Bergmann's Rule predicts larger body sizes in colder habitats, increasing organisms' ability to conserve heat. Originally formulated for endotherms, it is controversial whether Bergmann's Rule may be applicable to ectotherms, given that larger ectotherms show diminished capacity for heating up. We predict that Bergmann's Rule will be applicable to ectotherms when the benefits of a higher conservation of heat due to a larger body size overcompensate for decreased capacity to heating up. We test this hypothesis in the lizard Psammodromus algirus, which shows increased body size with elevation in Sierra Nevada (SE Spain). We measured heating and cooling rates of lizards from different elevations (from 300 to 2500 m above sea level) under controlled conditions. We found no significant differences in the heating rate along an elevational gradient. However, the cooling rate diminished with elevation and body size: highland lizards, with larger masses, have a higher thermal inertia for cooling, which allows them to maintain heat for more time and keep a high body temperature despite the lower thermal availability. Consequently, the net gaining of heat increased with elevation and body size. This study highlights that the heat conservation mechanism for explaining Bergmann's Rule works and is applicable to ectotherms, depending on the thermal benefits and costs associated with larger body sizes.
Altitudinal gradients offer a good opportunity to study organisms' adaptations to clinal environmental variables. Regarding altitude, the most influential variables on organisms are temperature and ultraviolet (UV) solar radiation, the first decreasing and the second increasing with altitude. Both variables affect ectotherms' biology, as ectotherms depend on environmental temperature for thermoregulation, frequently being heliotherms. Here, we studied dorsal coloration in the lizard Psammodromus algirus (Linnaeus, 1758) along a wide altitudinal gradient (2200 m) in Sierra Nevada (south-east Spain). We hypothesize that the skin will be darker with altitude, i.e. in environments with lower temperatures and higher UV radiation intensity. Results show that individual dorsal colorations became darker at high altitude. We propose two non-mutually exclusive explanations for this result: (1) darker dorsal surface would favour faster warming at high altitudes, where temperature is lower, and (2) darker dorsal surface would protect against UV radiation, stronger at high altitudes. We found significant relationships between both temperature and UV radiation and population dorsal darkness, giving mixed support for the two explanations. Moreover, dorsal hue was positively correlated with substrate hue, suggesting that hue evolved to maximize crypsis. Our study therefore suggests that geographical variation in dorsal coloration in this lizard is adaptive, and darkness coloration might have evolved in response to adverse conditions (low temperature and high UV radiation) at high altitudes.
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