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McGUIRE-SYSTEMATICS OF CROTAPHYTID LIZARDS Fig. 29.-A postfemonl mite pocket in a juvenile Croraphytus hicinctores. humeral fold was not observed in any of the outgroup taxa examined and, thus, appears to be unique to Croraphyttrs, excluding C. dickersonae. As discussed above, the antehumeral fold of C. dickersonae terminates further anteriorly than in any other crotaphytid, usually failing to reach the forelimb insertion, which probably explains the absence of an antehumeral mite pocket in this species. Pos~emoral Mire Pockets (Character 58; Fig. 29).-In most crotaphytids, subdermal mite pockets are present at the posterodorsal border of the hindlimb insertion where a patch of finely scaled or unscaled skin dips inward between M. iliofibularis and M. iliofemoralis. These pockets usually are inhabited by trombiculid mite larvae and occasionally ticks. Arnold (1 986) noted that mite pockets, which may occur in a variety of anatomical regions, often vary both intra- and interspecifically in terms of their presence, degree of development (e.g., depth), and in the nature of their squamation, and in this respect Crotaphytidae is no exception. However, pockets were absent only in Crorapl~us reticularus and occasionally in C. collaris and C. nebrius. In Croraphytus, the depth of the mite pocket may be correlated with the degree of development of the antehumeral mite fold. For example, in C. reficulatrrs, which lacks the postfemoral pocket, the mite pockets of the antehumeral fold (discussed above) are strongly developed. In contrast, the mite pockets of the antehumeral fold are absent in C. dickersonae. while the postfemoral pockets are the most strongly developed (deepest) of all Crotaph))rus. Postfemoral mite pockets are not unique to Crotaphytidae. Smith (1 939) noted that they are present in seven species of Sceloporucs, including the five species in his S. variabilis group, as well as S. macltlosus and S. gadoviae. Shallow postfemoral pockets were also observed in Uta squamara and U. paltneri, but not other Ura. Although not observed here, shallow mite pockets are occasionally present in several species of Urosaurus (J. Wiens, personal communication, 1994). However, the absence of postfemoral pockets in Pf?r)mosoma, the sand lizards, Petrosaurus, most Uta (in those species that lack pockets, mites may accumulate in the postfemoral region, but an obvious subdermal pocket is lacking), most Urosaurus. Sator, as well as most Sceloporus, suggests that the pockets observed in subsets of Uro-
BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32 sattrus. Uta. and Sceloporus are not homologous with crotaphytid postfemoral pockets. Most Stenocercus and at least two species for- merly referred to Ophryoessoides (S. ornatus and S. trachycephalus) have postfemoral mite pockets (Fritts, 1 974; Arnold, 1 986). However, the postfemoral pocket of those Stenocercus species examined here (S. trachycephaltrs. S. chrysopygus, S. guenth- eri, S. imitator, S. roseiventris) occurs as a vertical fold along the lateral body wall immediately pos- terior to the hindlimb insertion and, thus, does not appear to be homologous with the poslfemoral mite pocket of crotaphytids. Furthermore, postfemoral mite pockets appear to be absent from the basal lineages of Liolaeminae (Phymaturus and Ctenob- lepharys: species examined include Ctenoblepharys adspersus, Phyrnaturus sp., P. pallurna, P. patagon- icus. P. punae), Leiocephalinae (G. Pregill, personal communication, 1993; verified in Leiocephalus car- inatus, L. inaguae, L. macropus, L. melanochlorus, L. pratensis [folds present, but no reduction in squamation], L. psammodrotnus, L. schreibersi), and Tropidurini (Uranoscodon strperciliosus. Plesiotni- crolophus koepkeorum, Microlophus theresioides, M. tigris. M. stolzmani, personal observation). Thus, the postfemoral mite pockets of certain members of the Stenocercini are considered to be nonhomolo- gous with crotaphytid postfemoral mite pockets. Several oplurids have postfemoral mite pockets that appear to be structurally identical with those of crotaphytids. That is, the pocket occurs as an invagination between M. iliofibularis and M. iliofemoralis. Arnold (1 986) noted the presence of postfemoral mite pockets in Oplurus cuvieri and 0. cyclurus and I have observed them in 0. cyclurus, as well as in 0. saxicola, O.fierinensis, and C/talaradon madagascariensis. Postfemoral mite pockets appear to be absent in 0. quadrimaculatus. Because we have no hypothesis of phylogenetic relationships for oplurids, it is not possible to say whether the pockets are derived within the family or were present an- cestrally. Therefore, the possibility that postfemoral mite pockets were present in the common ancestor of Opluridae cannot be discounted. Among iguanids, Dipsosaurus dorsalis has a weakly developed postfemoral pocket that occurs in the same anatomical position as the postfemoral mite pocket of crotaphytids. Because Dipsosaurus (along with the fossil species Arrnandisaurus exploratod) is the sister taxon of the remaining iguanids (de Queiroz, 1987; Norell and de Queiroz, 199 l), the possibility that postfemoral pockets were present in the common ancestor of Iguanidae cannot be elim- inated. Postfemoral mite pockets appear to be absent from Corytophanidae, Hoplocercidae, Chamaeleonidae, and Polychrotidae, although all of their constituent species have not been examined. Although postfemoral mite pockets may have been present in the common ancestors of the families Opluridae and Iguanidae, their presence is most parsimoniously treated as the derived state for Crotaphytidae. Hemipenes (Character 59). - Hemipenes were examined for all of the crotaphytid species except Cro- taphytus reticularus. The hemipenes ofcrotaphytids are bulbous and weakly bilobed with a short median fissure separating the two lobes apically. The sulcus spermaticus is covered by a large fleshy flap of in- tegument that folds over it from its lateral margin. This fold does not project directly toward the apex but rather extends laterally toward the outer margin of the lateral lobe. The sulcus spermaticus itself appears to terminate in a broad, shallow depression at the base of the lobes. The entire sulcate surface of the hemipenis is only weakly ornamented with a fine papillate or dimpled texture. Immediately outside of the sulcus spermaticus, the surface is ornamented with plicae that are continuous with those of the asulcate surface. Distally, the lateral surfaces of the lobes bear small knob-like processes that are covered with extremely fine calyculae. The base of the asulcate surface of the hemipenis is naked. More distally, ornamentation is present in the form of plicae and calyculae. The proximal lateral surfaces of the ornamented region of the hemipenis are covered by fine plicae. These plicae grade medially into calyculae and this calyculate zone extends distally toward the apex of the hemipenis where it spreads laterally. As a result, the entire surface of the hemipenis distal to the median apical fissure is ornamented with minute calyces. The lateral surface of each lobe bears a shallow depression ornamented with extremely fine calyces. The calyces reach their smallest sizes here and in the apical region of the hemipenis. The only obvious difference between the hemipenes of Crotaphytus and those of Gambelia is in their relative size. The hemipenes of Gambelia are roughly twice the size of those of similar-sized Cro- taphytus. Although the hemipenes of Crotaphytus
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