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1996 McGUIRE-SYSTEMATICS OF CROTAPHYTID LIZARDS 2 1 - - Fig. 13.-Lingual view of the right mandible of (A) Croraphyrus Fig. 14.-hbial view of the right mandible of (A) Crotaphytw reticularus (REE 2912, adult male, SVL = 122 mm) and (B) rfliculallu (REE 291 2, adult male, SVL = 122 mm) and (B) Gatnbclia cowi (REE 2800, adult female. SVL = 123 mm). Ang Gamhflia coppi (REE 2800, adult female, SVL = 123 mm). Cor = angular, An - anicular. Cor = coronoid, Den = dentav, pmf = COronoid~ Den dcntav, PrC = preanicular, Sur = surangular. = = posterior mylohyoid foramen, Pre = preanicular, Spl = splenmm. ial. Sur = surangular. Scalc = 5 mm. completely encloses Meckel's cartilage. In crotaphytids, the tubular nature of the dentary is incomplete. The anterior end of the dentary is open, while posteriorly the groove is closed, but not fused. In Crotaphy~tu, with the exception of C. grismeri, the groove is usually closed over less than one-half of its length anterior to the splenial and there is relatively consistent interspecific variation in this characteristic. In C. collaris. C. nebrius. and C. reticulatus. Meckel's groove is often open over its entire length anterior to the splenial, and most of the remaining specimens have the groove closed over less than one-third of its length. In C. antiquus and C. dickrsonae, the groove is not open over its entire length, but as in the above-mentioned taxa, it was nearly always closed over less than one-third of its length. In C. bicinctores, C. insularis, and C. vestigiurn. Meckel's groove is usually closed over between one-third and one-half of its length anterior to the splenial and was only once observed to be open over its entire length (C, vestigittrn. REE 28 11). Crotaphytusgrismeri is unique among Crotaphytus in that Meckel's groove is closed over between approximately 50 percent and 70 percent of its length in all specimens examined (five of five). Norell (1989) noted that in Gambelia, the groove is usually closed over two-thirds of its length anterior to the splenial. Unfortunately, this condition is much more variable in Garnbelia than in Crotaphytus. and although the groove in most specimens is closed over greater than one-half of its length anterior to the splenial, 12 of 30 G. silus, two of nine G. copei, and 12 of 45 G. wislizenii had a condition similar to that observed in Crotaphytus, with the groove closed over less than half of its length anterior to the splenial. Because of this variation, this character was not considered in the phylogenetic analysis. Norell (1 989) also considered an elongate dentary (with a posterior process projecting posterior to the superior apex of the coronoid, Etheridge and de Queiroz, 1988) to be a synapomorphy for Crota- phytidae. Although this character state was found to be derived in their phylogenetic analysis of pleu- rodont iguanians (possibly a paraphyletic assem- blage with respect to acrodont iguanians [Chamae- leonidae]), this state is widespread within Iguania and may be a synapomorphy for a group more in- clusive than Crotaphytidae. The dentary bears between three and eight mental foramina anteriorly. In Crotaphytus, the mental fo- ramina are usually restricted to the distal end of the dentary, while in Gambelia they may extend pos- teriorly to the midpoint of the bone. Continuous variation in this feature prevented its inclusion in the phylogenetic analysis. A trgular (Characters 23, 24; Fig. 1 3). -In Crota- phytus, the exposed portion of the angular extends further anteriorly than in Garnbelia wislizenii and G. copei. Defining states for this character is com- plicated by the variation that exists in those struc- tures that may serve as reference points. For this reason, two points of reference are included in the description of this character. In adult Crotaphytus. with very few exceptions, the angular extends an- teriorly at least to the fourth tooth (counting from

BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32 the rear of the tooth row) and usually well beyond this point. Juveniles are not always comparable be- cause their teeth are relatively larger than those of adults and are often widely spaced. The angular also extends well beyond the anterior extent of the cor- onoid in both adults and juveniles. In G. tvislizenii and G. copei, the angular was never observed to reach the fourth tooth (from the rear of the tooth row) and rarely reached beyond the first. In most specimens, the angular does not extend as far an- teriorly as does the coronoid. In C. bicincfores, C. grismeri, and G. silus, the anterior extent of the angular shows continuous variation with most spec- imens having an intermediate condition but others with character states similar to those observed in G. wislizenii and G. copei or the remaining species of Crotaphytus. Because of this continuous variation in these three taxa, 1 have coded each as unknown for this character. With respect to the outgroup taxa, the angular projects well anteriorly in chamaeleon- ids, hoplocercids, the corytophanids Basiliscus bas- iliscus, B. vittatus, B. plumifom, some Coryto- phanes cristatus. C. percarinatus, some Laernanctus longipes, L. serratus, and many polychrotids, while it is short in tropidurids (except Uranoscodon su- perciliosus), phrynosomatids, oplurids (except 0. fierinensis), and iguanids. The angular bears the posterior mylohyoid fora- men. This foramen usually is positioned well pos- terior to the superior apex of the coronoid in Gam- belia (eight of eight G. copei, 26 of 29 G. silus. 50 of 51 G. wislizenii), while it is equidistant with, or anterior to, the superior apex in most Crotaphytus (posterior to the superior apex in two of 49 C. col- laris, three of 15 C. dickersonae. two of 17 C. ne- brius, two of 23 C. reficulafus, one of 27 C. vestig- ium). Although most of the outgroup taxa exhibit the condition observed in Crotaphyfus, the presence of the posterior mylohyoid foramen posterior to the apex of the coronoid in phrynosomatids, some tro- pidurids, and some polychrotids (Frost and Ether- idge, 1989) as well as some oplurids prohibits po- larization of this character. Coronoid (Character 25; Fig. 13, 14).-The angle of the posterolingual process of the coronoid is near- ly vertical in Crotaphytus, while it extends poster- oventrally at an angle of approximately 45 degrees in G. tvislizenii, G. copei, and G. corona? (Norell, 1989). Gambelia silus may be intermediate in this feature or may approach the conditions observed in Croraphytus or G. tvislizenii-G. copei. Therefore, G. silus was coded as unknown ("?") for this character. Most outgroup taxa have a condition similar to Cro- faphytus (state 0) or occasionally the intermediate condition usually present in G. silus. The outgroup taxa with the G. wislizenii-G. copei condition in- clude only Pefrosaurus mearnsi, Phrynosoma doug- lassi. P. coronatutn, Uromasfyx. Brookesia sfutnpfi, and Chamaeleo kersteni (chamaeleonines as a whole are variable with respect to this feature). Therefore, the angled posterolingual process of the coronoid (state 1) is considered to be derived and the vertical condition ancestral. Norell (1989) considered this feature to be an unambiguous synapomorphy of Garnbelia, presumably because he did not examine specimens of G. silus. Surangular (Characters 26-28; Fig. 13-1 5). - Im- mediately anterior to the articular facet lies a me- dially oriented knob-like process here referred to as the medial process. A thin shelf of bone may extend anteriorly between the distal extremity of the medial process and the body of the surangular (Fig. 15). This shelf is usually much more strongly developed in Gambelia and, to a lesser degree, Crotaphyfus insularis than in the remaining Crotaphytus species. Crotaphytus vestigium is variable with respect to this character with seven of 27 having a shelf present. A lesser amount of variation was observed with a smaller shelf present in C. bicincfores (one of 25), C. collaris (five of 50), C. dickersonae (two of 16), C. nebrius (one of 17), and C. reticulatus (one of 14). In Gambelia, the shelf may entirely fill this space such that its edge may be either straight or, more frequently, convex in shape. The strongly de- veloped condition present in Gambelia suggests that it may be a further modification or intensification of the condition observed occasionally in Crofa- phytus. Thin shelves of bone between the medial process and the ramus of the mandible are present in a small number of iguanian taxa, including Lei- olepis belliana, Oplurus cuvieri, some Brachylophus fasciatus, some Ufa stansburiana. Urosaurus auri- culatus, Microlophus grayi, and most Phymarurus taxa (absent only in P. palluma and some P. punae). The shelves only approached the condition of Gam- belia in the four Phymaturus pafagonicus subspe- cies. This character was coded as a binary character with the absence of a shelf coded as state 0 and its presence as state 1 (taxa with intermediate frequen- cies coded appropriately). The presence of thin shelves of bone between the medial process of the surangular and the ramus of the mandible is inter- preted as the derived state. An additional process of the surangular may be

1996 McGUIRE-SYSTEMATICS OF CROTAPHYTID LIZARDS 2 1<br />

- -<br />

Fig. 13.-Lingual view of the right mandible of (A) Croraphyrus Fig. 14.-hbial view of the right mandible of (A) Crotaphytw<br />

reticularus (REE 2912, adult male, SVL = 122 mm) and (B) rfliculallu (REE 291 2, adult male, SVL = 122 mm) and (B)<br />

Gatnbclia cowi (REE 2800, adult female. SVL = 123 mm). Ang Gamhflia coppi (REE 2800, adult female, SVL = 123 mm). Cor<br />

= angular, An - anicular. Cor = coronoid, Den = dentav, pmf = COronoid~ Den dcntav, PrC = preanicular, Sur = surangular.<br />

=<br />

= posterior mylohyoid foramen, Pre = preanicular, Spl = splenmm.<br />

ial. Sur = surangular. Scalc = 5 mm.<br />

completely encloses Meckel's cartilage. In crotaphytids,<br />

the tubular nature of the dentary is incomplete.<br />

The anterior end of the dentary is open, while<br />

posteriorly the groove is closed, but not fused. In<br />

Crotaphy~tu, with the exception of C. grismeri, the<br />

groove is usually closed over less than one-half of<br />

its length anterior to the splenial and there is relatively<br />

consistent interspecific variation in this characteristic.<br />

In C. collaris. C. nebrius. and C. reticulatus.<br />

Meckel's groove is often open over its entire<br />

length anterior to the splenial, and most of the remaining<br />

specimens have the groove closed over less<br />

than one-third of its length. In C. antiquus and C.<br />

dickrsonae, the groove is not open over its entire<br />

length, but as in the above-mentioned taxa, it was<br />

nearly always closed over less than one-third of its<br />

length. In C. bicinctores, C. insularis, and C. vestigiurn.<br />

Meckel's groove is usually closed over between<br />

one-third and one-half of its length anterior to the<br />

splenial and was only once observed to be open over<br />

its entire length (C, vestigittrn. REE 28 11). Crotaphytusgrismeri<br />

is unique among Crotaphytus in that<br />

Meckel's groove is closed over between approximately<br />

50 percent and 70 percent of its length in all<br />

specimens examined (five of five). Norell (1989)<br />

noted that in Gambelia, the groove is usually closed<br />

over two-thirds of its length anterior to the splenial.<br />

Unfortunately, this condition is much more variable<br />

in Garnbelia than in Crotaphytus. and although the<br />

groove in most specimens is closed over greater than<br />

one-half of its length anterior to the splenial, 12 of<br />

30 G. silus, two of nine G. copei, and 12 of 45 G.<br />

wislizenii had a condition similar to that observed<br />

in Crotaphytus, with the groove closed over less than<br />

half of its length anterior to the splenial. Because of<br />

this variation, this character was not considered in<br />

the phylogenetic analysis.<br />

Norell (1 989) also considered an elongate dentary<br />

(with a posterior process projecting posterior to the<br />

superior apex of the coronoid, Etheridge and de<br />

Queiroz, 1988) to be a synapomorphy for Crota-<br />

phytidae. Although this character state was found<br />

to be derived in their phylogenetic analysis of pleu-<br />

rodont iguanians (possibly a paraphyletic assem-<br />

blage with respect to acrodont iguanians [Chamae-<br />

leonidae]), this state is widespread within Iguania<br />

and may be a synapomorphy for a group more in-<br />

clusive than Crotaphytidae.<br />

The dentary bears between three and eight mental<br />

foramina anteriorly. In Crotaphytus, the mental fo-<br />

ramina are usually restricted to the distal end of the<br />

dentary, while in Gambelia they may extend pos-<br />

teriorly to the midpoint of the bone. Continuous<br />

variation in this feature prevented its inclusion in<br />

the phylogenetic analysis.<br />

A trgular (Characters 23, 24; Fig. 1 3). -In Crota-<br />

phytus, the exposed portion of the angular extends<br />

further anteriorly than in Garnbelia wislizenii and<br />

G. copei. Defining states for this character is com-<br />

plicated by the variation that exists in those struc-<br />

tures that may serve as reference points. For this<br />

reason, two points of reference are included in the<br />

description of this character. In adult Crotaphytus.<br />

with very few exceptions, the angular extends an-<br />

teriorly at least to the fourth tooth (counting from

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