Behavioral Thermoregulation in Australian Elapid Snakes

Author(s): Harvey B. Lillywhite

Source: Copeia, Vol. 1980, No. 3 (Sep. 6, 1980), pp. 452-458
Published by: American Society of Ichthyologists and Herpetologists
Stable URL: http://www.jstor.org/stable/1444521
Accessed: 29/06/2010 15:15

Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at
http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless
you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you
may use content in the JSTOR archive only for your personal, non-commercial use.

Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at
http://www.jstor.org/action/showPublisher?publisherCode=asih.

Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed
page of such transmission.

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.

American Society of Ichthyologists and Herpetologists is collaborating with JSTOR to digitize, preserve and
extend access to Copeia.

http://www.jstor.org
Copeia, 1980(3), pp. 452-458

Behavioral Thermoregulation in Australian Elapid Snakes

HARVEY B. LILLYWHITE

Behavioral thermoregulation in laboratory thermal gradients was studied in

seven species of Australian snakes of the Elapidae:Acanthophisantarcticus,Aus-
trelaps superbus, Notechis scutatus, Pseudechis porphyriacus, Pseudonaja nuchalis,
P. textilis and Unechisflagellum.All species exhibited well-developed thermoreg-
ulatory behavior and controlled body temperatureswith precision comparableto
that reported for various heliothermic lizards. Temperatureregulation is accom-
plished by shuttling and by adjustments in the snake's position or orientation
while basking. Thermal preferenda of the various species range roughly between
30 and 35 C and are similar to those of other terrestrial, Temperate Zone snakes
in which thermal preferences have been adequatelyassessed. Thermal preferenda
of adult snakes appear to be higher than those of newborn or juveniles (in four
species) and may vary with levels of critical thermal minima (in five species) and
geographic distribution.

studies have demonstrated er taxa of snakes, I sought to determine the

NUMEROUS
the abilities of reptiles to maintain a char-
acteristic range of body temperatures by pos-
tural adjustments and the selection of appro-
priate microenvironments (reviews by Cloud-
sley-Thompson, 1971; Templeton, 1970;
Brattstrom, 1965). In laboratory studies, ther-
mal preferenda (Licht et al., 1966) may be de-
scribed from analysis of behavioral responses
to conditions which allow the opportunity for
temperature regulation. These procedures are
valuable in ascertaining the capabilities of rep-
tiles for behavioral thermoregulation and in
studying the levels and plasticity of thermal
preferenda in relation to phylogenetic, ecolog-
ical or physiological variables. Inherent thermal
preferences may be important in limiting mi-
crohabitat distribution and the timing of activ-
ity (Huey and Slatkin, 1976), and may also un-
derlie a suite of physiological adaptations to
temperature (Dawson, 1975).
Although there exist numerous data on the
thermoregulatory behavior of lizards, our
knowledge of this subject in relation to other
reptilian taxa is sparse and fragmentary. There
has been strong interest in snake thermoregu-
lation, but most studies have been limited to
either of two families, the Colubridae and Boi-
dae (Hammerson, 1977, 1979; Johnson, 1973;
Dill, 1972; Webb and Heatwole, 1971; Osgood,
1970; Kitchell, 1969; Hutchison et al., 1966;
Stewart, 1965; Cogger and Holmes, 1960). In
the interest of extending our knowledge to oth-
thermoregulatory responses of seven species of
Australian elapid snakes in laboratory thermal
gradients.
MATERIALS AND METHODS
The snakes used in this study (Table 1) were
collected in Victoria, South Australia and New
South Wales, and were returned directly to the
laboratory at Monash University. The snakes
were maintained at 24-26 C in metal or wood
cages with continuous access to water. Most
snakes were tested in thermal gradients within
one week of capture. Some of the snakes were
held for 2-3 weeks and were fed mice during
this period. However, these snakes were fasted
for a minimum of four days prior to testing.
All of the snakes selected for testing presum-
ably were post-absorptive, appeared to be in a
vigorous state of health and were not about to
engage in skin-shedding.
Two enclosures were used for thermal gra-
dients. Larger snakes (>60 cm total length)
were tested in a metal cage 180 x 90 x 30 cm
having a thick metal mesh top. Smaller snakes
were tested in a wood cage 100 x 46 x 45 cm
having a glass front and a screen mesh top. The
floor of each cage was covered with dry soil
overlain by a thin layer of dried grass clippings.
Thermal gradients were established by posi-
tioning a reflecting infrared heat lamp (250
watt red bulb) 30-45 cm above the substrate at

? 1980 by the American Society of Ichthyologists and Herpetologists

 LILLYWHITE-SNAKE THERMOREGULATION 453

40o- Two types of transmitter were used. A short

range transmitter consisted of a squegging
o
-
Hartley oscillator with an operating frequency
L35 of approximately 1 mHz on the AM broadcast
band. A longer range, pulsed, crystal controlled
citizen's band transmitter was sometimes used
•- 30 NOTECHISSCUTATUS with an operating frequency of 27.24 mHz.
w -
PSEUDECHIS Both types of transmitter emitted pulse rates
PORPHYRIACUS
- PSEUDONAJA TEXTILIS dependent on temperature and were calibrated
25 I I I I by use of a mercury thermometer and a con-
TIME,HOURINTERVALS trolled temperature bath. Each transmitter was
coated with silicone before calibration and
Fig. 1. Telemetered body temperaturesdepicting
thermoregulatory behaviorof three different snakes. feeding (forcibly) to a snake. The size of each
Examples were selected to illustrate variabilityand transmitter with silicone was approximately 3
speciesdifferences. Note the early warm-up(by bask- x 1.5 cm.
ing) in all three individuals. Each snake equipped with its transmitter was
tested individually. A snake was introduced into
one end of each cage. The lowest temperature the thermal gradient usually about 0700 h and
in the gradient was determined by ambient observed at intervals thereafter. The snake's
room air which usually ranged between 18 and body temperature and position in the thermal
22 C. Body temperatures in excess of 40 C were gradient were determined every 20 min (but
possible if a snake was positioned directly be- sometimes shorter or longer intervals: see be-
neath the heat lamp. low) throughout the rest of the work day (end-
Body temperatures of larger snakes were de- ing at 1700-1800 h). Some snakes were ob-
termined at intervals from a temperature trans- served (similarly) for 2-5 days.
mitter positioned freely in the gut at mid-body. Snakes smaller than 60 cm were tested in

TABLE1. BODYTEMPERATURES
OF SNAKESIN LABORATORY
THERMALGRADIENTS.
The mean and range de-
scribe all temperaturesmeasured after each snake had first warmed (by basking)to its preferred or activity
range of temperatures. Voluntary maxima are indicated as a single temperature or as a range of single
temperaturesfrom different snakes.
No.
No. measure-
Species animals ments X + SD Range Voluntary maxima Method

Acanthophisantarcticus 1 72 32.9 ? 3.64 27.2-38.5 38.5 telemetry

Austrelapssuperbus
juveniles 2 38 31.0 ? 2.37 25.8-35.3 34.8-35.3 thermometer
adults 4 78 32.8 ? 2.12 26.6-37.1 34.6-37.1 telemetry
Notechis scutatus
newborn 8 69 29.6 ? 2.66 23.0-34.0 34.0 thermometer
adults 7 170 32.2 ? 1.58 25.9-35.5 32.2-35.5 telemetry
Pseudechisporphyriacus
newborn 7 61 31.0 ? 1.90 27.0-35.0 35.0 thermometer
adults 3 60 32.7 ? 1.70 28.1-36.3 33.7-36.3 telemetry
Pseudonaja nuchalis
juvenile 1 11 32.0 ? 1.01 30.2-33.8 33.8 thermometer
adult 1 23 34.0 ? 1.21 31.8-36.9 36.9 telemetry
Pseudonaja textilis 2 36 34.5 ? 1.81 28.7-37.8 36.5-37.8 telemetry
Unechisflagellum 3 51 30.1 ? 2.87 24.0-36.2 34.2-36.2 thermometer
454 COPEIA, 1980, NO. 3

TABLE 2. VARIABILITY OF THERMOREGULATION TABLE 3. CRITICAL THERMAL MINIMA DETERMINED

AMONG INDIVIDUAL ADULT TIGER SNAKES, Notechisscu- IN FIVE SPECIES OF ELAPID SNAKES.
tatus.Temperatureswere determined as in Table 1.
Critical thermal
Snake minimum, C
Body temperature,C
Indi- Body
vidual mass,g + SD Range Austrelapssuperbus A 1.0
B 0.1
A* 380 33.0 ? 1.35 30.0-34.6 C 1.8 a = 0.97
B* 300 31.5 ? 1.35 29.6-34.3
C 285 32.4 ? 1.20 29.7-34.3 Notechis scutatus A 2.0
D 200 30.7 ? 3.30 25.9-35.0 B 1.0
E 300 32.8 ? 1.36 30.2-35.0 C 0.1
F 296 30.4 ? 0.33 30.0-31.2 D 0.5
G 314 29.9 ? 4.16 23.8-35.5 E 1.0 f = 0.92
* Gravid female. Pseudechisporphyriacus A 3.0
B 3.0 I = 3.00
Pseudonaja nuchalis A 7.0
t = 7.25
thermal gradients without the use of transmit- Pseudonaja textilis A 7.5 = 7.25
ters. In these cases, the protocol was similar to
that described for the larger snakes except that
body temperatures were taken by a Schultheis spent near the heat source. All of the species
thermometer inserted (orally) a few cm into the exhibited thermal preferenda at or somewhat
neck. Because of the disturbance to snakes, above 30 C (Table 1), although a broader range
body temperatures were taken every hour or at of body temperatures was tolerated. With one
longer intervals (1.5-3 h), and the period of exception (see below), thermal preferenda of
measurements was protracted for several days. individual snakes differed little on different
The smallest snakes (less than 30 cm total days, and there were no consistent trends in the
length) were tested in groups of 3-8 individu- daily differences. Control of body temperature
als. was very precise in some individuals (F, Table
For purposes of correlating preferred tem- 2). Smaller snakes appeared to maintain lower
peratures with thermal tolerance, the critical body temperatures than did larger snakes (Ta-
thermal minimum (CTMin) was determined in ble 1), but these differences were not tested sta-
twelve snakes. In these tests, a snake was cooled tistically because the smaller snakes were un-
in chipped ice at a rate of about 1 C min-1 until marked in groups and the variance between or
immobilized. The criteria used to determine within individuals could not be distinguished.
each snake's CTMin were 1) inability of the an- Differences in thermal preferenda were cor-
imal to right itself or to twist the body when related with differences in voluntary maxima
turned ventral side up, and 2) the cessation of (r = 0.577, P < 0.01, N = 23 observation pairs)
tongue flicks. The test was ended only when and with levels of CTMin (Tables 1, 3).
both of these conditions occurred. Tests for During basking, snakes positioned their bod-
CTMin were conducted on snakes that had ies in configurations which varied from out-
been held in the laboratory (24-26 C) for at stretched to tightly coiled. Adult specimens of
least two weeks. Notechisscutatus,Pseudechisporphyriacusand Aus-
trelapssuperbussometimes were observed at 5 or
RESULTS 10 min intervals and were seen to adjust bask-
ing positions so as to vary the part of the snake
All of the snakes behaviorally regulated body which received the most heat (Fig. 2). It did not
temperature in the thermal gradients. Typical- appear that (over time) any particular part of
ly, each snake elevated its body temperature by the body was warmed preferentially, and usu-
warming itself (basking) beneath the heat ally all segments of the body length were posi-
source and then maintained a relatively narrow tioned to receive maximum heating at different
range of elevated body temperatures during times. Coil adjustments also were observed in
subsequent activity within the gradient (Fig. 1). the other species, but their temporal aspects
Periods of movement generally included peri- were not noted in detail. Newborn snakes fre-
odic sojourns to the warmer end of the thermal quently formed aggregations near the heat
gradient, and periods of inactivity usually were source.
 LILLYWHITE-SNAKE THERMOREGULATION 455

and is comparable in pattern and precision to

that reported for various heliothermic lizards.
Thermal preferenda of the seven species range
roughly between 30 and 35 C and are (gener-
ally) similar to preferred temperatures report-
0910 0920 0930 ed for other terrestrial, Temperate Zone snakes
27.1 31.2 32.2
in which thermal preferences have been stud-
Fig. 2. Sequence of body positions at ten minute ied adequately (Hammerson 1977, 1979;
intervalsof an adult red-belliedblacksnake,Pseudechis Moore, 1978; Jacobson and Whitford, 1971;
porphyriacus,baskingbeneath an infrared heat lamp. Platt, 1969). There is considerable convergence
The x represents the center of irradiancefrom the of
heat lamp, and the circle indicates roughly the iso- thermoregulation between Australian brown
and the colubrid Masticophis
therm where irradiationis reduced to 50% of maxi- snakes (Pseudonaja)
mum. Numbers indicate time and core temperature studied by Hammerson (1977, 1979). Both gen-
of the snake. era are characterized by relatively high thermal
preferenda, similar behavior in a thermal gra-
dient and generally comparable precision and
consistency of body temperature control. These
The single specimen of Acanthophisantarcticus snakes are taxonomically and geographically
was relatively sedentary, and its behavior dif- distant, but are similar ecologically in that both
fered from that of the other species. This snake are swiftly-moving diurnal animals which feed
usually remained in a stationary coil near the on small rodents and other reptiles sometimes
heat source, but occasionally changed the po- in xeric habitats. Hence, this example suggests
sition of its body as described above for N. scu- that predictable adaptive patterns of thermo-
tatus, P. porphyriacusand A. superbus. I did not regulatory behavior have evolved among
observe the nocturnal behavior of this snake, snakes.
although on two occasions the heat lamp was The nocturnal snakes A. antarcticus and U.
left on overnight. Both times the snake was flagellum regulated body temperature in pho-
found near the heat lamp on the following tothermal gradients but with less consistency
morning and had a body temperature compa- than was observed in the other species. This
rable to that which it maintained during the difference possibly was evoked by the testing
daylight observation periods. On one occasion procedure which was not entirely appropriate
the heat lamp was positioned directly over the for nocturnal or secretive snakes.
snake which allowed its body temperature to There is very little information on the body
rise to 38.5 C before moving. Conversely, when temperatures of elapid snakes under natural
the snake was lifted away from the lamp (two conditions. Heatwole (1976; Heatwole and
occasions), the animal returned to occupy a po- Johnson, 1979) reported ranges of 25.0-33.8
sition nearer the heat source. Body tempera- and 27.2-33.2 C for naturally active N. scutatus
ture did not fall below 27.2 C during these ob- and P. porphyriacus,respectively, and these tem-
servations. Unlike individuals of other species, peratures correspond reasonably well with the
the mean preferred body temperature of A. range and preferenda of body temperatures
antarcticus varied considerably on five different determined in the laboratory (Table 1). Ham-
days (daily thermal preferenda = 36.3, 29.4, merson (1979) found that body temperatures
29.7, 33.6, 31.3 C). of Masticophis in thermal gradients were very
Unechisflagellum is a secretive species, and the similar to those obtained in outdoor enclosures
specimens studied remained concealed beneath under a variety of weather conditions. My data
a dense layer of dried grass clippings which for laboratory thermal preferenda differ only
covered the cage floor. These snakes clearly a little from independent assessments by Geof-
thermoregulated and usually were found near frey Witten (pers. comm.) who determined a
the heat source, but their movements and be- thermal preferendum of 33.8 C for P. porphy-
havior were not observed. riacus; those of four other diurnal elapid
species, including N. scutatus and A. superbus,
DISCUSSION were ca. 31 C.
The behaviors of snakes altered both the di-
Thermoregulatory behavior is well devel- rection of heat exchange (e.g., basking) and the
oped in all of the snakes which were studied rate at which body temperatures changed (pos-
456 COPEIA, 1980, NO. 3
ture or orientation). Coarse body temperature
control was achieved principally by shuttling
between the heat source and other parts of the
thermal gradient, while adjustments in posture
and orientation during basking permitted finer
control of body temperature and probably min-
imized horizontal temperature gradients within
the snake. The dynamics of orientation behav-
iors observed in basking snakes (Fig. 2) impli-
cate a role of peripheral thermal receptors (Ca-
banac and Hammel, 1971) in regulating heat
uptake over the whole body and provide the
inference that body temperatures were rela-
tively uniform. Heatwole and Johnson (1979)
observed flattening and tilting in cold red-bel-
lied blacksnakes, P. porphyriacus, basking in
sunlight. They concluded that posture is a ma-
jor influence on the rate of change of core tem-
perature and suggested that horizontal gradi-
ents are more important than are vertical ones
in influencing the heating and cooling of the
body. I did not observe pronounced flattening
by snakes, but the snakes I observed were never
as cold as those studied by Heatwole and John-
son. Several reports have emphasized the ap-
parent tendencies of certain snakes to selective-
ly heat different parts of the body and to
maintain thermal gradients within body tissue
(Hammerson, 1977; Johnson, 1973; Webb and
Heatwole, 1971; Regal, 1966; Benedict, 1932).
I did not have the capability to monitor regional
body temperatures from snakes during the
present investigations; however, the shifting
behavior of these snakes while basking (Fig. 2)
probably eliminated appreciable thermal gra-
dients within their bodies.
Little attention has been paid to possible
changes in thermal preferenda during the on-
togeny of reptiles. In the present study, mean
preferred body temperatures of newborn orju-
venile snakes were lower than those of adults
of the same species (Table 1). This possibly in-
dicates an ontogenetic difference in thermal
choice, but alternative interpretations come to
mind. First, it is possible that in the newborn of
N. scutatus and P. porphyriacusthe tendency to
aggregate interfered with thermoregulatory
behavior, although I did not observe anything
to indicate that thermoregulation was socially
inhibited as reported for some lizards (Regal,
1971). Second, body temperatures of small
snakes were taken at intervals with a thermom-
eter whereas body temperatures of larger
snakes were obtained from radio transmitters
positioned in the gut. If indeed head-body tem-
perature differences were maintained by these
snakes, these differences might be reflected by
thermometer placement. The Schultheis meth-
od perhaps comes closer to measuring head
temperatures than does the transmitter meth-
od. It is further conceivable that the mere pres-
ence of an object in the gut causes the heat
seeking behavior reported to occur in certain
snakes and other ectothermic vertebrates
(Lang, 1979; Witten and Heatwole, 1978; Re-
gal, 1966; Cogger, 1974; Gatten, 1974; Lilly-
white et al., 1973). Digestion in various other
snakes (Hammerson, 1979; Kitchell, 1969) and
crocodilians (Diefenbach, 1975) is not accom-
panied by any significant upward shift in the
mean body temperature when compared with
fasted animals. In those snakes and crocodilians
which show a thermophilic response following
feeding, the elevation of body temperature
above fasting levels is proportionate to the vol-
ume or mass of ingested food (Saint Girons,
1975; Lang, 1979). The size of the transmitter
used in the present study was relatively small
and did not noticeably distend the gut. More-
over, experiments with toads indicated that the
elevation in preferred body temperature fol-
lowing meal ingestion cannot be attributed sole-
ly to the mechanical stimulus of objects in the
gut (Lillywhite et al., 1973).
Pough and Busack (1978) reported that sub-
adult Spanish fringe-toed lizards, Acanthodacty-
lus erythrurus, maintained a significantly lower
mean body temperature (35.4 C) than adults
(38.3 C) in the field. The activity of subadults
is curtailed by high temperatures during three
months of summer while the activity of the
more thermophilic adults is curtailed only dur-
ing July and August. The possibility that young
snakes have a thermal preferendum lower than
that of adults is consistent with certain obser-
vations from ecological studies. Young colubrid
snakes may be active later in the year than are
adults of the same species (Brown and Parker,
1976) and probably are less active than the
adults during the active season (Lillywhite, in
press). Possibly during these periods small
snakes are operating at lower body tempera-
tures than those which characterize the active
adults. Unfortunately, very little information is
available concerning ecological differences be-
tween different age classes of elapid or other
snakes.
Laboratory thermal preferenda are generally
correlated with cold tolerance (as measured by
CTMin) in five species of elapid snakes for
 LILLYWHITE-SNAKE
which both measurements are available (Tables
1, 2). Hence, both species of Pseudonaja dem-
onstrated the highest CTMin and also selected
high body temperatures in thermal gradients.
Lower CTMin and selected temperatures char-
acterized N. scutatus and A. superbus,with P. por-
phyriacus being intermediate. The species hav-
ing the lower CTMin (better cold tolerance) are
restricted to a southerly distribution in Austra-
lia, whereas the other species range much far-
ther north. The two species of Pseudonaja both
occur in open, dry and sometimes hot environ-
ments. My data are corroborated crudely by
those of Heatwole (1976) who reported CTMin
of N. scutatus to be lower than that of P. textilis,
but that of A. superbus only slightly lower than
that given for Pseudonaja. These data indicate
that attributes of thermal physiology are ad-
justed in relation to levels of preferred body
temperatures or vice versa.
ACKNOWLEDGMENTS
I am indebted to A. K. Lee for the use of his HUEY,R. B., ANDM. SLATKIN.1976. Costs and ben-
laboratory and to the Department of Zoology,
Monash University, for support during my stay
as a visiting scientist. Special thanks are extend-
ed to Ron Waters and Roger Martin who pro-
vided some of the snakes.
LITERATURE CITED
BENEDICT,F. G. 1932. The physiology of large rep-
tiles with special reference to the heat production JOHNSON, C. R. 1973. Thermoregulation in py-
of snakes,tortoises,lizardsand alligators.Carnegie
Inst. Wash. Publ. No. 425, Washington,D.C.
BRATTSTROM, B. H. 1965. Body temperatures of
reptiles. Amer. Midl. Nat. 73:376-422.
BROWN,W. S., ANDW. S. PARKER.1976. Movement
ecology of Coluber constrictornear communal hiber-
nacula. Copeia 1976:225-242.
CABANAC, H. P., ANDH. T. HAMMEL.1971. Periph-
eral sensitivity and temperature regulation in Tili-
qua scincoides. Internat. J. Biometeor. 15:239-243.
CLOUDSLEY-THOMPSON, J. L. 1971. The temperature
and water relations of reptiles. Merrow Publ. Co.,
Watford Herts.
COGGER,H. G. 1974. Thermal relations of the mal-
lee dragon Amphibolurusfordi (Lacertilia: Agami-
dae). Aust. J. Zool. 22:319-339.
. 1975. Reptiles and amphibians of Australia.
A. H. & A. W. Reed, Sydney.
, AND A. HOLMES. 1960. Thermoregulatory
behavior in a specimen of Morelia spilotes variegata ~-
Gray (Serpentes:Boidae). Proc. Linn. Soc. New
South Wales 85:328-333.
DAWSON, W. R. 1975. On the physiological signifi-
THERMOREGULATION 457
cance of the preferred body temperaturesof rep-
tiles, p. 433-473. In: Perspectives of biophysical
ecology, ecological studies. Vol. 12. D. M. Gates and
R. B. Schmerl (eds.). Springer-Verlag, N.Y.
DILL, C. D. 1972. Reptilian core temperatures: vari-
ation within individuals. Copeia 1972:577-579.
DIEFENBACH, C. 0. 1975. Thermal preferences and
thermoregulation in Caiman crocodilus. Copeia
1975:530-540.
GATTEN,R. E., JR. 1974. Effect of nutritional status
on the preferred body temperature of the turtles
Pseudemys scripta and Terrapene ornata. Copeia
1974:912-917.
HAMMERSON, G. A. 1977. Head-body temperature
differences monitored by telemetry in the snake
Masticophisflagellum piceus. Comp. Biochem. Phys-
iol. 57A:399-402.
- . 1979. Thermal ecology of the striped racer,
Masticophis lateralis (Serpentes: Colubridae). Her-
petologica 35:267-273.
HEATWOLE, H. 1976. Reptile ecology. Univ. Queens-
land Press, St. Lucia.
- , AND C. R. JOHNSON. 1979. Thermoregula-
tion in the red-bellied blacksnake, Pseudechis por-
phyriacus (Elapidae). Zool. J. Linn. Soc. 65:83-101.
efits of lizard thermoregulation. Quart. Rev. Biol.
51:363-384.
HUTCHISON, V. H., H. G. DOWLING AND A. VINE-
GAR. 1966. Thermoregulation in a brooding fe-
male Indian python, Python molurus bivittatus. Sci-
ence 151:694-696.
JACOBSON, E. R., ANDW. G. WHITFORD.1971. Phys-
iological responses to temperature in the patch-
nosed snake, Salvadora hexalepis. Herpetologica
27:289-295.
thons---II. Headbody temperature differences and
thermal preferenda in Australian pythons. Comp.
Biochem. Physiol.45A:1065-1087.
KITCHELL, J. F. 1969. Thermophilic and thermo-
phobic responses of snakes in a thermal gradient.
Copeia 1969:189-191.
LANG, J. W. 1979. Thermophilic response of the
American alligator and the American crocodile to
feeding. Copeia 1979:48-59.
LICHT, P., W. R. DAWSON, V. H. SHOEMAKERAND A.
R. MAIN. 1966. Observations on the thermal re-
lations of Western Australian lizards. Copeia
1966:97-110.
LILLYWHITE, H. B. 1980. Tracking as an aid in the
study of snake communities. In: Herpetological
communities: A Symposium of the Society for the
Study of Reptiles and Amphibians and the Her-
petologist's League, August 1977. N. J. Scott (ed.).
Assoc. Syst. Curators, Lawrence, Kansas. In press.
, P. LICHTANDP. CHELGREN.1973. The role
of behavioral thermoregulation in the growth en-
ergetics of the toad, Bufo boreas. Ecology 54:375-
383.
458 COPEIA, 1980, NO. 3

MOORE, R. G. 1978. Seasonal and daily activity pat-
terns and thermoregulation in the southwestern
speckled rattlesnake (Crotalus mitchellipyrrhus) and
the Colorado desert sidewinder (Crotaluscerastesla-
terorepens)Copeia 1978:439-442.
OSGOOD, D. W. 1970. Thermoregulation in water
snakes studied by telemetry. Copeia 1970:568-571.
PLATT, D. R. 1969. Natural history of the hognose
snakes Heterodon platyrhinos and Heterodon nasicus.
Univ. Kans. Publ. Mus. Nat. Hist. 18:253-420.
POUGH,F. H., ANDS. D. BUSACK. 1978. Metabolism
and activity of the Spanish fringe-toed lizard (Lac-
ertidae: Acanthodactyluserythrurus).J. Thermal Biol.
3:203-205.
REGAL, P. J. 1966. Thermophilic response following
feeding in certain reptiles. Copeia 1966:588-590.
. 1971. Long term studies with operant con-
ditioning techniques, of temperature regulation
patterns in reptiles. J. Physiol. 63:403-406.
SAINTGIRONS, H. 1975. Observations preliminaires
sur la thermoregulation des viperes d'Europe. Vie
Milieu Ser. C, Biol. Terr. 25:137-168.
STEWART, G. R. 1965. Thermal ecology of the garter
snakes Thamnophissirtalis concinnus (Hallowell) and
Thamnophis ordinoides (Baird and Girard). Herpe-
tologica 21:81-102.
TEMPLETON, J. R. 1970. Reptiles, p. 167-221. In:
Comparative physiology of thermoregulation, vol.
1. G. C. Whittow (ed.). Academic Press, New York.
WEBB, G., ANDH. HEATWOLE. 1971. Patterns of heat
distribution within the bodies of some Australian
pythons. Copeia 1971:209-220.
WITTEN, G. J., AND H. HEATWOLE. 1978. Preferred
temperature of the agamid lizard Amphibolurusnob-
bi nobbi.Copeia 1978:362-364.
DEPARTMENT OF PHYSIOLOGY AND CELL BIOLO-
GY, UNIVERSITY OF KANSAS, LAWRENCE, KAN-
SAS 66045. Accepted 22 May 1979.

Copeia,1980(3),pp. 458-462

Orientation to the Sun by the Iguanid Lizards Uta stansburiana

and Sceloporusundulatus: Hourly and Monthly Variations

STEVE WALDSCHMIDT

Populations of Uta stansburiana and of Sceloporus undulatus were studied in

western Colorado. The distribution of the behavioral responses of these lizards
within four orientational categories showed significant variations throughout the
summer. These variations were closely associated with both monthly and hourly
changes in the thermal environment. During the day, the percentage of lizards
found in the shade was positively associated with increasing air temperature. The
observed frequencies of lizards in both parallel and perpendicular orientations
to the sun's rays suggest that these orientations are important behavioral re-
sponses to the thermal environment in the early morning and late afternoon.

ectotherm can regulate its body temper-
AN ature
by selecting different microhabitats,
altering its activity periods, and changing its
body surface areas that are exposed to thermal
fluxes. Many studies have shown that lizards are
capable of modifying their energy (heat) bal-
ance and thus behaviorally regulating their
body temperature [see reviews by Brattstrom
(1965) and Huey and Slatkin (1976)].
For two lizard species, I show that the relative
frequencies of four behavioral categories
change daily and monthly throughout the sum-
mer. I hypothesize that these behavior patterns
@ 1980 by the American Society of Ichthyologists and Herpetologists
are thermoregulatory, and are used to alter the
body surface area exposed to direct solar ra-
diation. If other factors that affect the energy
balance of a lizard are held constant, changes
in surface area will affect a lizard's body tem-
perature by altering the amount of direct solar
radiation absorbed (Bartlett and Gates, 1967;
Porter et al., 1973; Muth, 1977).
MATERIALS AND METHODS
Study site.-The 1,200 m2 study site was lo-
cated on a south facing hillside (1,658 m ele-