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McGUIRE- SYSTEMATICS OF CROTAPHYTID LIZARDS 7 7 pale to fluorescent. The white component of the dorsal pattern is retained well in preservative and is easily characterized as nearly all populations have white spots on the body with spots or reticulations present on the tail and hindlimbs. Some populations from Coahuila, Durango, Nuevo Leon, San Luis Potosi, and Zacatecas may have a dorsal pattern consisting at least in pan of black spots that may or may not be surrounded by white, a pattern that is reminiscent of that of C. antiquus and C. reticulatus and potentially the result of introgression from the latter species (Montanucci, 1974). The forelimbs are generally patternless or only obscurely patterned, but may occasionally bear pale reticulations or spots. Transverse body bars are absent. Reticulations gen- erally are confined to the superficial mandibular and temporal regions, as well as the hindlimbs and tail. A broad white or off-white caudal vertebral stripe is lacking. The dorsal surface of the head is not pale- colored, and generally is covered with spots that range in color from rust to chocolate brown. Olive green or orange ventrolateral coloration is lacking. Most of the variation in gular pattern coloration observed within Crotaphytus is restricted to C. col- laris. The gular coloration observed in living adult males examined over the course of this study range between olive green, dark blue, turquoise blue, slate gray, yellow, or orange. However, a black central component is not found in this species. As stated above, the peripheral gular pattern is always com- posed of a white reticulated pattern. Anterior and posterior collar markings are always present and the posterior markings occasionally may contact mid- dorsally. The anterior collars are not complete ven- trally as black pigments are absent from the gular fold. A pair of black spots may be present middor- sally between the anterior collar markings. A pair of enlarged melanic axillary patches are variably present immediately posterior to the forelimb in- sertion, although they are restricted to populations from the western portion of the species' range (Ar- izona). Small melanic inguinal patches are also vari- ably present in adult males from this portion of the range. The femoral pores are generally off-white to gray in color. Paired, melanic keels may or may not be present on the ventral surface of the caudal ex- tremity. Female Crotaphytus collaris are much less con- spicuously marked than males, particularly in those populations characterized by green dorsal colora- tion. While females may retain a green component in their pattern, it is always of a much duller hue. As in other Crotaphyrw. the gular pattern of females is less developed. Inguinal patches, which are vari- ably present in adult males, are lacking in females. Females develop vivid orange or reddish lateral bars during the gravid period. The tail is not vividly colored in either adult or subadult females. Size. -This species exhibits strong sexual dimor- phism with males reaching larger adult size (maxi- mum observed SVL = 13 1 mm) than females (max- imum observed SVL = 106 mm). Distribution (Fig. 45). - Crotaphytus collaris has an extensive distribution in the western and south- central United States and northern Mexico extend- ing from northwestern Arizona, eastern Utah, and western Colorado eastward across the southern Great Plains into Missouri, northern Arkansas, and pos- sibly extreme northwestern Louisiana; and south- ward into extreme nonhern Sonora and nonhcen- tral mainland Mexico. Numerous isolated popula- tions occur on the eastern periphery of its range in Missouri and Arkansas. In Texas, the eastern dis- tributional extent of C. collaris is limited by the Balcones Escarpment as suitable rocky habitat does not extend east of this point. For this reason, a num- ber of localities that lie east of the escarpment are considered questionable (FMNH 1 1 7 1 16-1 8 -"Ce- dar Creek, Bastrop Co."; USNM 12762-"Tehu- acana, Limestone Co.," 145 18 -"Gainesville, Cooke Co."; UTA 892-"10 mi. S Dallas, Dallas Co."; see Axtell [1989a] for a more complete assessment of potentially erroneous localities for Texas speci- mens). In Mexico, C. collaris extends as far east as the eastern slopes of the Sierra Madre Oriental, while C. reliculatus occupies the flatland Tamaulipan thornscrub habitats to the immediate east. These two species approach one another closely in the vi- cinity of Allende, Coahuila, Mexico. In western and northern Arizona, the distributions of C. collaris and C. bicinctores abut one another and at least two hybrid zones occur (see description of the distri- bution of C. bicincrores). The questionable ("?") Colorado locality on the dot distribution map (Fig. 45) refers to a specimen (USNM 58603) from Ar- chuleta County, Colorado, for which no specific lo- cality data were given. The questionable ("?') 10- cality from near the border between Tamaulipas and San Luis Potosi, Mexico, represents a locality given for C. rcticulatus (AMNH 104448-"rte. 101, 12 mi. SW jct. with side rd. to Tula, 13 mi. NE San Luis Potosi state line"). This locality is dubious for C. reticularus, but would not be unexpected for C. collaris. An extremely detailed dot distribution map for C. collaris in Texas was provided by Axtell (I 989a).
7 8 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32 30 Pacific Ocean 110 100 Gulf of Mexico Crotaphytus collaris Crotaphytus reticulatus Fig. 45.-Geographic distribution of Crotaphyrics collaris and C, reticulatus. Thc "?" in southern Colorado denotes a specimen without precise locality data from Archuleta County. The "?"s along the eastern periphery of C. collaris' range in Texas represent dubious localities that Lie east of the Balcones Escarpment. The "?" locality from near the border between Tamaulipas and San Luis Potosi, Mexico, represents a locality given for C. reticulatus that is dubious for this species, but would not be unexpected for C. collaris. Dot distribution maps for the states of Colorado greet, Sabine Parish (cited as a personal commu- (Hammerson, 1986), Kansas (Collins, 1982), Mis- nication in Dundee and Rossman, 1989). souri (Johnson, 1987), and Oklahoma (Webb, 1970) Fossil Record. -Numerous Pleistocene fossils have also been published. from several western states have been referred to Dundee and Rossman (1989) questioned whether this taxon, including a number of fossils more rea- C. collaris occurs naturally in the state of Louisiana. sonably referred to other species (see C. bicinctores Two specimens are known, one of which may have and C. nebrius accounts). All of the fossils, with the been accidentally introduced (Frierson, 1927), while above exceptions, fall within the current distribu- the other was collected by D. Leslie at Boone's Land- tional limits of C. collaris (Estes, 1983). ing on the Toledo Bend Reservoir southwest of Ne- Natural History. -More has been written about
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McGUIRE- SYSTEMATICS OF CROTAPHYTID LIZARDS 7 7<br />
pale to fluorescent. The white component of the<br />
dorsal pattern is retained well in preservative and<br />
is easily characterized as nearly all populations have<br />
white spots on the body with spots or reticulations<br />
present on the tail and hindlimbs. Some populations<br />
from Coahuila, Durango, Nuevo Leon, San Luis<br />
Potosi, and Zacatecas may have a dorsal pattern<br />
consisting at least in pan of black spots that may or<br />
may not be surrounded by white, a pattern that is<br />
reminiscent of that of C. antiquus and C. reticulatus<br />
and potentially the result of introgression from the<br />
latter species (Montanucci, 1974). The forelimbs are<br />
generally patternless or only obscurely patterned,<br />
but may occasionally bear pale reticulations or spots.<br />
Transverse body bars are absent. Reticulations gen-<br />
erally are confined to the superficial mandibular and<br />
temporal regions, as well as the hindlimbs and tail.<br />
A broad white or off-white caudal vertebral stripe<br />
is lacking. The dorsal surface of the head is not pale-<br />
colored, and generally is covered with spots that<br />
range in color from rust to chocolate brown. Olive<br />
green or orange ventrolateral coloration is lacking.<br />
Most of the variation in gular pattern coloration<br />
observed within Crotaphytus is restricted to C. col-<br />
laris. The gular coloration observed in living adult<br />
males examined over the course of this study range<br />
between olive green, dark blue, turquoise blue, slate<br />
gray, yellow, or orange. However, a black central<br />
component is not found in this species. As stated<br />
above, the peripheral gular pattern is always com-<br />
posed of a white reticulated pattern. Anterior and<br />
posterior collar markings are always present and the<br />
posterior markings occasionally may contact mid-<br />
dorsally. The anterior collars are not complete ven-<br />
trally as black pigments are absent from the gular<br />
fold. A pair of black spots may be present middor-<br />
sally between the anterior collar markings. A pair<br />
of enlarged melanic axillary patches are variably<br />
present immediately posterior to the forelimb in-<br />
sertion, although they are restricted to populations<br />
from the western portion of the species' range (Ar-<br />
izona). Small melanic inguinal patches are also vari-<br />
ably present in adult males from this portion of the<br />
range. The femoral pores are generally off-white to<br />
gray in color. Paired, melanic keels may or may not<br />
be present on the ventral surface of the caudal ex-<br />
tremity.<br />
Female Crotaphytus collaris are much less con-<br />
spicuously marked than males, particularly in those<br />
populations characterized by green dorsal colora-<br />
tion. While females may retain a green component<br />
in their pattern, it is always of a much duller hue.<br />
As in other Crotaphyrw. the gular pattern of females<br />
is less developed. Inguinal patches, which are vari-<br />
ably present in adult males, are lacking in females.<br />
Females develop vivid orange or reddish lateral bars<br />
during the gravid period. The tail is not vividly<br />
colored in either adult or subadult females.<br />
Size. -This species exhibits strong sexual dimor-<br />
phism with males reaching larger adult size (maxi-<br />
mum observed SVL = 13 1 mm) than females (max-<br />
imum observed SVL = 106 mm).<br />
Distribution (Fig. 45). - Crotaphytus collaris has<br />
an extensive distribution in the western and south-<br />
central United States and northern Mexico extend-<br />
ing from northwestern Arizona, eastern Utah, and<br />
western Colorado eastward across the southern Great<br />
Plains into Missouri, northern Arkansas, and pos-<br />
sibly extreme northwestern Louisiana; and south-<br />
ward into extreme nonhern Sonora and nonhcen-<br />
tral mainland Mexico. Numerous isolated popula-<br />
tions occur on the eastern periphery of its range in<br />
Missouri and Arkansas. In Texas, the eastern dis-<br />
tributional extent of C. collaris is limited by the<br />
Balcones Escarpment as suitable rocky habitat does<br />
not extend east of this point. For this reason, a num-<br />
ber of localities that lie east of the escarpment are<br />
considered questionable (FMNH 1 1 7 1 16-1 8 -"Ce-<br />
dar Creek, Bastrop Co."; USNM 12762-"Tehu-<br />
acana, Limestone Co.," 145 18 -"Gainesville, Cooke<br />
Co."; UTA 892-"10 mi. S Dallas, Dallas Co."; see<br />
Axtell [1989a] for a more complete assessment of<br />
potentially erroneous localities for Texas speci-<br />
mens). In Mexico, C. collaris extends as far east as<br />
the eastern slopes of the Sierra Madre Oriental, while<br />
C. reliculatus occupies the flatland Tamaulipan<br />
thornscrub habitats to the immediate east. These<br />
two species approach one another closely in the vi-<br />
cinity of Allende, Coahuila, Mexico. In western and<br />
northern Arizona, the distributions of C. collaris<br />
and C. bicinctores abut one another and at least two<br />
hybrid zones occur (see description of the distri-<br />
bution of C. bicincrores). The questionable ("?")<br />
Colorado locality on the dot distribution map (Fig.<br />
45) refers to a specimen (USNM 58603) from Ar-<br />
chuleta County, Colorado, for which no specific lo-<br />
cality data were given. The questionable ("?') 10-<br />
cality from near the border between Tamaulipas and<br />
San Luis Potosi, Mexico, represents a locality given<br />
for C. rcticulatus (AMNH 104448-"rte. 101, 12<br />
mi. SW jct. with side rd. to Tula, 13 mi. NE San<br />
Luis Potosi state line"). This locality is dubious for<br />
C. reticularus, but would not be unexpected for C.<br />
collaris.<br />
An extremely detailed dot distribution map for<br />
C. collaris in Texas was provided by Axtell (I 989a).