Herpetologists' League

The Relationship of Morphology to Diet in the Horned Lizard Genus Phrynosoma

Author(s): Richard R. Montanucci
Source: Herpetologica, Vol. 45, No. 2 (Jun., 1989), pp. 208-216
Published by: Herpetologists' League
Stable URL: http://www.jstor.org/stable/3892163 .
Accessed: 31/07/2013 19:43

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .
http://www.jstor.org/page/info/about/policies/terms.jsp

.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of
content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms
of scholarship. For more information about JSTOR, please contact support@jstor.org.

Herpetologists' League is collaborating with JSTOR to digitize, preserve and extend access to Herpetologica.

http://www.jstor.org

This content downloaded from 128.112.200.107 on Wed, 31 Jul 2013 19:43:16 PM

All use subject to JSTOR Terms and Conditions
Herpetologica, 45(2), 1989, 208-216
C) 1989 by The Herpetologists'League, Inc.

THE RELATIONSHIP OF MORPHOLOGYTO DIET

IN THE HORNED LIZARD GENUS PHRYNOSOMA
RICHARD R. MONTANUCCI
Department of Biological Sciences, Clemson University,
Clemson, SC 29634-1903, USA

ABSTRACT: Horned lizards,genus Phrynosoma, show a range in dietary habits, such that some
species are more strongly myrmecophagous (ant-eating) than others. The relationship between
skeletalmorphologyand dietary specializationwas examinedby correlationand stepwise regression
analyses.Five of seven morphologicalvariablesfrom the mandibleand epipterygoidare significantly
correlatedwith the mean percentage of ants in the diet. Epipterygoidlength, height of the coronoid
process, vertical diameter of the mandible, and length of the mandible posteriorto the coronoid
process are all negatively correlated; length of the tooth row is positively correlated. Stepwise
regressionanalysisselected epipterygoid length as the single best predictorof diet in Phrynosoma.
The best two-variablemodel includes epipterygoid length and vertical diameter of the mandible,
and has an R2= 0.8377.
Key words: Lacertilia;Phrynosoma; Morphology;Foraging; Diet; Ants

HORNED lizards of the genus Phryno-
soma are primarily ant-eating reptiles,
whose dietary habitsare well known (Mon-
tanucci, 1981; Pianka and Parker, 1975,
and references cited therein; Powell and
Russell, 1984; Rissing, 1981; Turner and
Medica, 1982). The apparent relationship
between dentitional morphology and diet
in Phrynosoma was examined by Hotton
(1955) as part of a broader study of the
adaptive relationships of dentition to diet
in North American iguanids. He examined
three species (P. cornutum, P. douglassii,
and P. platyrhinos), concluding that they
were all similar in having blunt, non-cus-
pate, peg-like teeth, and that their diet
consisted almost entirely of ants. Presch
(1969) reiteratedHotton'sobservations,but
also noted that in three species (P. asio, P.
mcallii, and P. coronatum), the posterior
teeth show a hint of a tricuspid condition.
I could not confirm the difference between
anteriorand posteriorteeth in these species,
but the dentitional morphology of all
members of the genus should be reviewed.
Pianka and Parker (1975) determined
the percentage of ants (by number and
volume) in the diet of each species of Phry-
nosoma. In reviewing Pianka and Parker's
data, I concluded that the species of Phry-
nosoma are not all alike in their feeding
habits. Rather, there appears to be a range
of dietary habits within the genus, with
208
some species being more strongly myr-
mecophagous than others. Those species
with a comparatively lower percentage of
ants in the diet consume a greater variety
of terrestrialinsects and other arthropods.
When the mandibles of various species of
Phrynosoma were compared, I noted ap-
parent differences in robustnessand avail-
able area (on the coronoid and posterior
to it) for muscle attachment. Furthermore,
the epipterygoids, which are thought to
brace the palatal complex of the reptilian
skull (Romer, 1956), are reduced or absent
in Phrynosoma (Axtell, 1986; Presch,
1969). These observations led me to hy-
pothesize that in highly myrmecophagous
species of horned lizards, the mandible
should be relatively slender, area for mus-
cle attachment reduced, teeth reduced in
size, and epipterygoids greatly reduced or
absent. These anticipated relationshipsbe-
tween morphology and diet in Phrynoso-
ma are herein examined statistically.
METHODS AND MATERIALS
Mandiblesand epipterygoids from skel-
etons of Phrynosoma were examined un-
der a Wild M-5 dissecting microscope.The
left or right element from each specimen
was randomly selected for measurements,
which were made with an ocular microm-
eter. Adult specimens were selected in or-
der to minimize ontogenetic variationwith-

This content downloaded from 128.112.200.107 on Wed, 31 Jul 2013 19:43:16 PM

All use subject to JSTOR Terms and Conditions
June 1989] HERPETOLOGICA 209

in samples. The number of specimens K

examined for mandibles and epiptery-
goids, respectively, is indicated in paren-
theses as follows: Phrynosoma asio (8, 6);
P. braconnieri (4, 3); P. cornutum (19, 15);
P. coronatum (18, 19); P. ditmarsi (1, 1);
P. douglassii (15, 12); P. mcallii (8, 10); P.
modestum (5, 4); P. orbiculare (5, 4); P.
platyrhinos (19, 19); P. solare (8, 10); and
P. taurus (2, 2).
The following measurementsand counts
were obtained: (1) TP-the number of
tooth positions;(2) TH-average height of
the four tallest teeth in the tooth row; (3)
LC-length of the mandible from the pos-
terior edge of the coronoid process to the
posterioredge of the articularcondyle; (4)
TR-length of the tooth row from the an-
terior end of the dentary to the last tooth
position anterior to the coronoid process;
(5) CP-height of the coronoid process G
along its posterior border; (6) MD-ver-
tical diameter of the mandible taken at a
point midway between the last tooth po- AA BA
sition and the coronoid process; (7) EP-
length of the epipterygoid. FIG. 1. -The morphological variables used in this
Measurementsfrom each mandible were study: (1) number of tooth positions, and (2) average
standardizedby dividing each observation height of the four tallest teeth, between F and G; (3)
length of the mandible posterior to the coronoid pro-
by the total length of the mandible. Epi- cess, distance C-D; (4) length of the tooth row, dis-
pterygoid length was standardized by di- tance F-G; (5) height of the coronoid process, dis-
viding this measurement by the distance tance D-E; (6) vertical diameter of the mandible (two
between the snout and pineal foramen of arrows at H). Variables 1-6 were divided by mandible
the skull (Fig. 1). Standardized variables length, distance A-B. Epipterygoid length (I-J) was
divided by snout to pineal foramen distance (K-L).
are denoted by an asterisk (see Table 1 Skull modified from Jenkins and Tanner (1968).
below). Dietary data of species of Phry-
nosoma were obtained from Table 6 in
Pianka and Parker (1975). The percentage length. Therefore, the ratios seemed ap-
of ants (by volume) in the diet of each propriate for the statistical comparisons,
species was the datum used with the mor- and the use of analysis of covariance was
phological variablesfor statisticalanalyses. deemed unnecessary. Also, the standard-
The data represent average values of ant ized variables,as ratios,displayed a normal
consumption based on samples collected distribution, or did not depart sufficiently
from many localities and at different times from normality to warrant the use of non-
during the activity season. The percentage parametric regression techniques.
of ants in the diet of P. orbiculare is a Descriptive statistics (mean, standard
pooled average, incorporating the sample error of the mean, and observed limits)
from P. boucardi because the two taxa are were calculated for each morphological
consideredconspecific (Montanucci,1979). variable. In order to determine if associ-
The relationshipbetween total length of ations exist among the different measure-
the mandible and the other measurements ments, Pearson correlation coefficients
was determined to be essentially linear. were calculated for all variables,including
The same relationship appeared to exist the percentage of ants in the diet. The
between epipterygoid length and skull variablesthat were significantly correlated

This content downloaded from 128.112.200.107 on Wed, 31 Jul 2013 19:43:16 PM

All use subject to JSTOR Terms and Conditions
210 HERPETOLOGICA
with the proportionof ants in the diet were
then subjected to stepwise regressionanal-
ysis in order to confirm the anticipated
relationships between some of the inde-
pendent (morphological)variablesand the
dependent variable (percentage of ants).
The programused was the StatisticalAnal-
ysis System (SAS) STEPWISE procedure
(SAS Institute Inc., 1985). Results of the
stepwise regression analysis allowed selec-
tion of a model that best explains the re-
lationship between morphology and diet
in Phrynosoma.
RESULTS
Descriptive statistics for the seven mor-
phological variablesare presented in Table
1. Note that Phrynosoma ditmarsi is dis-
tinctive, compared with the other species,
in having a very long epipterygoid (EP*),
extensive vertical expansion of the man-
dible (MD*), tall coronoid process (CP*),
and the greatest distance from the coro-
noid processto the articularcondyle (LC*).
Phrynosoma solare is notable in having the
shortestteeth and the smallest mean value
for epipterygoid length. In some speci-
mens of P. solare, the elements are entirely
lacking. Phrynosoma solare, as well as P.
modestum and P. mcallii, have the short-
est coronoid process. The mandible is most
slender in P. modestum and P. solare, and
the shortest distance from the coronoid to
the articular process is seen in P. mcallii
and P. platyrhinos. Representative man-
dibles from the 12 species of Phrynosoma
are illustrated for comparison (Fig. 2).
The number of tooth positions in Table
1 represents standardized data. Thus, rel-
ative to jaw length, P. modestum has the
greatest number of teeth and P. asio has
the fewest teeth. The actual number of
tooth positions, based on a larger sample
(including left and right mandibles for
some specimens), is given below. The
species, sample size, mean, and observed
limits follow: P. asio (16), X = 20.8, 18-24;
P. braconnieri (5), X = 20.8, 15-23; P. cor-
nutum (31), X= 19.2,13-26; P. coronatum
(34),X = 21.3, 16-27; P. ditmarsi (2), Xt =
25.5, 25-26; P. douglassii (27), X = 20.5,
17-26; P. mcallii (13), X = 14.9, 11-18; P.
modestum(9), x = 23.2, 20-25; P. orbicu-
This content downloaded from 128.112.200.107 on Wed, 31 Jul 2013 19:43:16 PM
All use subject to JSTOR Terms and Conditions
[Vol. 45, No. 2
lare (10), x = 21.6, 19-23; P. platyrhinos
(32), xf = 17.4, 13-27; P. solare (14), x =
19.8, 17-23; P. taurus (4), x = 22.2, 21-
24. In terms of actual numbers then, P.
mcallii has the lowest average number of
tooth positions and P. ditmarsi has the
highest average number. However, the lat-
ter value may be an artifact of small sam-
ple size; note that four other species have
limits that overlap those of P. ditmarsi.
Associationsamong the seven standard-
ized morphological variables and the di-
etary percentage of ants are revealed by
correlation analysis (Table 2). Epiptery-
goid length (EP*) shows a strong, positive
correlationwith the height of the coronoid
process (CP*), and it is less strongly cor-
related with the vertical diameter of the
mandible (MD*) and tooth height (TH*).
Epipterygoid length is also negatively cor-
related with the length of the tooth row
(TR*). The vertical diameter of the man-
dible (MD*) shows highly significant pos-
itive correlation with height of the coro-
noid process (CP*)and with distance from
the coronoid process to the articular con-
dyle (LC*). Tooth height (TH*) is posi-
tively correlated with height of the coro-
noid process.Length of the tooth row (TR*)
is negatively correlatedwith distance from
the coronoid process to the articular con-
dyle (LC*), and with diameter of the man-
dible (MD*).
The percentage of ants in the diet shows
very significant negative correlationswith
epipterygoid length (EP*), height of the
coronoid process (CP*), and vertical di-
ameter of the mandible (MD*) and is neg-
atively correlated, though less so, with dis-
tance between the coronoid process and
the articular condyle (LC*). Ant con-
sumption is positively correlated with
length of the tooth row (TR*).
Stepwise regression analysis of the data
was performed in order to obtain a model
that best predicts the diet of the species of
Phrynosoma. Two morphological vari-
ables, TP* and TH*, were deleted from
the analysis because of the lack of signif-
icant correlationswith ants in the diet. Sev-
eral methods of model selection were em-
ployed to find the variables among the
remaining five that contribute most im-
TABLE 1.-Mean ? 1 SE, and observed limits for seven morphological measurements, and mean percentage of a
Abbreviations are: TP = number of tooth positions; TH = mean height of four tallest teeth; LC = length of mandib
length of tooth row; CP = height of coronoid process; MD = vertical diameter of mandible; EP = epipterygoid length
(see Methods and Fig. 1).
Species TP* TH* LC* TR* CP*

asio 0.170 ? 0.004 0.059 ? 0.001 0.320 ? 0.006 0.538 ? 0.006 0.123 ? 0.003 0.171
0.158-0.196 0.050-0.067 0.302-0.354 0.500-0.560 0.107-0.135 0.15
braconnieri 0.258 ? 0.025 0.052 ? 0.003 0.342 ? 0.002 0.511 ? 0.017 0.127 ? 0.003 0.143
0.182-0.289 0.042-0.060 0.337-0.349 0.463-0.540 0.120-0.135 0.10
cornutum 0.190 ? 0.006 0.057 ? 0.001 0.338 ? 0.003 0.518 ? 0.004 0.115 ? 0.002 0.182
0.130-0.260 0.043-0.070 0.313-0.365 0.489-0.560 0.097-0.137 0.16
coronatum 0.200 ? 0.005 0.063 ? 0.001 0.325 ? 0.002 0.525 ? 0.003 0.120 ? 0.003 0.150
0.151-0.240 0.051-0.079 0.303-0.343 0.500-0.556 0.088-0.142 0.12
ditmarsi 0.224 0.062 0.353 0.491 0.146 0
douglassii 0.194 ? 0.007 0.072 ? 0.001 0.338 ? 0.006 0.531 ? 0.006 0.133 ? 0.002 0.177
0.145-0.250 0.059-0.084 0.298-0.385 0.496-0.566 0.114-0.150 0.15
mcallii 0.192 ? 0.005 0.067 ? 0.001 0.304 ? 0.005 0.533 ? 0.005 0.099 ? 0.005 0.143
0.164-0.211 0.061-0.074 0.280-0.329 0.506-0.554 0.082-0.121 0.13
modestum 0.276 ? 0.032 0.047 ? 0.001 0.321 ? 0.007 0.538 ? 0.006 0.084 ? 0.002 0.123
0.149-0.333 0.042-0.052 0.302-0.346 0.512-0.552 0.076-0.092 0.10
orbiculare 0.196 ? 0.008 0.073 ? 0.005 0.330 ? 0.005 0.530 ? 0.002 0.130 ? 0.007 0.173
0.165-0.211 0.053-0.086 0.316-0.345 0.523-0.538 0.115-0.156 0.15
platyrhinos 0.206 ? 0.008 0.056 ? 0.001 0.304 ? 0.004 0.520 ? 0.003 0.108 ? 0.003 0.130
0.152-0.300 0.042-0.070 0.258-0.330 0.490-0.552 0.073-0.126 0.11
solare 0.208 ? 0.008 0.031 ? 0.002 0.316 ? 0.008 0.539 ? 0.006 0.085 ? 0.002 0.129
0.163-0.234 0.021-0.040 0.285-0.351 0.516-0.574 0.070-0.092 0.11
taurus 0.207 ? 0.015 0.058 ? 0.002 0.325 ? 0.013 0.533 ? 0.018 0.127 ? 0.001 0.152
0.192-0.223 0.056-0.060 0.312-0.339 0.514-0.552 0.126-0.128 0.14

This content downloaded from 128.112.200.107 on Wed, 31 Jul 2013 19:43:16 PM

All use subject to JSTOR Terms and Conditions
212 HERPETOLOGICA [Vol. 45, No. 2

ii Ui u u r nri
a U v | 1-m1 " | | uuriii
I a .

onG S~~~~~~~~~~~~~~~~~~

g ~n ~~ H
~ ~ ~ ~ ~I | ~~~~1
| |. I N I 11 II II It Ii U II

FIG. 2.-Mandibles of Phrynosoma: (A) P. asio (AMNH 72636); (B) P. braconnieri (AMNH 90833); (C)
P. cornutum (AMNH77117);(D) P. coronatum (AMNH99685);(E) P. ditmarsi (UAZ 35511); (F) P. douglassii
(AMNH 70639); (G) P. mcallii (AMNH 77042); (H) P. modestum (AMNH 74499); (I) P. orbiculare (RRM
2379); (J) P. platyrhinos (RRM 2323); (K) P. solare (RRM 2324); (L) P. taurus (CAS 141361). Scale is in
millimeters.

This content downloaded from 128.112.200.107 on Wed, 31 Jul 2013 19:43:16 PM

All use subject to JSTOR Terms and Conditions
June 1989] HERPETOLOGICA 213

TABLE 2.-Pearson correlation coefficients (below the diagonal) and the probabilities (above the diagonal)
that the observed r values are significantly different from zero. See Table 1 for abbreviations.

TP* TH* LC* TR* CP* MD* EP* Ants

TP* - 0.1754 0.4714 0.4924 0.4278 0.4961 0.7342 0.8007
TH* -0.418 0.5394 0.6140 0.0201 0.1552 0.0146 0.0951
LC* 0.230 0.197 0.0243 0.0113 0.0070 0.0587 0.0340
TR* -0.219 -0.162 -0.642 0.0448 0.0127 0.0426 0.0358
CP* -0.252 0.657 0.699 -0.587 0.0040 0.0009 0.0006
MD* -0.218 0.437 0.730 -0.691 0.761 0.0277 0.0033
EP* 0.109 0.681 0.559 -0.592 0.828 0.631 0.0002
Ants 0.081 -0.503 -0.613 0.608 -0.843 -0.771 -0.869

portantly to the predictive power of the
model.
The forward selection procedure added
variables one by one to the model until no
remaining variable produced a significant
F statistic. Two variables, EP* and MD*,
entered the model at the 0.10 level of sig-
nificance. The first variable which entered
the model was EP*, with an R2 = 0.7555.
The backward elimination procedure
deleted variables(all were initially entered
into the model) one by one until all vari-
ables remaining in the model produced F
statisticssignificant at the 0.10 level. Three
variables were removed in the following
sequence: TR*, LC*, and CP*, and only
EP* and MD* were retained. The final R12
for the two-variable model, using either
the backward elimination or forward-se-
lection procedures, was R2 = 0.8377.
The maximum R2 improvement pro-
cedure was used to find the best one-vari-
able model, the best two-variable model,
and so forth. This procedure selected vari-
able EP* as having the highest R12. The
second variable added to the model was
MD*, as it produced the greatest increase
in R2= 0.8377 (which is the same as in the
previously mentioned procedures). Under
the maximum R2improvement procedure,
other variables could be substituted after
comparisons with the two variables al-
ready in the model. The comparisonswould
determine if removing one variable and
replacing it with another increased the R2.
Such comparisons did not produce an in-
crease in the R2, and the best two-variable
model, with EP* and MD*, stands. The
best three-variablemodel, containing EP*,
MD*, and CP*, has an R2of 0.8435, which
is not much higher than the R2 value of
the best two-variable model. The equation
for the regressionline in the best two-vari-
able model is Y = -1.205(EP*) -
207.544(MD*) + 122.683. The results of
the three model selection procedures are
identical and are summarized in Table 3.
DISCUSSION
Significantintercorrelationsexist among
some of the morphological variables, as
previously mentioned (Table 2). Epipter-
ygoid length (EP*), which is the single best
predictor of diet in Phrynosoma, shows
strong positive correlation with height of
the coronoid process (CP*), and it is also
correlated,though less strongly, with tooth
height (TH*) and diameter of the man-
dible (MD*). Tooth height is not signifi-
cantly correlated with ants, but variables
CP* and MD* are. The strong intercor-
relation of EP* and CP* apparently pre-
cluded the entry of the latter variable into
the models obtained in the stepwise regres-
sion analysis.In other words, CP* was con-
tributing very little additional variance to
the model not already explained by EP*.
Note the small increase in R2 between the
best two-variable and the best three-vari-
able models obtained by the maximum R2
improvement procedure. This assertion is
also supported in that when EP* was de-
leted from the analysis, CP* was chosen
by stepwise regressionas the variable with
the highest R2 (= 0.7121). A two-variable
model with CP* and MD* had an R2 =
0.7514, which is substantially less than R2
= 0.8377 of the previously obtained two-

This content downloaded from 128.112.200.107 on Wed, 31 Jul 2013 19:43:16 PM

All use subject to JSTOR Terms and Conditions
214 HERPETOLOGICA
TABLE 3. -Summary of the results of stepwise regres-
sion analysis.
Best two-variable model
Source df SS F Prob. > F
Regression 2 3862.3454 23.23 0.0003
Error 9 748.0919
Total 11 4610.4373
Variable TYPE II SS F Prob. > F
MD 379.3226 4.56 0.0614
EP 1120.4696 13.48 0.0051
variable model (with EP* and MD*). Thus
CP* is not as good a predictor of diet in
Phrynosoma as is EP*, either alone or in
combination with MD*.
In the lacertilian skull, the elongate, cy-
lindrical epipterygoid extends almost ver-
tically from the pterygoid to reach the pa-
rietal, or it may slope slightly to attach to
the prootic; see illustrations in McDowell
and Bogert (1954). The epipterygoid is re-
duced or absent in burrowing forms (di-
bamids, amphisbaenians),reduced in some
agamids, and entirely lacking in chamae-
leontids (Camp, 1923). In the genus Phry-
nosoma, the epipterygoid is reduced; the
ventral end of the element rests in a slight
depression in the middle of the pterygoid,
and its dorsalend reaches the anterior bor-
der of the prootic. There is variation in
the length of the element within and be-
tween species, and in P. solare, it may be
greatly reduced or absent (see below and
Table 1). The shorterthe element, the low-
er is its point of attachment to the prootic.
According to Romer (1956), the epipter-
ygoid assistsin stabilizing the palatal com-
plex (vomer, palatine, ectopterygoid, and
pterygoid) of the reptilian skull. In lizards
with well-developed epipterygoids, the
vertical placement of the element between
the pterygoid and the parietal would pro-
vide a strong brace, allowing the palatal
area to better withstand pressuresexerted
during mastication and swallowing of the
prey. In Phrynosoma, the shorter, more
sloping epipterygoid appears to offer a
much less effective brace for the palatal
area. Apparently, selection to maintain the
epipterygoid has been relaxed as a con-
sequence of ant-eating, and evolution has
This content downloaded from 128.112.200.107 on Wed, 31 Jul 2013 19:43:16 PM
All use subject to JSTOR Terms and Conditions
[Vol. 45, No. 2
proceededtowardreductionandlossof the
element. Presch (1969:251)mentioned that
the epipterygoid has lost its function of
support in Phrynosoma, but he did not
elaborate, or discuss its significance in the
context of dietary specialization.
Presch (1969) indicated that the paired
epipterygoidsare absent in P. solare (based
on 18 specimens), whereas Axtell (1986)
found at least some evidence of the ele-
ments in four specimens. Among 10 spec-
imens examined by me, the epipterygoids
are lacking in seven, present in one, and
present but with cartilaginousattachments
to the prootic in two specimens. In the
latter two specimens, the cartilaginousex-
tensions varied from 33-66% of the total
length of the element. Among the speci-
mens lacking the elements, it could not be
determined whether the absence was nat-
ural or due to preparation techniques.
Phrynosoma cornutum is another
species which shows evidence of further
reduction of the epipterygoids. Among 15
specimens examined, four have cartilagi-
nous extensions reaching the prootic. The
cartilage comprises 10-22% of the total
length of the element. In four other spec-
imens, the epipterygoids (one or both) fail
to reach the prootic. Whether cartilagewas
present prior to cleaning of the skull is
uncertain. Phrynosoma cornutum con-
sumes a relatively high proportionof ants,
but less so than in P. mcallii or P. modes-
tum (Table 1), both of which appear to
have fully ossified epipterygoids.
The intercorrelationsamong some mor-
phological variables (Table 2) provide in-
sight as to how structural features are co-
evolving. The strong positive correlation
between epipterygoid length and height
of the coronoid process, for example, sug-
gests that both are functionally related and
responding in the same way to selection
pressuresimposed by dietary change. The
coronoid is the site of insertion for the M.
adductor mandibularis externus and the
M. levator angularis oris (Jenkins and
Tanner, 1968). Presumably,the higher the
coronoid process, the more area is avail-
able for attachment of these muscles,
thereby increasing the masticating power
of the mandibles. In turn, a strengthening
June 1989]
of the epipterygoids would be required to
brace the pterygoids when prey items are
being subdued and crushed. The vertical
diameter of the mandible (MD*)and tooth
height (TH*) are also positively correlated
with the height of the coronoid process, as
would be expected. The variable MD* is
a measure of the robustness of the man-
dible, and TH* may reflect the grasping
and masticating capabilities of the teeth.
The distance from the coronoid process to
the articular condyle (LC*) is negatively
correlated with the length of the tooth row
(TR*). A slight anterior shift of the coro-
noid process would diminish TR*, but in-
crease LC*, thus providing more area for
muscle attachment. Two muscles, the M.
pseudotemporalis profundus and the M.
adductor mandibularis posterior, insert
onto this area (Jenkinsand Tanner, 1968).
A slight anterior shift of the coronoid
process would perhaps also enhance the
lever action of the mandibles by moving
the main point of muscle pull (coronoid
process) closer to the area bearing the
weight (TR*), and farther from the ful-
crum (articularcondyle). Note that TR* is
also negatively correlated with MD*, CP*,
and EP*.
Specialization for myrmecophagy in
Phrynosoma appears to have involved the
following morphological changes: (1) a re-
duction or loss of the epipterygoid; (2) re-
duction of the coronoid process; (3) re-
duction in the diameter of the mandibular
ramus; (4) reduction in the area posterior
to the coronoid process, and a concomitant
increase in the length of the tooth row,
possibly accomplished by a slight shift in
the position of the coronoid. Such mor-
phological changes have evolved, in var-
ious combinations,in Phrynosoma mcallii,
P. modestum, P. platyrhinos, and most
notably P. solare. Phrynosoma solare has
the highest consumption of ants compared
with other members of the genus (Table
1). The specialization for myrmecophagy
is corroborated not only by morphology
and dietary information,but by behavioral
observationsas well. When fed large prey
items such as crickets, adult P. solare at-
tempted to catch the insects by extruding
the tongue. Regardless of size, the prey
This content downloaded from 128.112.200.107 on Wed, 31 Jul 2013 19:43:16 PM
All use subject to JSTOR Terms and Conditions
HERPETOLOGICA 215
items were treated as though they were
ants, and such attempts to ingest prey were
usually unsuccessful. The prey was simply
too large to be held by the sticky tongue
without assistance from the jaws. Even-
tually, some lizards learned to catch the
large prey by using the jaws. Phrynosoma
solare also has the smallest teeth of any
member of the genus (Table 1). The small
size of the teeth is probably a consequence
of diet, although there is no significantcor-
relation between tooth height (TH*) and
ant consumption across species (Table 2).
The remaining species appear to be less
morphologically specialized for ant-eat-
ing. Phrynosoma ditmarsi appears to con-
sume the lowest proportion of ants (Table
1), but comparatively little is known of its
feeding habits in nature. Roth (1971, per-
sonal communication) found weevils, oth-
er beetles, true bugs, grasshoppers,spiders,
and scorpions, in addition to ants, in the
stomachs of specimens. Phrynosoma dit-
marsi has enormous vertical expansion of
the mandible, a tall coronoid process,more
extensive area posteriorto that process,and
a long epipterygoid. The teeth are not ex-
ceptionally tall; four other species surpass
P. ditmarsi in relative tooth height. How-
ever, the crowns of the teeth are quite
flattened. This suite of morphological fea-
tures suggests greater crushing power in
P. ditmarsi compared with other Phry-
nosoma. Behavioralobservationslend sup-
port to this idea. P. ditmarsi is more effi-
cient in capturing large prey than is P.
douglassii or P. orbiculare (Montanucci,
unpublished data). The two latter species
frequently beat the prey against the sub-
strate with lateral movements of the head,
apparently to subdue it; P. ditmarsi sel-
dom, if ever, does this. Moreover, P. dit-
marsi masticates its prey somewhat more
than do the other two species. Although P.
ditmarsi may be regarded as a dietary gen-
eralist, its morphology suggests specializa-
tion for eating large, chitinous arthropods.
I speculate that such prey items may be
periodicallyabundantand importantin the
diet of this horned lizard, but field studies
are needed to provide concrete data on
various aspects of the natural history of
this rare and interesting species.
216 HERPETOLOGICA
Acknowledgments.-I thank the following cura-
tors and museum staff for the loan of skeletal speci-
mens used in this study (museumabbreviationsfollow
Leviton et al., 1980): G. W. Foley, R. G. Zweifel
(AMNH);R. C. Drewes, A. E. Leviton, J. V. Vindum,
L. Wishmeyer(CAS);R. F. Inger, H. Marx,A. Resetar
(FMNH); W. E. Duellman, S. K. Dutta, D. R. Frost
(KU);R. L. Bezy, J. W. Wright (LACM);P. Alberch,
J. P. Rosado (MCZ);D. A. Good, H. W. Greene, D.
B. Wake (MVZ);C. H. Lowe (UAZ); D. M. Harris,
A. G. Kluge(UMMZ);R. W. McDiarmid,A. D. Yoder,
G. R. Zug (USNM);J. A. Campbell (UTA);T. R. Van
Devender (TRV, personal collection). My personal
specimens (RRM)used in this study will be deposited
in the LACM. Computer time and facilities were
made available through the Clemson University
Computer Center; William C. Bridges,Jr. and Louis
Deluca kindlyprovidedtechnicalassistanceand help-
ful discussions.
LITERATURE CITED
AXTELL, R. W. 1986. The epipterygoids in Phry-
nosoma solare, still present, but disappearing. J.
Herpetol. 20:79-81.
CAMP, C. L. 1923. Classification of the lizards. Bull.
Am. Mus. Nat. Hist. 48:289-481.
HOTTON, N., III. 1955. A survey of adaptive rela-
tionships of dentition to diet in the North American
Iguanidae. Am. Midl. Nat. 53:88-114.
JENKINS, R. L., AND W. W. TANNER. 1968. Os-
teology and myology of Phrynosoma p. platyrhinos
Girard and Phrynosoma d. hernandesi Girard.
Brigham Young Univ. Sci. Bull. Biol. Ser. 9(4):1-
34.
LEVITON, A. E., R. McDIARMID, S. MOODY, M.
NICKERSON, J. ROSADO, 0. SOKOL, AND H. VORIS.
1980. Museum acronyms-Second edition. Her-
petol. Rev. 11:93-102.
This content downloaded from 128.112.200.107 on Wed, 31 Jul 2013 19:43:16 PM
All use subject to JSTOR Terms and Conditions
[Vol. 45, No. 2
MCDOWELL, S. B., JR., AND C. M. BOGERT. 1954.
The systematic position of Lanthanotus and the
affinities of the anguinomorphan lizards. Bull. Am.
Mus. Nat. Hist. 105:1-142.
MONTANUCCI, R. R. 1979. Notes on systematics of
horned lizards allied to Phrynosoma orbiculare
(Lacertilia: Iguanidae). Herpetologica 35:116-124.
. 1981. Habitat separation between Phry-
nosoma douglassi and P. orbiculare (Lacertilia:
Iguanidae) in Mexico. Copeia 1981:147-153.
PIANKA, E. R., AND W. S. PARKER. 1975. Ecology
of horned lizards: A review with special reference
to Phrynosomaplatyrhinos.Copeia 1975:141-162.
POWELL, G. L., AND A. P. RUSSELL. 1984. The diet
of the eastern short-horned lizard (Phrynosoma
douglassi brevirostre)in Alberta and its relation-
ship to sexual size dimorphism. Can. J. Sci. 62:428-
440.
PRESCH, W. 1969. Evolutionary osteology and re-
lationships of the horned lizard genus Phrynosoma
(family Iguanidae). Copeia 1969:250-275.
RISSING, S. W. 1981. Prey preferences in the desert
horned lizard: Influence of prey foraging method
and aggressive behavior. Ecology 62:1031-1040.
ROMER, A. S. 1956. Osteology of the Reptiles. Uni-
versity of Chicago Press, Chicago.
ROTH, V. D. 1971. Food habits of Ditmars' horned
lizard with speculations on its type locality. J. Ar-
izona Acad. Sci. 6:278-281.
SAS INSTITUTE INC. 1985. SAS User's Guide: Sta-
tistics, 1985 ed. SAS Institute, Cary, North Carolina.
TURNER, F. B., AND P. A. MEDICA. 1982. The dis-
tribution and abundance of the flat-tailed horned
lizard (Phrynosomamcallii). Copeia 1982:815-823.
Accepted: 22 August 1988
Associate Editor: William Cooper, Jr.