Accurate predictions of the potential distribution of range-shifting species are required for effective management of invasive species, and for assessments of the impact of climate change on native species. Range-shifting species pose a challenge for traditional correlative approaches to range prediction, often requiring the extrapolation of complex statistical associations into novel environmental space. Here we take an alternative approach that does not use species occurrence data, but instead captures the fundamental niche of a species by mechanistically linking key organismal traits with spatial data using biophysical models. We demonstrate this approach with a major invasive species, the cane toad Bufo marinus in Australia, assessing the direct climatic constraints on its ability to move, survive, and reproduce. We show that the current range can be explained by thermal constraints on the locomotor potential of the adult stage together with limitations on the availability of water for the larval stage. Our analysis provides a framework for biologically grounded predictions of the potential for cane toads to expand their range under current and future climate scenarios. More generally, by quantifying spatial variation in physiological constraints on an organism, trait-based approaches can be used to investigate the range-limits of any species. Assessments of spatial variation in the physiological constraints on an organism may also provide a mechanistic basis for forecasting the rate of range expansion and for understanding a species' potential to evolve at range-edges. Mechanistic approaches thus have broad application to process-based ecological and evolutionary models of range-shift.

During the month of February 1979, several hundred hatchling land iguanas (Conolophus pallidus) were observed emerging from their natal burrows in a 2 ha communal nesting area on Isla Santa Fe, Galapagos Islands. During this emergence, as many as nine Galapagos hawks were observed to patrol the nesting area and attack hatchling iguanas.The hypothesis that the ability of hatchling land iguanas to escape predation could be influenced by the interaction of the physiological state of the lizards and the thermal environment was analyzed using (1) empirical data on the effect of body temperature (T ) on locomotory ability of iguanas and (2) biophysical modeling of the T's of hatchlings under natural conditions. This hypothesis was tested by assessing the success of natural hawk attacks on lizards exposed to different thermal environments.During those periods when predicted T 's of hatchlings were always<32°C, (at which temperatures land iguanas were shown to have less than maximal ability to sprint rapidly) hawks were successful in 67% of the observed attacks. However, when T 's of hatchlings were always ≧32° C, hawks were successful on only 19% of observed attacks. During periods when hatchling T's could be <32° C or 32-40° C (depending upon which microhabitat the hatchling occupied before the attack), the hawks were successful in 46% of the observed attacks.These data indicate that the physical environment, as mediated through the physiological state of the lizards and to correlated locomotary abilities, significantly affects the ability of hatchling land iguanas to escape predation.

The aims of this paper are to compare the thermal ecology of four species of varanid lizards that occupy a range of habitats and climatic regions, and to assess the efficacy of methods for evaluating the extent to which ectothermic animals exploit their thermal environments. Hertz et al. (1993) have proposed several indices of thermoregulation, and these are evaluated with respect to our data from varanid lizards. The thermoregulatory characteristics of three tropical monitor lizards (Varanus panoptes, V. gouldii, and the semiaquatic V. mertensi), and the temperate—zone V. rosenbergi were studied throughout the year. Radiotelemetry was used to measure the body temperatures (Tbs) of free—ranging animals, and microclimatic data were collected to determine the range of possible Tbs that an animal could achieve. Operative temperatures (Tb's) were estimated by biophysical models for each set of animal characteristics and microclimatic conditions. The Tb's selected by animals in a laboratory thermal gradient were used to determine the set—point range of Tb's that the animals voluntarily select. Plots that superimpose Tb's, Tb's, and the set—point range across the day are extremely useful for describing the thermoregulatory characteristics of ectotherms. These plots can be used to determine the extent to which the animals exploit their thermal environment: we define an index of thermal exploitation (Ex) as the time in which Tb's are within the set—point range, divided by the time available for the animal to have its Tb's within the set—point range. Only V. mertensi was active throughout the year. In general, during seasons of inactivity, the Tb's of inactive species fell outside the set—point range, but during periods of activity all species selected Tb's within their set—point range. The temperate—zone species (V. rosenbergi) thermoregulates very carefully during periods when environmental conditions allow the animals to attain the set—point range, and V. gouldii also thermoregulates carefully in the wet season. V. mertensi selects Tb's that are significantly lower than the other species both in the field and in the laboratory, and thermoregulatory indices of this species were intermediate relative to the other species. The amount of time spent in locomotion each day was not correlated with the indices of thermoregulation: the most active species, V. panoptes, was, with respect to several indices, the least careful thermoregulator. The type of question that is being addressed, with respect to the interactions between an animal's thermal environment and its thermoregulatory behavior, determines the appropriateness of the various indices of thermoregulation. The Ex index describes the thermoregulatory characteristics of ecotherms in a heterogeneous thermal environment, and in such an environment a large amount of information can easily be interpreted graphically. This index is less useful in a thermally homogeneous environment.

Most frog species show little resistance to evaporative water loss (EWL), but some arboreal species are known to have very high resistances. We measured EWL and cutaneous resistance to evaporation (Rc) in 25 species of frogs from northern Australia, including 17 species in the family Hylidae, six species in the Myobatrachidae, and one each in the Bufonidae and the Microhylidae. These species display a variety of ecological habits, including aquatic, terrestrial, and arboreal specialisations, with the complete range of habits displayed within just the one hylid genus, Litoria. The 25 species measured in this study have resistances that range from Rc=0 to 63.1. These include low values indistinguishable from a free water surface to high values typical of "waterproof" anuran species. There was a strong correlation between ecological habit and Rc, even taking phylogenetic relationships into account; arboreal species had the highest resistance, aquatic species tended to have little or no resistance, and terrestrial species tended to have resistance between those of arboreal and aquatic frogs. For one species, Litoria rubella, we found no significant changes in EWL along a 1,500-km aridity gradient. This study represents the strongest evidence to date of a link between ecological habits and cutaneous resistance to water loss among species of frogs.

We used simulations from a biophysical model that integrates interlinked exchanges of energy and water between frogs and their environments to address questions about the limits to thermoregulation and about adaptations for arboreality. Body size and cutaneous resistance (Rc) both significantly affected body temperature (Tb) and the time to desiccate to 70% of standard mass (an ecologically relevant metric of desiccation). Cutaneous resistances < 25 s/cm allow basking frogs to elevate their Tb several degrees above ambient, but Rc above 25 had little additional effect on Tb. Small frogs (<10 g) are able to elevate their Tb above ambient while basking, even with small Rc. Large frogs must have greater skin resistances to be able to elevate body temperatures above ambient, yet large frogs take longer to desiccate to 70% of their standard mass. Frogs can avoid rapid desiccation with high Rc, a large body size, or some combination of these traits. Our literature survey indicates that frogs with a combination of Rc and body size that would result in long times to desiccate to 70% of standard mass tend to be arboreal, suggesting that those species may be selectively favored in a niche that often requires frogs to be away from water sources for extended periods of time.