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Ecology of the whiptail lizard Cnemidophorus deppii on a tropical beach

Article  in  Canadian Journal of Zoology · February 2011

DOI: 10.1139/z93-334

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 Ecology of the whiptail lizard Cnemidophorus deppii on a tropical beach
LAURIEJ. VITT, PETERA. ZANI,JANALEE
P. CALDWELL,
A N D RICHARD
D. DURTSCHE
Oklahoma Museum of Natural History and Zoology Department, University of Oklahoma,
Can. J. Zool. Downloaded from www.nrcresearchpress.com by "Institute of Vertebrate Paleontology and Paleoanthropology,CAS" on 06/10/13

Norman, OK 73019, U.S.A.

Received May 26, 1993
Accepted September 10, 1993

VITT, L.J., ZANI,P.A., CALDWELL, J.P., and DURTSCHE,R.D. 1993. Ecology of the whiptail lizard Cnemidophorus deppii
on a tropical beach. Can. J. Zool. 71: 2391 -2400.
The whiptail lizard Cnemidophorus deppii was studied during late dry season on a tropical beach on the Pacific coast of
Nicaragua. Most aspects of the ecology of this species are similar to those of other active foraging lizard species studied.
Individual C. deppii spend most of a typical daily activity period on sand moving from vegetation patch to vegetation patch,
presumably in search of food. The amount of time spent in the sun is greatest in early morning and at its lowest level at
+
midday. The average rate of movement was 0.048 0.004 m/s. Body temperatures of active lizards averaged 40.0 0.25"C, +
and most activity occurred during morning and late afternoon. Body temperatures were significantly lower in whiptails active
during the morning than later in the day. Forty-two types of prey were identified in stomachs, with termites, spiders, and
various orthopterans accounting for most of the diet volumetrically. There was no correlation between lizard size and prey
size. There was a significant negative relationship between prey width and the number of prey in stomachs. Snout-vent
length (SVL) at sexual maturity was 60 mm for females and 58 mm for males. Mature females averaged 63.8 f 0.7 mm
SVL and produced clutches varying from 1 to 3 eggs ( 1 = 1.8 +
0.2). Oviductal eggs averaged 13.6 +
0.64 x 7.7 +
0.2 1 mm in size. There was no significant relationship between female SVL and clutch size. Relative clutch mass was similar
to that for other active foraging lizard species. Sexual dimorphism was apparent in coloration (males brightly colored), body
size (males larger), and relative head size (male heads larger independent of body size differences). These differences pre-
sumably are due to sexual selection.

VITT, L.J., ZANI,P.A., CALDWELL, J.P., et DURTSCHE, R.D. 1993. Ecology of the whiptail lizard Cnemidophorus deppii
on a tropical beach. Can. J. Zool. 71 : 2391 -2400.
For personal use only.

Le lkzard Cnemidophorus deppii a fait l'objet d'une ktude a la fin de la saison skche, sur une plage tropicale de la cate
du Pacifique au Nicaragua. Presque tous les aspects de l'kcologie de cette espkce s'apparentent a ceux qui prkvalent chez
d'autres espkces de lkzards qui doivent faire une recherche active de leur nourriture. Les lkzards passent la plus grande partie
de leur pkriode quotidienne d'activitk sur le sable, se dkpla~antd'une touffe de vkgktation a une autre, probablement en qu2te
de nourriture. La durke des pkriodes passkes au soleil est maximale au dkbut de la matinke et minimale au milieu de la journke.
La vitesse moyenne des dkplacements est de 0,048 +0,004 m/s. La tempkrature corporelle des lkzards actifs est de 40 +
0,25"C en moyenne et les lkzards sont actifs surtout le matin et a la fin de l'aprks midi. La tempkrature du corps est
significativement plus basse chez les lkzards actifs le matin que chez les lkzards actifs plus tard au cours de la journke.
Quarante-deux types de proies ont kt6 reconnus dans les estomacs et ce sont les termites, les araignkes et des orthoptkres
divers qui constituent le plus gros du volume du rkgime alimentaire. I1 n'y a pas de corrklation entre la taille des lkzards
et la taille des proies. I1 y a une corrklation nkgative significative entre la largeur des proies et leur nombre dans les estomacs.
La taille a la maturitk sexuelle est de 60 mm (SVL = longeur museau-kvent) chez les femelles et de 58 mm chez les miiles.
Les femelles a maturitk mesurent en moyenne 63,8 f 0,7 (longueur SVL) et pondent des couvkes de 1 a 3 oeufs ( 1 =
+
1,8 0,2). Les oeufs dans l'oviducte mesurent en moyenne 13,6 0,64 x 7,7 + + 0,21 mm. I1 n'y a pas de corrklation
significative entre la longueur SVL et le nombre d'oeufs par couvke. La masse relative d'une couvke est semblable a celle
des couvkes d'autres espkces de lkzards qui recherchent activement leur nourriture. I1 existe un dimorphisme sexuel kvident
dans la coloration (miiles a coloration vive), la taille (miles plus grands) et la taille relative de la t2te (miiles a t2te plus grande,
indkpendamment de la taille globale). Ces diffkrences sont sans doute dues a la sklection sexuelle.
[Traduit par la rkdaction]

Introduction Cnemidophorus deppii on a tropical beach in Central America.

Lizards in the teiid genus Cnemidophorus are widespread in These sand-covered beaches offer a habitat low in structural
the New World, occurring from the northern United States to diversity that is exposed to high daily temperatures owing to
Argentina (Wright 1993). They have caught the attention of direct sun on western exposures during most of the day. Further-
behaviorists (Crews and Moore 1993; Leuck 1993), ecologists more, the potential exists for a high resource base comprising
(Pianka 1970, 1986; Schall 1977, 1993; Case 1983; Milstead intertidal invertebrates associated with the advance and retreat
1957a, 1957b, 1965; Price et al. 1993; Cuellar 1993), physiol- of the tide. The costs of harvesting this resource may be high,
ogists (Bowker 1993; Garland 1993; Wit and Sellars 1993) and including digging and increased exposure to high environmental
evolutionary biologists (See citations in Wright 1993) because temperatures and predators. We asked the following questions:
of their high visibility, observable behavio~s(e.g., Anderson (i) Given the low structural diversity of the beach habitat, how
1993), high levels of activity, and the occurrence of both sexu- do these lizards distribute themselves among microhabitats?
ally and parthenogenetically reproducing species (Cuellar 1971; (ii) How do these lizards balance daily activity associated with
Cole 1975; many others). These lizards often occur in environ- foraging against the thermal extremes of the beach habitat?
ments that would appear to cause extreme thermal stress as a (iii) Given the apparent availability of marine crustaceans, iso-
result of high temperatures and low humidity (e.g ., Pianka 1970). pods, and molluscs, has the diet of C. deppii shifted to take
In this study, we examined the ecology of a population of advantage of this presumably rich food source? In addition, we
Printed in Canada 1 Imprime au Canada
 2392 CAN. J . ZOOL. VOL. 71. 1993

present limited information on reproductive characteristics of TABLE1. Microhabitat, exposure, and behavior
this population and make comparisons with another study con- of individual Cnemidophorus deppii observed
ducted on this species (Fitch 1973). on a tropical beach in Nicaragua based on
haphazard transects
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Methods No. of Percentage

The field site, located 5.0 km north of Salinas in Departamento lizards of total
Rivas on the Pacific coast of Nicaragua (1 1"28'N, 86"8'W), is a
sandy beach with scattered vegetation creating a habitat that is exposed Microhabitat
to direct sunlight throughout the day. Only two other lizard species Rock
were observed in this habitat. Sceloporus variabilis (Phrynosomatidae) Leaf litter
was common and partially arboreal, and Phyllodactylus tuberculosus, Tree
a nocturnal gekkonid, was found under surface objects. Adjacent habitat Open sand
is xeric woodland (see Terin and Incer Baquero 1964), typical of the Sand under grass
Pacific coast of Central America, and includes several additional lizard Sand under shrub
species (e.g., Ameiva undulata, Mabuya unimarginata, and others). Exposure
Field studies were conducted during March 1993, late in the dry Sun
season. The west-facing beach increased exposure to afternoon sun. Shade
Our procedures for collecting data on activity, microhabitat use, Clouds
behavior, exposure, and activity temperatures were as follows.
Behavior
First, two investigators haphazardly walked through the habitat
Foraging
collecting the following data for each individual lizard observed or
Basking
collected: (i) Exposure. Three categories were identified: sun, shade,
Social interaction
and clouds. We considered clouds to be a distinctly different category
from shade because lizards active when clouds blocked the sun did
not have the option of moving into the sun for thermoregulatory pur-
poses as did lizards active in shade. (ii) Behavior. Three categories individual lizards were actually moving, as opposed to not moving,
were identified: foraging, basking, and social interaction. Lizards while on the surface. (ii) Exposure. We recognized four categories:
moving around searching were considered foraging for prey. Lizards shade, filtered sun, sun, and clouds. Filtered sun was added to our
For personal use only.

that were stationary, usually pressed against the substratum regardless exposure categories to represent lizards under the canopy of vegeta-
of exposure, were considered basking. Lizards clearly interacting tion receiving a combination of sun and shade owing to the filtering
with a conspecific were considered engaging in social interactions. effect of the vegetation. Specifically, we use "filtered" to refer to
(iii) Microhabitat. Six microhabitats were used by C. deppii: rock, a nearly uniform infusion of light mixed with shadows from small
leaf litter, in shrub or tree (trunk, branch, or limb), open sand, sand leaves in the vegetation ranging from 0.5 to 2.5 m off of the ground.
under grass, and sand under shrubs. For lizards observed on parts of (iii) Corrected distance. This is the actual distance moved in a 600-s
trees or shrubs, we estimated distance off the ground in cm. (iv) Activity. trial. We corrected the distance data for the six individuals observed
The time each lizard was observed was recorded to estimate frequency for shorter periods by calculating a distance per second rate and
of encounter throughout the activity period. An independent estimate multiplying by 600. (iv) Overall rate of movement. The rate was the
of hourly activity was made by having the other two investigators distance moved on a per unit time (s) basis. No animal was used for
simultaneously walk parallel transects at hourly intervals and count more than one trial.
active lizards. The transects were selected a priori based on observed Body size was estimated for each collected lizard by measuring the
high abundances of lizards, and the same transect was monitored in distance from snout to vent (SVL) to 1 mm with a linear rule. Addi-
exactly the same manner during each trial. This provides an unbiased tional morphological measurements included tail length, length of the
estimate of activity, removing the effects of varying habitats, varying regenerated portion of the tail (if any) to 1 mm, total mass to 0.1 g
personnel, and unequal effort. (v) Thermul ecology. A sample of lizards (taken with Pesola@spring balances), and the following linear measure-
was collected with air rifles (using BBs) and cloacal, substratum, and ments to 0.1 mm (taken with electronic calipers): head width measured
air temperatures were taken immediately (within 1 min) and recorded at the widest point of the skull, head length as the distance from the
using Miller and Webera rapid-registering thermometers. These lizard anterior edge of the tympanum to the tip of the snout, head height at
samples were used for other purposes as well (see below). As an the highest point of the skull, body width and height at mid-body,
independent measure of the thermal milieu these lizards experience, hind-leg length and fore-leg length as the distance from the body to
we placed three thermocouples at various places in the microhabitats the tip of the longest toe. Our sample of 81 lizards (53 males and
used by C. deppii. One was placed 1 mm below the sand surface in 28 females) was the minimum we considered adequate to accurately
shade, one was placed 1 mm below the sand surface in sun, and a represent the population. We are confident that our sampling had
third was shaded and placed 1 cm above the substratum. Tempera- little if any impact on the population for two reasons. First, this wide-
tures were recorded at 15-min intervals beginning just before lizard spread whiptail lizard is likely the most abundant species in beach
activity commenced (08:00), and ended when lizard activity ceased habitats of dry forest on the Pacific coast of Central America (e.g.,
(19:00) using an Omega OM550 data logger. Fitch 1973). Second, on a return trip to the same locality in April,
To further assess activity, particularly as it related to thermoregula- lizards appeared more abundant than they had been in March, possibly
tion and foraging, we conducted observational studies on 39 focal as a result of immigration from adjacent dry forest habitats that were
animals. Individual lizards were randomly selected and observed con- being burned.
tinuously. Six of the 39 lizards were observed for time periods ranging Lizards were properly tagged, preserved in 10% buffered formalin,
from 226 to 510 s, and the other 33 were observed for 600 s each. and later transferred to 70% ethanol. Most preserved specimens were
Typically, lizards were observed from approximately 10 m. These deposited in the herpetological collections at the Oklahoma Museum
were considered to be behaving normally if they continued to forage, of Natural History (OMNH), although a sample of the collection was
bask, or engage in social interactions. During the focal studies, obser- deposited at the National Museum of Nicaragua in Managua.
vations were recorded on microcassette recorders and later transcribed. Preserved lizards were opened within three weeks after collection
This allowed maximum information collection and continuous monitor- for the examination of reproductive tracts and stomachs. Females
ing of individual lizards. Our efforts were directed toward measuring were considered sexually mature if they contained oviductal eggs or
the following variables: (i) Movement. We recorded the amount of time enlarged vitellogenic follicles. Males were considered sexually mature
 VITT ET AL.

TABLE2. Summary of focal animal studies on 39 individual Cnemidophorus deppii

Category .f k SE Min./max.

Movement
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% time not moving

% time moving
Overall rate of movement (rnls)
Corrected rate of movement (rnls)
Corrected total distance traveled (m)
Exposure
% time in sun
% time in shade
% time in filtered sun
% time under clouds
NOTE:The overall rate of movement is the rate based on the distance moved during the entire time period
(600 s). The corrected rate of movement is based on the distance moved during the proportion of time actually
in movement.

if they contained enlarged testes and convoluted epididymides. Eggs

or vitellogenic follicles were measured for length and width (to the shade
Cnemidophorvs deppii sun
nearest 0.01 mm) with electronic calipers, and egg volume was calcu-
filtered sun
lated with a modified version of the formula for a prolate spheroid:
volume = 4 / 3 ~ ( 1 / 2length)(l/2 width)2
Removed stomachs were opened and prey items were separated and
identified to family or the lowest taxon possible. Individual prey items
For personal use only.

were measured for length and width with electronic calipers (to the
nearest 0.01 mm). Volume for each individual prey item was esti-
mated with the above formula.
The reciprocal of Simpson's (1949) niche breadth measure was cal-
culated for numerical and volumetric diet data:
. . . . . .
9:OO 1O:OO 11:OO 12:OO 13:OO 14:OO 1 5 0 0 16:OO 17:OO 18:OO

Time of Day
where i is the resource category, p is the proportion of resource FIG. 1. Exposure of individual Cnemidophorus deppii based on
category i, and n is the total number of categories. Niche breadth focal animal studies. The time value for each exposure category is a
values vary from 1 (exclusive use of one category) to n (even use of percentage based on means for 39 individuals.
all categories) (Pianka 1986).
Most statistical comparisons were made with standard parametric
tests when assumptions could be reasonably met. To compare male and number of lizards first sighted under cloud cover simply reflects
female morphology, we calculated the log,, of each morphological the absence of clouds during much of the time period that we
characteristic, regressed these on the log,, of SVL, and performed studied these lizards. Approximately half of the lizards were
ANOVAs on the regression residuals to remove the effect of body observed foraging, approximately half were basking, and very
size, with sex as the factor. ANOVAs were used to compare body few were engaged in social interactions (Table 1).
temperatures of lizards during various activities, experiencing different In focal animal studies, we added the category "filtered
exposures, and occurring in different microhabitats. This was consid- sun. " Focal animal studies revealed that C. deppii spend most
ered justified even though there were significant correlations between of their active time in shade or filtered sunlight (Table 2), thus
body temperatures and environmental temperatures (substratum and reducing their exposure to direct sun. A significant portion of
air), because the R2 values and slopes were low. Moreover, an
time is spent basking against the substratum for thermoregula-
analysis of body temperatures on an hourly basis revealed that much
of the correlation was due to low early morning body temperatures tory purposes. During early morning they bask on sand in sun-
when lizards first became active (see below). Body temperatures light, presumably to gain heat because the sand surface in the
remained relatively high throughout the remainder of the activity period. sun heats rapidly (Fig. 5). During midday and later, most of
Most statistical tests were performed with the Macintosh version of the "basking" takes place in shade, presumably facilitating heat
Statview (Abacus Concepts 1987). Unless otherwise indicated, means loss. The amount of time spent in the shade increased through-
are presented k one standard error. out the morning to midday (Fig. 1). More than half of the time
is spent moving through the habitat in search of prey, mates,
Results or thermoregulatory sites. The distance moved during given
time periods varied considerably among individuals as did the
Microhabitat distribution, exposure, and bihavior rate of movement.
More than half of the 98 lizards for which we collected data
based on haphazard transects were first sighted in shade with sun Thermal ecology and time of activity
available (Table 1). In this data set, lizards under the canopy of The body temperatures of 61 active C. deppii varied from
plants were considered to be in the shade even though they may 33.6 to 43.5"C (1= 40.0 f 0.25"C) with most active body
have experienced filtered sun (see below). The relatively low temperatures registering above 39 "C (Fig. 2). The lowest
 2394 CAN. J . ZOOL. VOL. 71, 1993

Cnemidophorus deppii
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32 34 36 38 40 42 44
Body Temperature ('C)
FIG. 2 . Histogram showing the distribution of body temperatures
for active Cnemidophorus deppii on a coastal beach in western
Nicaragua. The mean body temperature of active individuals is indi- 8:OO
I

9:OO
I

1O:OO
I

11:OO
I

12:OO
I

13:OO
I

14:OO
I

15:OO
I

16:OO
I

17:OO 18:OO
cated by the arrow.
Time of Day
FIG.4 . Relationship between hourly mean body temperatures and
time of day for active Cnemidophorus deppii. Means f 1 SE are
Cnemidophorus deppii
shown. Numbers above each hourly mean indicate sample size. N D
0 indicates no data for a given time period. See text for statistical
comparisons.
For personal use only.

. .
29 30 3i 32 33 34 35 36 3>

Air Temperature ("C)

FIG.3. Relationship between lizard body temperature and air tem-
perature for active Cnemidophorus deppii. The 95 % confidence limits
for the true mean of y are shown as is the regression line.

temperatures were recorded from individuals collected early

in the morning, apparently just after emergence from burrows.
Substratum and air temperatures associated with active lizards
averaged 37.6 f 0.61 and 32.9 f 0.2g°C, respectively. The
lowest substratum and air temperatures recorded during lizard
activity were 29.5 and 29.0°C, respectively, whereas the highest
substratum and air temperatures associated with lizard activity
were > 50 and 36.8"C. There were significant correlations
between body temperatures and both substratum and air tem-
peratures. Substratum temperatures accounted for only 26% Time of Day
of the variance in lizard body temperatures (F[I,59]= 20.3, FIG. 5 . Daily progression of environmental temperatures in the
p < 0.001) and the slope of the regression line was only 0.2. microhabitat used by Cnemidophorus deppii (see Methods).
Air temperatures accounted for 32% of the variation in body
temperatures (F[ = 27.3, p < 0.00 1) and the slope of the
regression line was 0.5 (Fig. 3). Even though body temperatures differences resulting from time of activity (F15,51= 6.3, p <
were correlated with environmental temperatures, a simple 0.001). The body temperatures of lizards on the surface between
regression analysis does not accurately reflect determinants of 09:OO and 10:OO were significantly lower (based on pairwise
body temperatures for C. deppii. Typically, these lizards emerge post-hoc Scheffk's F tests, p < 0.05) than the body tempera-
early in the morning from night-time retreats with relatively tures of lizards active from 11:00 to 12:00, 12:00 to 13:00, and
low body temperatures, heat up relatively rapidly, remain very 15:00 to 16:00. No other comparisons were significant.
warm throughout activity (although most individuals become An ANOVA on lizard body temperatures with behavior
inactive during midday), and experience reduced body tem- (foraging, basking, social interaction) as the factor shows that
peratures only at the very end of the day (Fig. 4). Indeed, a lizard body temperatures were not influenced by behavior
significant relationship exists between body temperatures and (FL2+58]= 1.0, p = 0.385). There was, however, a significant
time of activity (R2 = 0.12, F11,591 = 27.3, p < 0.001), which effect of exposure on lizard body temperatures (F[2,581= 6.4,
is primarily a result of lower lizard body temperatures early p = 0.003), with lizards active while sun was available having
in the morning (Fig. 4). An ANOVA on temperatures grouped body temperatures 3.1 "C (Scheffk's F = 6.1 1, p < 0.05) and
by hour (including only samples with n 1 3) revealed significant 2.9"C (Scheffk's F = 5.69, p < 0.05) higher (sun and shade,
 VITT ET AL.

TABLE3. Composition of the diet of Cnemidophorus deppii from the Pacific coast
of Nicaragua

Volume Volume
Prey tY Pe No. No. (%) (mm3) (%I Frequency
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Thy sanura 1
Odonata 1
Orthoptera (unid.) 1
Tetrigidae 1
Tettigoniidae 6
Gry llidae 5
Mantidae 2
Phasmatidae 6
Blattidae 6
Acrididae 8
Isoptera
Termitidae 884
Dermaptera 1
Hemiptera
Miridae 2
Reduviidae 1
Pentatomidae 3
Homoptera
Cicadidae 1
Cercopidae 1
Achilidae 1
For personal use only.

Coleoptera (unid.) 1
Cicindelidae 1
Carabidae 3
Staphylinidae 3
Buprestidae 1
Tenebrionidae 5
Cerambycidae 1
Chrysomelidae 1
Diptera (unid.) 18
Culcidae 7
Drosophilidae 1
Hymenoptera (unid.) 1
Formicidae 25
Insect larvae
Eruciform 12
Elateriform 1
Campodeiform 3
Vermiform 4
Chilopoda 1
Pseudoscorpionida 1
Phalangida 1
Araneae 21
Crustacea 6
Mollusca 1
Plant parts (seeds) 20
Totals 1070
NOTE:Frequency is the number of lizards containing a particular prey item. unid., indicates individuals identi-
fied only to order.

respectively) than lizards active when the sun was covered by in which lizards were actually sampled (F[4,56j = 0.7,
clouds. These differences appear to result, at least partly, from p = 0.578).
differences in substratum (F[2,581= 1.0, p = 2.98, p = 0.059, Surface temperatures of the beach in sun vary greatly during
marginally significant) and air temperatures (F[2,58]= 6.2, p = the day, ranging from less than 30°C in early morning to 60°C
0.004) resulting from exposure. There were no significant in mid-afternoon (Fig. 5). Sand temperatures in the shade are
differences in substratum temperatures among microhabitats considerably lower, but do not decrease rapidly at the end of
 CAN. J. ZOOL. VOL. 71. 1993

Haphazard census
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Hourly census

- -1 -0.5 0 0.5 1 1.5 2 2.5 3 3.5

- Log ,,(Prey Volume)

- - FIG.7 . Distribution of the log,, of individual prey volumes show-

- - ing the predominance of small prey (mostly termites) in the diet of
Cnemidophorus deppii.
7 - 7 - . . . . . - r .

Time of Activity (0.10- 1772.75 mm3). The prey size distribution was heavily
FIG.6. (A) Daily activity of Cnemidophorus deppii based on obser- skewed toward small prey even though some individual lizards
vations made during haphazard lizard sampling. (B) Daily activity of consumed a single very large prey item. There were no signifi-
Cnemidophorus deppii based on a repeated hourly census conducted cant correlations between any measure of prey size (length,
on a single transect. width, or volume) and lizard SVL (all p values > 0.19). Loglo
transformation of prey size data followed by similar regression
For personal use only.

analysis produced poorer relationships (all p values > 0.35).

the day as do temperatures in the sun, most likely because of There was, however, a significant negative relationship between
low rates of convective heat loss. Air temperatures remain prey width and the number of prey in stomachs (F11,63j = 10.6,
below surface temperatures in sun and shade. Based on obser- p = 0.002). The slope of the regression was low (slope =
vations of active lizards during the sampling period, lizard -0.01, t = 3.25, p = 0.002).
activity is greatest during mid-morning through early after-
noon (Fig. 6). Hourly lizard censuses revealed a similar pat- Reproduction and sexual dimorphism
tern of activity (Fig. 6). Females varied from 48 to 72 mm SVL, and the smallest
female showing evidence of reproduction was 60 mm, which
Prey utilization we consider size at sexual maturity. Twenty sexually mature
Sixty-three C. deppii contained from 1 to 234 (1= 16.7 f females averaged 63.8 f 0.7 rnrn SVL. Clutch size varied from
5.1) prey items. There was no relationship between the number 1 to 3 eggs averaging 1.8 f 0.2 (n = 17). Seven oviductal eggs
of prey items and lizard SVL (FL1 ,621 = 1.2, p = 0.2651). from three females averaged 13.60 f 0.64 mm in length,
Forty-two prey categories were identified, including insects, +
7.66 0.21 mm in width, 420.36 f 29.85 mm3 in volume,
their larvae, arachnids, other arthropods, molluscs, and seeds and 0.37 f 0.02 g in mass. Although larger females tended
(Table 3). Termites made up more than 80% of the prey items to produce larger clutches, there was no significant relation-
taken by C. deppii on a numerical basis, with ants, spiders, ship between female SVL and clutch size (F, 161 = 0.5, p =
seeds, and unidentified flies making up most of the remainder. 0.485). The three females with oviductal eggs averaged 7.43 f
Volumetrically, termites were also important, accounting for 1.03 g in total mass (including eggs), their clutches averaged
14.5% of the diet. However, orthopterans (roaches, walking +
0.87 0.18 g, clutch mass / total mass averaged 0.11 0.01, +
sticks, grasshoppers, and crickets) as a group made up more and clutch mass / body mass averaged 0.12 +
0.01. Males
than 50% of the diet by volume. Niche breadth for prey num- reach sexual maturity at 58 mm SVL. At least some reproduc-
bers was 1.46, indicating many individuals of just a few prey tion takes place during the late dry season based on the presence
types (termites), whereas niche breadth for prey volumes was of females containing oviductal eggs or corpora lutea indicat-
10.89, indicating that on a volumetric basis several prey items ing recent oviposition. Four females contained simultaneously
were important components of the diet. The occurrence of many enlarged vitellogenic follicles and corpora lutea indicating that
termites and an array of other prey that are not surface active at least two clutches of eggs are laid by individual females.
coincident with lizard activity is consistent with our observa- Males attain larger size than females (Fig. 8, Table 3) and
tions of these lizards frequently digging for prey. Only two mean SVL of mature males is significantly greater than for
prey items, termites and spiders, were taken by more than females Mann-Whitney U test, Z = -5.13, p < 0.001).
30% of the lizards sampled (Table 3). Mean total mass was also greater for males than for females
Mean prey size was highly variable, but most prey were small (Mann-Whitney U test, Z = -5.34, p < 0.001). Heads of
in size (Fig. 7). Prey length averaged 4.44 f 0.18 mm (1.56 - males are significantly larger than heads of females when the
144.53 mm), prey width averaged 1.42 f 0.03 mm (0.35 - effect of body size is removed (Table 4, Fig. 9) and the same
10.43 m), and prey volume averaged 14.99 f 2.63 mm3 is true for hind leg length.
 VITT ET AL.

Cnernidophorus deppii
1 Cnemidophorus deppii
l3 1
males
Can. J. Zool. Downloaded from www.nrcresearchpress.com by "Institute of Vertebrate Paleontology and Paleoanthropology,CAS" on 06/10/13

A females

45 50 55 60 65 70 75 80 85 90

Snout-Vent Length (mm)

FIG. 8. Distribution of snout-vent lengths for Cnemidophorus
deppii. Arrows indicate size at sexual maturity for each sex. L ~ g , ~ ( S n o u t - V e nLength)
t

FIG. 9. Regression of log,, of head width versus log,, snout-vent

length for Cnemidophorus deppii showing divergence in head size in
TABLE4. Descriptive statistics for morphological charac-
teristics of sexually mature Cnemidophorus deppii adults (see Table 5 for statistical comparisons).

Males Females
season. On the other hand, the beaches may provide a rich
Snout -vent length (mm) resource base for lizards. A diversity of invertebrates is asso-
ciated with litter on the beach, including numerous small spe-
Mass (g) cies of crabs, isopods, and molluscs. In addition, at least some
of the driftwood deposited high on the beaches is utilized by
Head width (mm) termites and numerous insect larvae. These invertebrates should
For personal use only.

provide a water source (preformed) as well as a food supply

Head length (mm)
for lizards, although the potential exists for solute levels to be
Head height (mm) high in beach invertebrates. Presumably utilization of these
beaches by lizards represents a balance between the costs of
Body width (mm) thermal stress (Huey and Slatkin 1976) or reduced activity,
and the benefits of foraging (Anderson 1993; Anderson and
Body height (mm) Karasov 1981) in relatively rich areas.
A majority of C. deppii activity occurs during mid to late
Hind-leg length (mm) morning and late afternoon. This activity pattern is similar to
that observed in many tropical and temperate teiid lizards (Fitch
Fore-leg length (mm) 1973; Hirth 1963; Vitt and Carvalho 1992) as well as many
other lizard taxa occurring in open habitats (Pianka 1986). Even
NOTE: Means are presented f 1 SE with min./max. values in parentheses though substratum temperatures in the shade remain below the
underneath. The minimum value for SVL indicates size at sexual maturity,
the maximum indicates maximum SVL for each sex.
body temperatures of C. deppii during the activity periods,
substratum temperatures in the sun exceed 50°C before mid-
day, not retreating below 50°C until approximately 17:OO.
Presumably high microenvironmental temperatures partially
Discussion explain the reduced activity of these lizards at midday. Activity
Typically, Cnemidophorus species are well known for their at high body temperatures is not uncommon in tropical teiid liz-
high body temperatures (Anderson and Karasov 1981; Karasov ards (e.g., L.J. Vitt and G.R. Colli, unpublished data; Fitch
and Anderson 1984; Bowker 1993; Schall 1977) and activity 1973; Casas-Andreu and Gurrola-Hidalgo 1993).
levels (Garland 1993; Anderson 1993). These characteristics Mean active body temperatures for this population of C. deppii
seem tied in with a set of ecological (Huey and Pianka 1981; are higher than those reported for most other lizard species
Magnusson et al. 1985; Anderson and Karasov 1988), physio- (Brattstrom 1965; Huey and Slatkin 1976; Schall 1977) but are
logical (Anderson and Karasov 1981; Karasov and Anderson similar to those reported for a Costa Rican beach population
1984), and behavioral (Regal 1978, 1983; Garland 1993; Vitt of C. deppii (Fitch 1973) and some desert lizards (Anderson
and Cooper 1986) attributes of active foraging lizards. and Karasov 1981; Karasov and Anderson 1984). Several obser-
The tropical teiid lizard C. deppii inhabits a variety of habitats vations suggest that C. deppii prefer activity at a high body
associated with the dry forests of western Central America. It temperature. Microhabitat temperatures become extreme before
does not occur on beaches associated with the tropical moist 10:00, exceeding 40°C on sand surfaces in the sun. Cnemido-
forests of the Atlantic coast (e.g., Wright 1993). During the phorus deppii can and do spend considerable time in the shade
tropical dry season, Pacific coast habitats are xeric and can be or in filtered sun, presumably to reduce exposure to high sub-
considered stressful for lizards in terms of water balance. stratum temperatures in the sun. They frequently press them-
Because the beach sand is quick to drain after rainfall, there selves against the substratum in the shade, presumably to lose
is no surface fresh water even following brief rains in the dry heat to the substratum. The substratum in the shade remains
 CAN. 1. ZOOL. VOL. 7 1. 1993

TABLE5. Analyses of variance on residuals of regressions of log,, and of specified morphological characters versus logs
of snout-vent length between sexes of Cnemidophorus deppii

Morphological
character ANOVA on residuals Intercept Slope f SE R2 Regression F test
Can. J. Zool. Downloaded from www.nrcresearchpress.com by "Institute of Vertebrate Paleontology and Paleoanthropology,CAS" on 06/10/13

Mass (g)
Both sexes
Head width (mm)
Males*
Females
Head length (mm)
Males*
Females
Head height (mm)
Males*
Females
Body width (mm)
Both sexes
Body height (mm)
Both sexes
Hind-leg length (mm)
Males*
Females
Front-leg length (mm)
Both sexes
--
For personal use only.

- - -

NOTE:Where sexual differences exist (indicated by p > 0.05 in ANOVA on residuals), separate regression models are presented, otherwise models common
to both sexes appear. Asterisks indicate which sex is larger with respect to the given character. Note that the data set here is larger than that in Table 4 owing
to the inclusion of immature animals.

below 35°C until about 15:00. Our focal animal studies show detected by visual cues similar to those used by other lizard
that individuals spend much more time in the shade as environ- species (Cooper 1981). Cnemidophorus deppii did not, however,
mental temperatures increase. Nevertheless, even though micro- take advantage of the plethora of marine invertebrates, partic-
habitats with lower temperatures are available, the lizards ularly crustaceans, that were abundant on the beach.
maintain their high body temperatures. In addition, during The patchy distribution of prey (i-e., most prey are associated
cloudy periods there is the suggestion that substratum temper- with vegetation) results in lizards moving across the sand in
atures in the sun may decrease because of lack of exposure. the sun to search for prey among patches. This is reflected in
Some of the variation in surface sun temperatures (Fig. 5) the high rates of movement as well as the large distances moved
between 12:OO and 16:OO resulted from clouds blocking the during the activity sampling periods. The high rates and dis-
sun. Finally, the amount of time in a given day that C. deppii tances moved appear typical for whiptail lizards. It has been
maintain high body temperatures may be low (see below). suggested that the costs of this activity are most likely accounted
Yet another observation adds credence to the notion that for by high foraging efficiency (Etheridge and Wit 1993). Fitch
C. deppii prefer activity at very high body temperatures. We (1973) reported distances moved by individual C. deppii in
found no differences in body temperatures between lizards Costa Rica ranging from 0 to 170 m, but 76 of 81 moved less
active in the sun versus those found active in the shade, as long than 60 m, and most moved less than 24 m. His estimates
as sun was available. However, during brief cloudy periods, differ from ours partly as a result of differences in how move-
body temperatures of C. deppii were significantly lower than ment was measured. Our estimates were based on timed inter-
body temperatures of lizards active while sun was available. vals (600 s) for individual lizards during a single daily activity
We suspect that the reduced body temperatures under cloudy period. Fitch's movement estimates were based on linear dis-
conditions reflect the inability of C. deppii to gain the heat neces- tances between where individuals were successively captured
sary to maintain high body temperatures in the absence of the on different days. Nevertheless, the distances that C. deppii
sun. Similar effects of clouds on body temperatures of active move in a single day, or over longer time periods, are large
teiid lizards have been reported by Vitt and Carvalho (1992). compared with the distances moved by typical similarly sized
The diet of C. deppii is similar to that reported for other spe- sit-and-wait foraging lizards. The daily duration of activity for
cies of Cnemidophorus (e.g ., Mitchell 1979; Pianka 1970; Vitt individual C. deppii remains unknown. However, individual
1991; Schall 1993). Most species of Cnemidophorus tend to activity may actually be restricted to a relatively short period
eat more termites, spiders, and orthopterans'than other prey of time. The number of hours per day that other species of
types. Many of these prey are not surface active at the same Cnemidophorus are active varies from 2 to 5 (Etheridge and
time as the lizards. The lizards detect prey by chemical means Wit 1993). In addition, because these lizards ingest many hidden
(Cooper 1990) and dig them from the ground or locate them prey (see Vitt and Cooper 1986), considerable searching is
under surface items. Other prey, particularly orthopterans, are required to locate them. Consequently, a major cost of forag-
 VITT ET AL. 2399

ing in this thermally extreme environment is activity at very Acknowledgments

high body temperatures. We thank Milton Camacho of the Departamento de Fauna
The reproductive characteristics of C. deppii are similar to Silvestre of Nicaragua for making this research possible and
those of other species of Cnemidophorus with the same body for providing excellent logistic aid. Magdiel Gonzales of the
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size (Dunham et al. 1988; Vitt and Breitenbach 1993). In a Departamento de Fauna Silvestre aided in the field studies, and
study of C. deppii in Costa Rica, Fitch (1973) reported reproduc- to him we extend our appreciation. Financial aid for this and
tive characteristics nearly identical to those reported here. other Nicaraguan studies stemmed from a research grant from
Females in the Costa Rican population reached sexual maturity the Faculty Research Council at the University of Oklahoma
at 58 mm SVL, averaged 70 mm SVL, produced clutches to L.J.V. and J.P.C.
averaging 2.8 eggs, and females reproduced at least 2 times
per year. Most of the differences between the Costa Rican Abacus Concepts. 1987. Statview 11. Abacus Concepts Inc., Berkeley,
population and that in Nicaragua can be attributed to differ- California.
ences in mean adult female size and their effects on reproduc- Anderson, R.A. 1993. An analysis of foraging in the lizard, Cnemido-
tive characteristics. phorus tigris. In Biology of whiptail lizards (Genus Cnemidophorus).
Some females in our Nicaraguan samples contained oviductal Edited by J.W. Wright and L.J. Vitt. Oklahoma Museum of Natural
eggs or corpora lutea, indicating that at least some reproduc- History, Norman, Oklahoma. pp. 83 - 116.
Anderson, R.A., and Karasov, W.H. 1981. Contrasts in energy
tion was taking place late in the dry season. Most tropical
intake and expenditure in sit-and-wait and widely foraging lizards.
Cnemidophorus that have been studied either curtail (e.g., Leon Oecologia, 49: 67 -72.
and Cova 1973) or reduce (Vitt 1983; Magnusson et al. 1985) Anderson, R.A., and Karasov, W .H., 1988. Energetics of the lizard
reproduction during the dry season, with all or most reproduc- Cnemidophorus tigris and life history consequences of food-
tion occurring during the wet season. Fitch (1985) recognized acquisition mode. Ecol. Monogr. 58: 79- 110.
that the "long and severe dry season [in southwest Nicaragua Anderson, R.A., and Vitt, L.J. 1990. Sexual selection versus alterna-
and northwest Costa Rica] inhibits egg production," and he tive causes of sexual dimorphism in teiid lizards. Oecologia, 84:
went on to state, "Based on our limited data from SW 145- 157.
Nicaragua, it is clear that some reproduction does take place Bowker, R.C. 1993. The thermoregulation of the lizards Cnemido-
during the dry season." Consequently, Fitch's observations phorus exsanguis and C. velox: some consequences of high body
temperature. In Biology of whiptail lizards (Genus Cnemidophorus).
For personal use only.

correspond directly with ours.

Edited by J.W. Wright and L.J. Vitt. Oklahoma Museum of
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