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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Studying the causes of parasite geographic distribution is relevant to understand ecological and evolutionary processes that affect host populations as well as for species conservation. Temperature is one of the most important environmental variables affecting parasite distribution, as raising temperatures positively affect development, reproduction, and rate of transmission of both endo- and ectoparasites. In this context, it is generally accepted that, in mountains, parasite abundance decreases with elevation. However, empirical evidence on this topic is limited. In the present study, we analyzed the elevational variation of hemoparasites and ectoparasites of a lizard, Psammodromus algirus, along a 2,200-m elevational gradient in Sierra Nevada (SE Spain). As predicted, ectoparasite (mites, ticks, mosquitoes, and sandflies) abundance decreased with elevation. However, hemoparasite prevalence and intensity in the lizard augmented with altitude, showing a pattern contrary to their vectors (mites). We suggest that tolerance to hemoparasites may increase with elevation as a consequence of lizards at high altitudes taking advantage of increased body condition and food availability, and reduced oxidative stress. Moreover, lizards could have been selected for higher resistance against hemoparasites at lowlands (where higher rates of replication are expected), thus reducing hemoparasite prevalence and load. Our findings imply that, in a scenario of climate warming, populations of lizards at high elevation may face increased abundance of ectoparasites, accompanied with strong negative effects.

Achieving optimal body temperature maximizes animal fitness. Since ambient temperature may limit ectotherm thermal performance, it can be constrained in too cold or hot environments. In this sense, elevational gradients encompass contrasting thermal environments. In thermally pauperized elevations, ectotherms may either show adaptations or suboptimal body temperatures. Also, reproductive condition may affect thermal needs. Herein, we examined different thermal ecology and physiology capabilities of the lizard Psammodromus algirus along a 2200-m elevational gradient. We measured field (T(b)) and laboratory-preferred (T(pref)) body temperatures of lizards with different reproductive conditions, as well as ambient (T(a)) and copper-model operative temperature (T(e)), which we used to determine thermal quality of the habitat (d(e)), accuracy (d(b)), and effectiveness of thermoregulation (de-db) indexes. We detected no Tb trend in elevation, while T(a) constrained T(b) only at high elevations. Moreover, while Ta decreased more than 7 °C with elevation, T(pref) dropped only 0.6 °C, although significantly. Notably, low-elevation lizards faced excess temperature (T(e) > T(pref)). Notably, de was best at middle elevations, followed by high elevations, and poorest at low elevations. Nonetheless, regarding microhabitat, high-elevation de was more suitable in sun-exposed microhabitats, which may increase exposition to predators, and at midday, which may limit daily activity. As for gender, d(b) and d(e)-d(b) were better in females than in males. In conclusion, P. algirus seems capable to face a wide thermal range, which probably contributes to its extensive corology and makes it adaptable to climate changes.

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