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BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32
of Cuba (Hardy, 1958; Ruibal and Williams, 196 I)
and in several species of Leiocephalus (C. A. Haas,
S. B. Hedges, personal communication, 1994; K. de
Queiroz, personal communication, 1995-although
they are described as coiling their tails vertically
over their backs by Schwanz and Henderson, 199 I).
However, these groups are nested within Polychrotidae
and Tropiduridae, respectively, indicating that
their behaviors are convergent with that observed
in Crotaphyrus. The presence of lateral tail coiling
is considered to be the derived state.
Constrtnption oj Vertebrares. -All crotaphytids
except Crotaphytus antiquus. C. gristneri, and C.
nebrius have either been documented in the literature
to include vertebrates in their diets (C. bicinctores:
Banta, 1960; Snyder, 1972; Nussbaum et al.,
1983; C. collaris: Filch, 1956; McAllister and Trauth,
1982; C. reticulatus: Klein, 195 1; Montanucci, 197 1 ;
Gambelia copei: Banta and Tanner, 1968; Montanucci,
1965; G. ~vislizenii: Stejneger, 1 893; McCoy,
1967; Montanucci, 1967; Snyder, 1972; Tanner and
Krogh, 1974a; Parker and Pianka, 1976; Tollestrup,
1979, 1983; Pietruszka et al., 198 1; Crowley and
Pietruszka, 1983) or have been observed to do so
by the author. The primary vertebrate prey is other
lizards, although rodents and snakes also have been
recorded. There appears to be variation in the relative
proportion of vertebrates included in the diets
of the various species, with Gatnbelia btblizenii
(Parker and Pianka, 1976; Tollestrup, 1979, 1982,
1983) and G. copei consuming a greater proportion
of vertebrate prey than other species.
Many other iguanian species are known to eat
vertebrates, including the phrynosomatid genera Petrosaurus,
Uma, Holbrookia, and Sceloporus, which
are all known to include other lizards in their diets
(Stebbins, 1 985); the corytophanid Basiliscus (Van
The following character list includes the morpho-
logical characters (informative or uninformative)
discussed in the text, as well as the nine informative
allozyme characters (characters 89-98) that could
be coded using the Manhattan distance frequency
approach discussed in Wiens (1 995). One multistate
morphological character (3 1) was also coded using
the Manhattan distance frequency approach. Its step
matrix is presented in Appendix 4 along with the
step matrices for the allozyme characters. Character
descriptions followed by (P) are polarized, those fol-
lowed by (U) are unpolarized, and those followed
CHARACTER LIST
Devender, 1 982); the pol ychrotid A nolis equestris
(Ruibal, 1964); and the chamaeleonids Chlatnydo-
saurus kingii and Physignathus Icsueurii (Comer,
1992). I have not attempted to review the feeding
habits of all of the potential outgroup taxa, but it is
likely that many other species have similar feeding
habits. Thus, the presence or absence of carnivory
may not be a polarizable character, limiting its use-
fulness in this analysis. Furthermore, since most
lizards will eat anything palatable that they are able
to overcome, the inclusion of vertebrates in the diet
may be, at least in pan, a function of maximum
adult size. For these reasons, this characteristic was
not included in this analysis. However, the carniv-
orous predatory habits of Crotaphyttcs and Gam-
belia are consistent with a hypothesis ofcrotaphytid
monophyly.
Vocalization. -The ability to vocalize is rare in
squamates, with gekkotans being the only family in
which it is known to occur commonly. Within ig-
uanian lizards, vocalization is apparently limited to
crotaphytids and certain polychrotids. A squealing
sound is known to be emitted by Gatnbelia wislizenii
(Jorgenson et al., 1963; Wever et al., 1966; Smith,
1974) and Crotaphytus bicinctores (Smith, 1974)
during periods of stress. Similar vocalizations were
discussed by Ruibal(1964) in three species of Cuban
anoles, Anolis isolepis, A. lucius, and A. vermiculatus
and by Lynn and Grant (1940) in A. grahami and
A. opalinus (also noted in A. grahalni by Etheridge,
1955). Because vocalization data are lacking for the
majority of crotaphytid species, I have not included
this character in the phylogenetic analysis. How-
ever, as with carnivory, the presence of vocalization
in some species of Crotaphytus and Gambelia is
consistent with the hypothesis of crotaphytid mono-
P~Y~Y.
by (UO) are unordered. Characters 28, 68, 75, 84,
and 85 were not analyzed using frequency coding
(see Materials and Methods).
SKULL AND MANDIBLE
1. Nasal process of the premaxilla (P): (0) broad,
(I) narrow and elongate.
2. Ventral suture between vomers and premaxilla
(P): (0) does not form a strong vertical ridge,
(1) forms a strong vertical ridge.
3. Nasals (P): (0) do not overlap nasal process of
the premaxilla anterior to posterior extent of