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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
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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- thermoregulatory responses of seven species of
acteristic range of body temperatures by pos- Australian elapid snakes in laboratory thermal
tural adjustments and the selection of appro- gradients.
priate microenvironments (reviews by Cloud-
sley-Thompson, 1971; Templeton, 1970; MATERIALS AND METHODS
Brattstrom, 1965). In laboratory studies, ther-
mal preferenda (Licht et al., 1966) may be de- The snakes used in this study (Table 1) were
scribed from analysis of behavioral responses collected in Victoria, South Australia and New
to conditions which allow the opportunity for South Wales, and were returned directly to the
temperature regulation. These procedures are laboratory at Monash University. The snakes
valuable in ascertaining the capabilities of rep- were maintained at 24-26 C in metal or wood
tiles for behavioral thermoregulation and in cages with continuous access to water. Most
studying the levels and plasticity of thermal snakes were tested in thermal gradients within
preferenda in relation to phylogenetic, ecolog- one week of capture. Some of the snakes were
ical or physiological variables. Inherent thermal held for 2-3 weeks and were fed mice during
preferences may be important in limiting mi- this period. However, these snakes were fasted
crohabitat distribution and the timing of activ- for a minimum of four days prior to testing.
ity (Huey and Slatkin, 1976), and may also un- All of the snakes selected for testing presum-
derlie a suite of physiological adaptations to ably were post-absorptive, appeared to be in a
temperature (Dawson, 1975). vigorous state of health and were not about to
Although there exist numerous data on the engage in skin-shedding.
thermoregulatory behavior of lizards, our Two enclosures were used for thermal gra-
knowledge of this subject in relation to other dients. Larger snakes (>60 cm total length)
reptilian taxa is sparse and fragmentary. There were tested in a metal cage 180 x 90 x 30 cm
has been strong interest in snake thermoregu- having a thick metal mesh top. Smaller snakes
lation, but most studies have been limited to were tested in a wood cage 100 x 46 x 45 cm
either of two families, the Colubridae and Boi- having a glass front and a screen mesh top. The
dae (Hammerson, 1977, 1979; Johnson, 1973; floor of each cage was covered with dry soil
Dill, 1972; Webb and Heatwole, 1971; Osgood, overlain by a thin layer of dried grass clippings.
1970; Kitchell, 1969; Hutchison et al., 1966; Thermal gradients were established by posi-
Stewart, 1965; Cogger and Holmes, 1960). In tioning a reflecting infrared heat lamp (250
the interest of extending our knowledge to oth- 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 perature differences were maintained by these
control was achieved principally by shuttling snakes, these differences might be reflected by
between the heat source and other parts of the thermometer placement. The Schultheis meth-
thermal gradient, while adjustments in posture od perhaps comes closer to measuring head
and orientation during basking permitted finer temperatures than does the transmitter meth-
control of body temperature and probably min- od. It is further conceivable that the mere pres-
imized horizontal temperature gradients within ence of an object in the gut causes the heat
the snake. The dynamics of orientation behav- seeking behavior reported to occur in certain
iors observed in basking snakes (Fig. 2) impli- snakes and other ectothermic vertebrates
cate a role of peripheral thermal receptors (Ca- (Lang, 1979; Witten and Heatwole, 1978; Re-
banac and Hammel, 1971) in regulating heat gal, 1966; Cogger, 1974; Gatten, 1974; Lilly-
uptake over the whole body and provide the white et al., 1973). Digestion in various other
inference that body temperatures were rela- snakes (Hammerson, 1979; Kitchell, 1969) and
tively uniform. Heatwole and Johnson (1979) crocodilians (Diefenbach, 1975) is not accom-
observed flattening and tilting in cold red-bel- panied by any significant upward shift in the
lied blacksnakes, P. porphyriacus, basking in mean body temperature when compared with
sunlight. They concluded that posture is a ma- fasted animals. In those snakes and crocodilians
jor influence on the rate of change of core tem- which show a thermophilic response following
perature and suggested that horizontal gradi- feeding, the elevation of body temperature
ents are more important than are vertical ones above fasting levels is proportionate to the vol-
in influencing the heating and cooling of the ume or mass of ingested food (Saint Girons,
body. I did not observe pronounced flattening 1975; Lang, 1979). The size of the transmitter
by snakes, but the snakes I observed were never used in the present study was relatively small
as cold as those studied by Heatwole and John- and did not noticeably distend the gut. More-
son. Several reports have emphasized the ap- over, experiments with toads indicated that the
parent tendencies of certain snakes to selective- elevation in preferred body temperature fol-
ly heat different parts of the body and to lowing meal ingestion cannot be attributed sole-
maintain thermal gradients within body tissue ly to the mechanical stimulus of objects in the
(Hammerson, 1977; Johnson, 1973; Webb and gut (Lillywhite et al., 1973).
Heatwole, 1971; Regal, 1966; Benedict, 1932). Pough and Busack (1978) reported that sub-
I did not have the capability to monitor regional adult Spanish fringe-toed lizards, Acanthodacty-
body temperatures from snakes during the lus erythrurus, maintained a significantly lower
present investigations; however, the shifting mean body temperature (35.4 C) than adults
behavior of these snakes while basking (Fig. 2) (38.3 C) in the field. The activity of subadults
probably eliminated appreciable thermal gra- is curtailed by high temperatures during three
dients within their bodies. months of summer while the activity of the
Little attention has been paid to possible more thermophilic adults is curtailed only dur-
changes in thermal preferenda during the on- ing July and August. The possibility that young
togeny of reptiles. In the present study, mean snakes have a thermal preferendum lower than
preferred body temperatures of newborn orju- that of adults is consistent with certain obser-
venile snakes were lower than those of adults vations from ecological studies. Young colubrid
of the same species (Table 1). This possibly in- snakes may be active later in the year than are
dicates an ontogenetic difference in thermal adults of the same species (Brown and Parker,
choice, but alternative interpretations come to 1976) and probably are less active than the
mind. First, it is possible that in the newborn of adults during the active season (Lillywhite, in
N. scutatus and P. porphyriacusthe tendency to press). Possibly during these periods small
aggregate interfered with thermoregulatory snakes are operating at lower body tempera-
behavior, although I did not observe anything tures than those which characterize the active
to indicate that thermoregulation was socially adults. Unfortunately, very little information is
inhibited as reported for some lizards (Regal, available concerning ecological differences be-
1971). Second, body temperatures of small tween different age classes of elapid or other
snakes were taken at intervals with a thermom- snakes.
eter whereas body temperatures of larger Laboratory thermal preferenda are generally
snakes were obtained from radio transmitters correlated with cold tolerance (as measured by
positioned in the gut. If indeed head-body tem- CTMin) in five species of elapid snakes for
LILLYWHITE-SNAKE THERMOREGULATION 457

which both measurements are available (Tables cance of the preferred body temperaturesof rep-
1, 2). Hence, both species of Pseudonaja dem- tiles, p. 433-473. In: Perspectives of biophysical
onstrated the highest CTMin and also selected ecology, ecological studies. Vol. 12. D. M. Gates and
R. B. Schmerl (eds.). Springer-Verlag, N.Y.
high body temperatures in thermal gradients. DILL, C. D. 1972. Reptilian core temperatures: vari-
Lower CTMin and selected temperatures char- ation within individuals. Copeia 1972:577-579.
acterized N. scutatus and A. superbus,with P. por- DIEFENBACH, C. 0. 1975. Thermal preferences and
phyriacus being intermediate. The species hav- thermoregulation in Caiman crocodilus. Copeia
ing the lower CTMin (better cold tolerance) are 1975:530-540.
restricted to a southerly distribution in Austra- GATTEN,R. E., JR. 1974. Effect of nutritional status
lia, whereas the other species range much far- on the preferred body temperature of the turtles
ther north. The two species of Pseudonaja both Pseudemys scripta and Terrapene ornata. Copeia
occur in open, dry and sometimes hot environ- 1974:912-917.
ments. My data are corroborated crudely by HAMMERSON, G. A. 1977. Head-body temperature
differences monitored by telemetry in the snake
those of Heatwole (1976) who reported CTMin
of N. scutatus to be lower than that of P. textilis, Masticophisflagellum piceus. Comp. Biochem. Phys-
iol. 57A:399-402.
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temperatures or vice versa. land Press, St. Lucia.
- , AND C. R. JOHNSON. 1979. Thermoregula-
tion in the red-bellied blacksnake, Pseudechis por-
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SAINTGIRONS, H. 1975. Observations preliminaires

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