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BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32
casionally present in C. nebrim (I 2 of 5 I). The spots,
which occur between the dorsal extensions of the
anterior collar markings, appear to be homologous
with the first transversely arranged row of blackfilled
hexagonal reticulations seen in C. reticulatus.
In C. antiquus, the nuchal spots are always present,
but often incompletely separated from the remainder
of the anterior collar markings. Black nuchal
spots are not present in the outgroup taxa and their
presence is coded as the derived state.
Ingitinal Patches (Characters 77,78; Fig. 32C, 34,
35).-In several species of Crotaphytus, adult males
develop dark brown or black ventral patches in the
inguinal region. These patches vary considerably in
size with C. bicinctores, C. dickersonae, C. grismeri,
C. insularis, and C. vesrigiittn having large patches
and C. antiquus, C. nebrius, and C. collaris having
smaller ones. All adult male C. bicinctores, C. antiquus,
C. dickersonae, C. grismeri, C. insularis, C.
nebrius, and C. vesrigiurn develop these patches while
only some C. collaris have them. Interestingly, only
C. collaris from the western periphery of its range
(in the area usually referred to the subspecies C. c.
baileyi) are known to have inguinal patches. Thus,
there are at least two characters associated with inguinal
patches: size of the patches and the frequency
with which they occur. Homology of the patches
seems likely. Both large and small patches begin
development as small ventral spots near the hind
limb insertion and the large patches differ only in
that they continue to become larger (and probably
grow faster). Inguinal patches of the type present in
some Crolaphj)rr& are extremely rare in the outgroup
taxa. Similar markings are present in Uma exsuland
LI. paraphygas (de Queiroz, 1989; although they occur
more laterally than in Crotaphyrus), Ufa nolascensis,
Uromasryx hardwickii (concentrated on the
thigh), and Enyalius iheringii (again, more laterally
oriented). This character has been coded two ways:
first, as a binary character with the absence of inguinal
patches (of any size) as state 0 and the presence
ofpatches as state 1 ; and secondly, as a separate
binary character with the presence of small patches
as state 0 and the presence of large patches as state
1. Taxa without inguinal patches were scored as
unknown ("?') for this second character. Because
the first character (77) considers the frequency in
which patches are present, the second character (78)
does not take frequency into consideration. For
character 78, the presence of small patches in any
frequency is assigned state 0 and the presence of
large patches in any frequency is coded as state 1.
Fetnoral Pore Secreriotu (Character 79; Fig. 22,
23, 33-35).-The femoral pore secretions of male
Crotaphyt us retictrlattrs and C. anriquus are jet black.
Unlike other Crotaphytus species, such as C. ne-
brim, which often have grayish secretions, the sub-
cutaneous glands themselves are also jet black. This
condition was not observed in other species of ig-
uanian lizards and is treated as the derived state.
Gular Pattern (Characters 80-82; Fig. 33-35).-
There is much variation in the gular pattern of male
Crotaphytus, especially in the wide-ranging species
C. collaris. However, the general arrangement of the
gular colors is similar in all of the species. For ex-
ample, each has a relatively uniformly colored cen-
tral gular region that is surrounded by a peripheral
reticulated or spotted pattern superficial to the man-
dibles. It is in the context of this general pattern that
the following discussion of variation is based. Be-
cause the pattern and extent of the gular coloration
is sexually dichromatic, the following discussion
pertains only to adult male Crotaphyrus.
Adult male Crotaphytus bicinctores, C. antiquus,
C. dickersonae, C. grismeri, C. insularis. C. rericu-
lattrs, and C. vestigiunt (Fig. 35) have a patch of
black pigment in the posteromedial portion of the
gular region. This pigmentation corresponds with
that portion of the gular pouch that is depressed by
the second ceratobranchials of the hyoid apparatus,
and thus presumably increases the visibility of the
depressed gular pouch during aggressive display. The
black patch is continuous with the black pigrnen-
tation of the gular fold and the ontogenetic devel-
opment of the gular patch suggests that it may be
an extension of the gular fold coloration. However,
the presence of black pigmentation in the gular fold
and in the posteromedial portion of the central gular
region are treated as separate characters because the
presence of black pigmentation in the gular fold is
not always associated with a black central gular patch
(e.g., C. nebrius). Because the outgroups do not have
a gular pattern that is similar to Crotaphytus, out-
group analysis cannot be utilized to assess polarity.
Therefore, this character was left unpolarized.
There is considerable variation in the peripheral
gular pattern of Croraphytus as well. In all Crora-
phytus except C. nebrius, the peripheral gular pattern
is composed of white reticulations on a dark field.
In most C. collaris, the dark pigmentation within
each subquadrate reticulation has a light center,
which results in a pattern reminiscent of the dorsal
pattern of a jaguar. A similar pattern is sometimes
evident in other species, such as C. bicinctores. In