[April 1990 Asiatic Herpetological Research Vol. 3, pp.

54l9]

Thermal Sensitivity of Sprinting and Clinging Performance in the Tokay

Gecko (Gekko gecko)

JONATHAN B. LOSOS 1

1
Museum of Vertebrate Zoology and Department of Integrative Biology,
University of California, Berkeley, CA 94720 USA

Abstract. -The thermal sensitivity of sprinting and clinging ability was measured in tokay geckos
(Gekko gecko). Sprinting performance was maximal at high (35-4 1°C) temperatures, as is the case for
other nocturnal lizards, but the optimal temperature for clinging was considerably lower (approximately
17°C). These different optima could be adaptive if maximal sprinting and clinging capabilities are needed at
different temperatures. Alternatively, they could result from constraints on adaptive evolution.

Key Words: Reptilia, Sauria, Gekkonidae, Gecko gecko, thermal sensitivity, optimal performance.

Introduction frequently experience in nature (Bennett

1980); however, nocturnal lizards appear
Most physiological processes are exceptional (Huey and Bennett 1987; Huey
temperature-dependent. Ectothermic etal. 1989). The thermal sensitivity of few
animals, which do not maintain a constant other whole-organism locomotor
body temperature, are thus
subject to performance measures has been determined
fluctuation in the rate at which they can for lizards. Here I report the thermal
perform many vital tasks. By regulating sensitivity of tokay geckos (Gekko gecko)
body temperature behaviorally, however, [Fig. 1] for two ecologically relevant tasks,
many reptiles can maximize performance maximum sprinting and clinging capability.
(Huey 1982a). In some cases,
capability I ask whether maximum performance
maximal performance temperature might capability occurs at the relatively low
differ for different tasks.For example, a temperatures most commonly experienced
wealth of behavioral data indicates that by geckos (Huey et al. 1989) and whether
many lizards and snakes increase their body the optimal temperature is the same for the
temperature after feeding, which suggests two performance measures.
that digestion has a higher optimal
temperature than other activities (reviewed Methods
in Huey 1982a).
Tokays are relatively large geckos found
Early work centered on the thermal on trees and walls throughout southeastern
dependence of sub-organismal traits (e.g., Asia (Smith 1935). Lizards were captured
enzyme activity, muscle contractile speed on Phuket Island, Phuket Province,
[Huey and Stevenson 1979]). In many Thailand, and transported to the University
cases, however, the effect of temperature of California, Berkeley in late September
on the functional capacities (e.g., sprint 1987. They were maintained in 30 x 17 x 9
speed, critical thermal maximum cm plastic shoeboxes at ambient
temperature) cannot be predicted by study temperatures and provided with water and
at sub-organismal levels (e.g., Licht 1967; crickets ad libitum. An ontogenetic series
Marsh and Bennett 1985, 1986). was used in this study (11 individuals;
Consequently, recent studies have focused snout-vent length: 85-180 mm; mass: 10-
on whole-organism performance. The 140 g).
thermal dependence of sprinting ability has
been studied in great detail. Maximum All trials were conducted in a walk-in
sprinting speed and/or endurance in many environmental chamber in the Museum of
species occurs at the temperature they most Vertebrate Zoology, University of

1990 by Asiatic Herpetological Research

April 1990 Asiatic Herpetological Research Vol. 3, p. 55

FIG. 1 . Gekko gecko from Phuket Province, Thailand.

California, Berkeley, within a month of from the experimental plate.

capture. Lizards were placed in the chamber
at least one hour prior to performance One per lizard was conducted per
trial
measurement. Humidity, which could not temperature. Performance at nine
be regulated, was determined on several temperatures (12, 16, 17, 22, 24, 31 [three
occasions using a Bacharach sling times], 34, 35, 41; the order of
psychrometer. temperatures is presented in figure 2) was
measured over a six-day period. On some
Clinging capability was measured by days, two trials were conducted.
placing lizards on a plexiglass plate and, at
a gradual and steady rate, lifting the end of Sprint capability was measured by
the plate so that the lizard was clinging with placing lizards at the end of a 2.25 m
its head directed down. The angle at which trackway covered with a rough rubber
the lizard fell from the plate was recorded surface and inducing them to run by
(protocol modified from Emerson and Diehl repeated taps to the tail (protocol following
1980; Alberch 1981). Lizards that jumped Huey 1982b; Huey et al. 1989; Garland
from the plate were not included in the 1985). As
the lizard ran, it interrupted light
analysis. As lizards began to slide, they beams stationed every 0.25 m. The time
usually attempted to maintain their grip by elapsed during each interval was computed
moving and re-applying their toe pads by a Compaq personal computer, the fastest
regardless of temperature. There was no single interval during four trials, conducted
evidence that temperature affected the at hourly intervals, was considered the
lizards' efforts to prevent sliding and falling maximum speed for that lizard at that
Vol. 3, p. 56 Asiatic Herpetological Research April 1990

155 1.50

.25

1.00
20 30

Temperature
April 1990 Asiatic Herpetological Research Vol. 3, p. 57

Eremiascincus fasciolatus (Huey and evolve high critical thermal maxima. The
Bennett 1987). Clinging capability, the high sprint performance maximum might be
thermal dependence of which has never a correlated effect of this physiological
previously been investigated, is maximal at adaptation to high temperatures and not be
considerably lower (approx. 17°C) adaptive per se (Huey et al. 1989).
temperatures.
Clinging capability depends upon both
It is difficult to envision how such physical and physiological processes.
differentoptima could evolve adaptively. Geckos cling to smooth surfaces by dry
Most lizards sprint maximally at adhesion. The subdigital lamellar pads of
temperatures close to those they normally geckos are covered with millions of
experience (Huey 1982a; Huey et al. microscopic setal hairs. When the pads are
1989). The low optimal temperature for adpressed to a surface, these hairs form
clinging matches the field temperatures of intermolecular bonds with molecules on the
many active nocturnal geckos (Huey et al. surface of the substrate (Hiller 1975). If
1989). However, many nocturnal geckos the surface energy (a measure of the
bask and/or are active to some extent during number of free electrons on the surface of
the day (Bustard 1967, 1968; Werner and the substrate) is relatively high, then
Whitaker 1978; Nagy and Knight 1989). enough bonds can form to support the
Consequently, the high optimal temperature lizard. Because these bonds result from the
for sprinting seen in nocturnal lizards
many activityof electrons, the forces theoretically
might represent adaptation for diurnal should be temperature-independent over the
capability (Huey and Bennett 1987; Huey et range of temperatures in this study.
al.1989). Nonethless, it seems implausible
that one aspect of locomotion, sprint However, geckos have an elaborate
performance, should be selected at high muscular and vascular system for the
temperatures, whereas another important adpression and removal of their toe pads
component of effective movement, (Russell 1975); the thermal dependence of
clinging, should be favored at considerably these muscles has not been investigated.
lower temperatures. The poor clinging performance of geckos at
10-12°C is clearly the result of
As an alternative explanation, the physiological incapacitation. At that
differences in thermal optima might result temperature, geckos were generally
from differences in evolutionary lability of inactive, moved slowly and infrequently,
the two performance capabilities and thus and even rarely bit or barked when
represent constraints on adaptive evolution handled. In contrast to trials at higher
in either sprinting or clinging ability. To temperatures, the lizards did not attempt to
understand why these performance abilities adjust their pads or posture when the plate
are affected differently by temperature, a was tilted and quickly lost their hold. More
better understanding is needed of their research is required to determine whether
underlying physical and physiological the performance decline at temperatures
bases (e.g., Garland 1984, 1985; Marsh above 17-1 8°C results from decreased
and Bennett 1985, 1986). In many species capability of the muscles and enzymes
of lizards, sprint performance is maximal at involved in clinging.
temperatures close to the critical thermal
maximum (Huey and Bennett 1987; Huey One possible confounding effect in the
et al. 1989). The cause for this linkage is clinging experiments is the variation in
unclear. However, if geckos must be able absolute humidity. Moisture might
to survive diurnal temperatures (either decrease the formation of high-energy
because they intentionally maintain high intermolecular bonds. Absolute humidity
temperatures to maximize other processes, in the environmental chamber was
or because environmental conditions measured at several temperatures and
preclude the maintenance of lower increased from 8.5 Barrs at 11.9°C to 25.1
temperatures), then they would have to Barrs at 34.3°C (relative humidity,
Vol. 3, p. 58 Asiatic Herpetological Research April 1990

however, was greatest at intermediate EMERSON, S. B., AND D. DIEHL. 1980. Toe pad

temperatures). Consequently, the greater morphology and mechanisms of sticking in

absolute humidity at higher temperatures in frogs. Biological Journal of the Linnaen
the environmental chamber might have Society 13:199-216.
caused a decrease in clinging ability.
Further research is needed to investigate to GARLAND, T., JR. 1984. Physiological correlates
of locomotory performance in a lizard: an
what extent humidity affects clinging.
allometric approach. American Journal of
Physiology 247:R806-R815.
Although different processes may often
be maximized at different temperatures, GARLAND, T., JR. 1985. Ontogenetic and
rarely is the difference as great as is individual variation in size, shape and speed in
observed between sprinting and clinging in the Australian agamid lizard Amphibolurus
tokay geckos (Huey 1982a). One would nuchalis. Journal of Zoology, London (A)
not expect these geckos to need maximum 207:425-439.
ability at these aspects of locomotion at
such different temperatures, but neither HILLER, U. 1975.Comparative studies on the
would one expect the physiological functional morphology of two gekkonid lizards.
Journal of the Bombay Natural History Society
sensitivity of different systems to be so
73:278-282.
different. Interestingly, the optimal
temperature for hearing sensitivity in the
HUEY, R. B. 1982a. Temperature, physiology,
tokay gecko is intermediate between the and the ecology of reptiles. Pp. 25-91. In C.
sprinting and clinging optima (Werner Gans and F. H. Pough (Eds.), Biology of the
1976). Further research is required to
Reptilia, Vol. 12. Academic Press, New York,
understand the processes shaping and USA.
constraining performance evolution.
HUEY, R. B. 1982b. Phylogenetic and
Acknowledgments ontogenetic determinants of sprint performance
in some diurnal Kalahari lizards. Koedoe 25:43-

Ithank T. Papenfuss and R. Macey for 48.

providing the geckos, R. Jones for

assistance with equipment, R. Huey for HUEY, R. B., AND A. F. BENNETT. 1987.

advice on racing geckos, J. Herron for Phylogenetic studies of coadaptation: preferred

temperatures versus optimal performance
providing me with a pre-publication
temperatures of lizards. Evolution 41:1098-
manuscript, and A. Bauer, H. Greene and 1115.
R. Huey for constructive comments on a
previous version of this paper. HUEY, R. B., P. H. NIEWIAROWSKI, J.
KAUFMANN, AND J. C. HERRON. 1989.
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