Serpientes

THERMAL ECOLOGY OF BLACK RAT SNAKES (ELAPHE OBSOLETA) IN A THERMALLY CHALLENGING ENVIRONMENT
GABRIEL BLOUIN-DEMERS1
AND

PATRICK J. WEATHERHEAD2

Department of Biology, Carleton University, 1125 Colonel By Drive, Ottawa, Ontario, Canada K1S 5B6

Abstract. A general model in thermal ecology predicts that ectotherms should stop thermoregulating when the costs outweigh the benefits. Supportfor this model comes from studies of warm-temperate species, but the extent to which the model can be extrapolated to species living in climatic extremes is unknown, because of the lack of information regarding the thermoregulatory behavior of such species. We tested the applicability of this cost–benefit model using data for black rat snakes (Elaphe obsoleta obsoleta) studied at the northernextreme of their range in Ontario. During 1997–1999, we used automated temperature-sensitive radiotelemetry to collect 150 000 body temperatures from 53 freeranging rat snakes. Simultaneously, we used physical models of snakes to measure the environmental operative temperatures available to black rat snakes, and we determined their preferred body temperature range in a laboratory thermal gradient.The mostly forested habitats inhabited by rat snakes in Ontario were more thermally challenging than the habitats of other species studied to date. The preferred body temperature ranges of male, nongravid female, and gravid female black rat snakes were not significantly different (preferred body temperature averaged across all individuals, 28.1 C). However, free-ranging gravid females tended tomaintain higher body temperatures in order to thermoregulate more effectively as well as exploit their thermal environment more than males and nongravid females. This difference was most pronounced during the day and prior to egg laying, and constituted the first documentation of such a phenomenon in an oviparous snake. Black rat snakes had indices of thermoregulation effectiveness similar to otherspecies but tended to exploit opportunities for thermoregulation less. Overall, our data provided support that was at best ambiguous for the current cost–benefit thermoregulation model, suggesting that this model may generally be less applicable to species inhabiting climatic extremes. We propose that, for species in extreme climates, the costs associated with thermoconformity may be more importantthan previously recognized. We identified several problems associated with the index of thermoregulation effectiveness used by previous researchers, and we propose a mathematically simpler alternative that circumvents these problems. We also make recommendations regarding the future use of the various indices of thermoregulation developed in recent years.
Key words: black rat snake; cost–benefitthermoregulation; ectotherm; Elaphe obsoleta obsoleta; Ontario, Canada; reptile; thermal challenge; thermal exploitation; thermoconformer; thermoregulation; thermoconformity.

INTRODUCTION Some reptiles are precise thermoregulators and are only active under a narrow range of body temperatures (Tb) (Adolph 1990). Other reptiles are thermal conformers and are active under a broader range of Tb(Ruibal and Philibosian 1970, Moore 1978, Hertz 1992). Differences between species in the costs and benefits of thermoregulation are assumed to account for this variation (Huey and Slatkin 1976). The most
Manuscript received 24 August 2000; revised 15 December 2000; accepted 23 December 2000; final version received 23 January 2001. 1 Present address: Department of Evolution, Ecology, and OrganismalBiology, Botany and Zoology Building, Ohio State University, 1735 Neil Avenue, Columbus, Ohio 432101293 USA. E-mail: gblouind@ccs.carleton.ca 2 Present address: Department of Natural Resources and Environmental Sciences, University of Illinois, 1102 South Goodwin Avenue, Urbana, Illinois 61801 USA.

obvious cost is time, because time spent thermoregulating, or waiting for conditions that allow...