguinal patches (of any size) as state 0 and the presence ofpatches as state 1 ; and secondly, as a separate binary character with the presence of small patches as state 0 and the presence of large patches as state 1. Taxa without inguinal patches were scored as unknown ("?') for this second character. Because the first character (77) considers the frequency in which patches are present, the second character (78) does not take frequency into consideration. For character 78, the presence of small patches in any frequency is assigned state 0 and the presence of large patches in any frequency is coded as state 1. Fetnoral Pore Secreriotu (Character 79; Fig. 22, 23, 33-35).-The femoral pore secretions of male Crotaphyt us retictrlattrs and C. anriquus are jet black. Unlike other Crotaphytus species, such as C. nebrim, which often have grayish secretions, the sub- cutaneous glands themselves are also jet black. This condition was not observed in other species of ig- uanian lizards and is treated as the derived state. Gular Pattern (Characters 80-82; Fig. 33-35).- There is much variation in the gular pattern of male Crotaphytus, especially in the wide-ranging species C. collaris. However, the general arrangement of the gular colors is similar in all of the species. For ex- ample, each has a relatively uniformly colored central gular region that is surrounded by a peripheral reticulated or spotted pattern superficial to the man- dibles. It is in the context of this general pattern that the following discussion of variation is based. Be- cause the pattern and extent of the gular coloration is sexually dichromatic, the following discussion pertains only to adult male Crotaphyrus. Adult male Crotaphytus bicinctores, C. antiquus, C. dickersonae, C. grismeri, C. insularis. C. rericu- lattrs, and C. vestigiunt (Fig. 35) have a patch of black pigment in the posteromedial portion of the gular region. This pigmentation corresponds with that portion of the gular pouch that is depressed by the second ceratobranchials of the hyoid apparatus, and thus presumably increases the visibility of the depressed gular pouch during aggressive display. The black patch is continuous with the black pigrnen- tation of the gular fold and the ontogenetic development of the gular patch suggests that it may be an extension of the gular fold coloration. However, the presence of black pigmentation in the gular fold and in the posteromedial portion of the central gular region are treated as separate characters because the presence of black pigmentation in the gular fold is not always associated with a black central gular patch (e.g., C. nebrius). Because the outgroups do not have a gular pattern that is similar to Crotaphytus, outgroup analysis cannot be utilized to assess polarity. Therefore, this character was left unpolarized. There is considerable variation in the peripheral gular pattern of Croraphytus as well. In all Crora- phytus except C. nebrius, the peripheral gular pattern is composed of white reticulations on a dark field. In most C. collaris, the dark pigmentation within each subquadrate reticulation has a light center, which results in a pattern reminiscent of the dorsal pattern of a jaguar. A similar pattern is sometimes evident in other species, such as C. bicinctores. In
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