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4 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32
throughout its range. Bun (1 9286) synonymized
Crotaphytus collaris baileyi with C. c. collaris on the
basis of extensive variation in the interorbital scale
characteristics used to separate the two forms, a
taxonomy that was not followed by subsequent
workers. Allen (1 933) reduced Crotaphytus dicker-
sonae to a subspecies of C. collaris, citing intergra-
dation in the hindlimb and tail length characters
that Schmidt (1922) used to distinguish C. dicker-
sonae from C. c. baileyi. Allen ( 1 933) did not follow
Bun's (1928b) synonymy of C. c. baileyi with C. c.
collaris.
Mittleman (1942) discussed the higher level phy-
logenetic relationships within North American ig-
uanian lizards. His diagrammatic representation of
relationships placed Crotaphytus as the sister taxon
of Petrosaurus and Streptosaurus. This group was
in turn depicted as the sister group of the phryno-
somatid sand lizards Ulna, Callisaurus, and Hol-
brookia. Despite the relationships implied by his
tree, he appears to have considered Crotaphytus to
be a relatively primitive iguanid (senso lato), be-
cause he suggested that the sand lizards were derived
from Crotaphytus-like stock, as was Sauromalus.
Smith (1946) separated Crotaphyrus wislizenii
from C. collaris and C. reticitlalus by placing it in
the genus Gambelia, thus elevating Baird's (1 858)
subgenus to generic rank. This controversial deci-
sion initiated much debate among various workers
on the group. Furthermore, Smith (1946) reduced
G. silus to a subspecies of G. wislizenii. With respect
to higher taxonomic relationships within the Igu-
ania, Smith followed Mittleman (1942) in placing
Crotaphyrus and Gambelia as the sister group of
Srreprosaurus plus Petrosaurus, and this group as
the sister taxon of the phrynosomatid sand lizards.
Smith and Taylor (1950) elevated dickersonae from
a subspecies of Crotaphytus collaris to the rank of
full species.
Fitch and Tanner (1 95 l), reinterpreting the data
of Bun (1928b), recognized Crotaphytus collaris
baileyi as a subspecies distinct from C. c. collaris.
This taxonomy had generally been followed in the
literature despite the earlier synonymy of the two
by Burt (1928b). In addition, they described a new
subspecies of Crotaphytus, C. c. atrriceps, from the
upper Colorado River basin.
Returning to the higher-level relationships within
the Iguania, Savage (1958) presented a phylogeny
that differed radically from that of Mittleman (1 942)
and Smith (1946). In his classification, Savage pro-
posed a new subgrouping, the iguanines, that in-
cluded Croraphytus plus those genera later placed
in the Iguanidae by Frost and Etheridge (1 989).
Cochran (1 96 1) recognized Crotaphytus silus as a
full species. Robison and Tanner (1962) attempted
to resolve the Crotaphytus-Gambelia debate by examining
osteological and myological evidence. As
a result, they chose not to recognize Gambelia as a
genus distinct from Crotaphytus.
Tanner and Banta (1963), in the first of a threepan
series examining the systematics of leopard lizards,
described a new subspecies, Croraplvtus wislizeni
punctatus, from the upper Colorado River basin
of Utah and Colorado. Like Cochran (1961),
those authors did not recognize the genus Gambelia.
Etheridge (1 964) removed Crotaphytus from Savage's
(1 958) iguanines because he was unable to find
any character or combination of characters that
would serve to diagnose the iguanines if Crotaphytus
was included. Furthermore, he hypothesized that
Crotaphytus may be the sister taxon to the sceloporines
(= Phrynosomatidae) plus tropidurines (=
Tropiduridae).
Leviton and Banta (1964) resurrected the name
copei for the Baja California populations of Crotaphyrus
\vislizenii, recognizing C. w. copei.
Weiner and Smith (1965) attempted to resolve
the Garnbelia-Crotaphytus controversy by examining
the osteology of the group. They placed all
members of Crotaphytus (including those that had
been placed in the genus Gambelia) into a grouping
they referred to as the "crotaphytiform" lizards. They
recognized only four species of crotaphytiform lizards:
C. collaris, C. reticulatus. C. insularis, and C.
~vislizeni (again relegating silus to a subspecies of C.
wislizeni). Thus, without presenting evidence, Weiner
and Smith (1 965) reduced C. dickersonae to the
rank of subspecies within C. collaris. Those members
of the genus with a superficial resemblance to
C. collaris (C. collaris, C. reticulatus, and C. insularis)
were further separated into the "collariform"
group. Finally, with respect to the Gambelia-Croraphytus
debate, they concluded that the subgeneric
rankings, Crotaphytus (Garnbelia) ~r~islimi first proposed
by Baird (1858) and Crotaphyrus (Crotaphytus)
were the lowest levels of taxonomic segregation
that could be justified by the data.
SoulC and Sloan (1 966) followed Weinerand Smith
( 1965) in recognizing dickersonae as a subspecies of
C. collaris and reduced insularis to a subspecies of
C. collaris as well.
Banta and Tanner (1968), in their second study
of leopard lizard systematics, provided a redescrip-