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BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY<br />

Fig. 17. -Ventral view of the fifth metatarsal bone of Crotaphytw<br />

collaris showing the contact of the medial and lateral plantar<br />

tubercles forming an arch (redrawn from Snyder, 1954).<br />

may be referring to ventromedial curvature). They<br />

may have been refemng to the presence of a pro-<br />

portionally shorter and broader pelvic girdle in east-<br />

em populations of Crotaphytus collaris (the only<br />

representative of the "collariform" group that they<br />

examined) than in other Crotaphytus species or<br />

Gambelia. This difference appears to be related, at<br />

least in part, to modification of the pubic rami, which<br />

are nearly transverse in orientation, rather than<br />

acutely angled anteriorly. However, the condition<br />

in the remaining populations of C. collaris (formerly<br />

referred to C. c. fuscus, C. c. baileyi, and C. c. au-<br />

riceps) appears to be intermediate in each of these<br />

features. Coding of this variation is further com-<br />

plicated by individual variation in pelvic girdle<br />

structure, such that some individuals approach the<br />

eastern C. collaris condition, while others approach<br />

the condition of other Crotaphytus species. Short,<br />

broad pelvic girdles are often observed in crevice-<br />

dwelling species (e.g., Sauromalus) and the rela-<br />

tively short, broad, pelvic girdles of eastern C. col-<br />

laris may be related to the crevice-dwelling behavior<br />

observed in these lizards.<br />

LIMBS<br />

(Character 47; Fig. 17)<br />

On the plantar surface of the fifth metatarsal are<br />

two large tubercles termed the medial and lateral<br />

plantar tubercles by Robinson (1975). These tuber-<br />

cles serve as attachment points for the tendons of<br />

M. gastrocnemius. In the majority of iguanian spe-<br />

cies, a groove runs between the two tubercles and a<br />

NO. 32<br />

tendon of M. flexor digitorum longus passes within<br />

it (Robinson, 1975). In Crotaphytus. the medial<br />

plantar tubercle usually curves laterally such that it<br />

contacts the lateral plantar tubercle forming a complete<br />

arch (Fig. 17), through which passes the tendon<br />

of M. flexor digitorum longus (noted and figured by<br />

Snyder, 1954). The contact ofthe tubercles is usually<br />

extensive and in some individuals, the tubercles may<br />

fuse completely. The arch condition was absent in<br />

the entire available series of Gambelia (41 specimens)<br />

and, in adults, it was always present in the<br />

20 C. bicinctores, four C. antiquus. 12 C. dickersonae,<br />

five C. gristneri, and 22 C. vestigium examined.<br />

It was complete on at least one pes in 28 of 36 C.<br />

collaris, three of five C. insularis, i 1 of 12 C. nebrius,<br />

and six of seven C. reticulatus. The majority of specimens<br />

that lacked the complete arch were juveniles,<br />

and in most cases the gap between the medial and<br />

lateral plantar tubercles was narrow. Therefore, this<br />

character was scored only for adults. Among the<br />

outgroup taxa examined, the arched form of the<br />

medial and lateral plantar tubercles was present only<br />

in the phrynosomatid sand lizards (Uma, Callisau-<br />

rtcs, Cophosaurus, and Holbrookia). This feature ap-<br />

pears to represent a synapomorphy for Crotaphytus,<br />

as well as providing additional character support for<br />

the monophyly of the phrynosomatid sand lizards.<br />

The hindlimb of Crotaphytus is much longer than<br />

that of Gambelia of similar SVL. A relatively long<br />

hindlimb is typical of lizard species that utilize bi-<br />

pedal locomotion (Snyder, 1952, 1954, 1962), al-<br />

though agamines provide an interesting exception.<br />

Much of the variation in hindlimb length between<br />

Crotaphytus and Gambelia is realized in the longer<br />

crus of the former, while the pes appears to be of<br />

relatively similar length. Although a greater relative<br />

hindlimb length appears to be a derived character-<br />

istic of Crotaphytus, there is great variation in the<br />

outgroup taxa and this feature was not included in<br />

the phylogenetic analysis.<br />

The dorsal body squamation of Crotaphytus and<br />

Gambelia is remarkably similar in that both genera<br />

are characterized by relatively undifferentiated head<br />

scales and fine homogeneous dorsal body squama-<br />

tion. However, despite many similarities in scale<br />

patterns and scale sizes on the various regions of<br />

the body, phylogenetically useful variation in squa-<br />

mation exists. A more detailed description of the<br />

squamation of crotaphytids is provided in the tax-<br />

onomic accounts of the family, genera, and species.

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