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Fig. 18.-Squamation of the dorsal portion of the head of Cro- raphyrus collaris (USNM 17 183. adult male). Scale = 5 mm. Rostra1 Scale (Character 48).- In all crotaphytids except Crotaphytus dickersonae, the rostral scale is approximately four times wider than high. In C. dickersonae, the rostral is less elongate and approximately two times wider than high. There is much variation in the outgroups, although most taxa have a rostral that is much wider than high. Consequent- ly, this character was left unpolarized. Siipraorbital Semicircles (Character 49; Fig. 18, 19). - Crotaphytus have supraorbital semicircles composed of scales that are much larger than the adjacent supraoculars. In Gatnbelia, obvious supraorbital semicircles are absent, with the supraoculars tending to grade into the frontal series. The outgroups vary considerably in the presence of discrete supraorbital semicircles. They are present in all oplurids and polychrotids examined (except Cha- rnaeleolis), and variable within the remaining fam- ilies. Within Hoplocercidae, they are absent in En- yalioides laticeps, but present in E. praestabilis and E. oshaugnessyi. Within Phrynosomatidae, they are present in Petrosaurus and the Sceloporus group, Lrt?ia notata. U. scoparia, and U. inornata, but absent in Phrynosoma and Uma exsul. In tropidurids, they are present in some Pttyrnaturus patagonicus, Leiocephalus, Liolaemus, Stenocercini, basal Tro- pidurini (except Uranoscodon stcperciliosus), and absent in Ctenoblepharys adspersics, most Phyma- OF CROTAPHYTID LIZARDS Fig. 19.-Squamation of the dorsal portion of the head of Gani- bclia ~vislizcnii (SDSNH 68662, adult fcmale). Scale = 5 mm. turtis, and Uranoscodon superciliosus. In iguanids, they are present in Dipsosaurus, absent in Brachy- lophus fasciatus, and generally absent in the remaining taxa. In chamaeleonids, they are absent in Hydrosaicrus pusttclatus, Leiolepis belliana. Uro- mastyx loricatus, and U. ocellatus, variable in U. geyrii, U. rnicrolepis, and U. acanfhinurus, and present in U. aegypticus, U. asmtcssi, U. hardwickii, U. rnacfadyeni, U. philbyi, and U. fhontasi. In corytophanids, they are present in t3asiliscus plumifrons, B. vitatrtcs, Corytophanes hernandczi, absent in C. cristatus and C. percarinattcs, and variable in Lae- manctics. Because of this extensive variation, this character was left unpolarized. Subociilars (Character 50; Fig. 20, 21).-In Cro- taphyttcs, the suboculars are subquadrate, with the third scale occasionally larger than the others, whereas in Garnbelia, the second subocular is four to five times larger than the others. Assessing po- larity of this feature is difficult because both states are widespread within the Iguania. At least one subocular is much longer than the others in phrynosomatids except Phrynosorna, the oplurids Oplurus
BULLETIN CARNEGIE MUSE .UM OF NATURAL HISTORY NO. 32 Fig. 20.-Squamation of the lateral portion of the head of Cro- taphytuscollaris (USNM 17183, adult male). Scale = 5 mm. saxicola. O.jierinensis, and 0. qttadrimaculattts, the chamaeleonid Leiolepis belliana, the tropidurids Phyrnaturus patagoniclrs, Leiocepltaltrs, Liolaernus, Stenocercini, Microlophtts, Plesio~~ticrolophus, and all but terminal Tropidurtcs (Frost, 1992), the iguanid Dipsosaurus, and the polychrotids Anisolepis, Pristidactylus, and Enyalitcs bilineatus. Multiple subequal suboculars are present in the oplurids Oplurus cyclurus, 0. ctrvieri, and Chalaradon, the chamaeleonids Uromastyx and Hydrosaurus pwtltlatus, hoplocercids, the tropidurids Phyrnaturtts punae. P. palluma, Ctenoblepharys, and Uranoscodon slcperciliosus, the polychrotids Urostrophus, Polychrus, Phenacosaurus, Charnaeleolis. Anolis, and Enyalius (except E. bilineatus), iguanids (except Dipsosaurus), and corytophanids. An elongate subocular appears to be the ancestral state in Phrynosomatidae, Tropiduridae, and Opluridae, and equivocal in Iguanidae, and Polychrotidae. The presence of subequal suboculars is the ancestral state for Corytophanidae, Hoplocercidae, and Chamaeleonidae. Therefore, this character could not be polarized. Terminal Supradigital Scales (Character 5 1). -In Gambelia, C. collaris, and C. reticulatus, the terminal supradigital scales nearly always lie flat against the dorsal surface of the claws. In the remaining Crotaphytus, the terminal supradigitals project dorsally such that each is elevated from the claw. A similar elevated condition occurs occasionally in various iguanians including the phrynosomatids Petrosaurus, Uta stansburiana (three of four), U. palmeri (one of four), and Uta sqitarnata (one of three), Fig. 2 I .-Squamation of the lateral portion of the head of Gam- hclia \visIizenii (SDSNH 68662, adult fcmale). Scale = 5 mm. the tropidurids Plesiomicrolophus koepkeorurn (one of four), Microlophus grayi (one of four), M. ther- esioides (one of four), M. tigris (one of four), and M. stolzntanni (three of four), and the hoplocercid En- yalioides laticeps (one of five). Despite this variation, the presence of elevated terminal supradigital scales is most parsimoniously considered lo be the derived state. Fenioral Pores (Characters 52, 53; Fig. 22, 23).- In Gambelia, the femoral pores extend distally at least to the inferior angle of the knee. The femoral pore series of G. silus usually just reaches this point, while the femoral pore series of G. tvislizenii and G. copei almost always extend beyond and may even arch posteriorly onto the lower leg. The femoral pore series of Crotaphytus does not reach the inferior angle of the knee and usually terminates well prox- imal to this point. Polarization of this character is complicated by the absence of femoral pores in the Tropiduridae, Opluridae, and Corytophanidae. However, in the remaining outgroups, the femoral pore series always terminates before reaching the inferior angle of the knee (Phrynosoma coronatun1 is variable with re- spect to this character). Therefore, the condition observed in Garnbelia is interpreted as the derived state. In Gambelia lvislizenii and G. copei, the femoral pores of females are large and contain substantial quantities of exudate, although the pores are usually slightly larger in males. In G. silus, Crotaphjaus, and all of the outgroup taxa examined that have femoral pores except Enyalioides laticeps, they are much larger and fuller in males than in females and, in- deed, in females the pores may be devoid ofexudate. Therefore, the condition observed in G. wislizenii and G. copei is considered to be the derived state.
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