Age structure of a lizard along an elevational gradient reveals nonlinear lifespan patterns with altitude.

Lifespan is one of the main components of life history. Shorter lifespans can be expected in marginal habitats. However, in the case of ectotherms, lifespan typically increases with altitude, even though temperature-one of the main factors to determine ectotherms' life history-declines with elevation. This pattern can be explained by the fact that a shorter activity time favors survival. In this study, we analyzed how lifespan and other life-history traits of the lizard Psammodromus algirus vary along a 2,200 m elevational gradient in Sierra Nevada (SE Spain). Populations at intermediate altitudes (1,200-1,700 m), corresponding to the optimal habitat for this species, had the shortest lifespans, whereas populations inhabiting marginal habitats (at both low and at high altitudes) lived longest. Therefore, this lizard did not follow the typical pattern of ectotherms, as it also lived longer at the lower limit of its distribution, nor did it show a longer lifespan in areas with optimal habitats. These results might be explained by a complex combination of different gradients along the mountain, namely that activity time decreases with altitude whereas food availability increases. This could explain why lifespan was maximum at both high (limited activity time) and low (limited food availability) altitudes, resulting in similar lifespans in areas with contrasting environmental conditions. Our findings also indicated that reproductive investment and body condition increase with elevation, suggesting that alpine populations are locally adapted.

Telomere length mirrors age structure along a 2200-m altitudinal gradient in a Mediterranean lizard.

The timing of organisms' senescence is developmentally programmed but also shaped by the interaction between environmental inputs and life-history traits. In ectotherms, ageing dynamics are still poorly understood even though their body temperature, metabolism, or growth trajectory are very sensitive to environmental changes. Here, we investigated the role of life-history traits such as age, sex, body size, body condition, and tail autotomy (i.e self-amputation) in shaping telomere length in six populations of the Algerian sand lizard (Psammodromus algirus) distributed along an elevational gradient from 300 to 2500 m above the sea level. Additionally, we compiled the available information on reptiles' telomere length in a review table. Our cross-sectional study shows that older lizards have longer telomeres, which might be mostly linked to the selective disappearance of individuals with shorter telomeres or, alternatively, mediated by a higher expression of telomerase across their life. In fact, variation in telomere length across elevation was explained by age structure of lizards; thus, in contrast to our predictions, altitude had no effect on telomere length in this study system. Telomere length was unaffected by tail regeneration and was sex-independent, but positively correlated with body condition, which might be linked to high somatic investment. Hence, our results suggest that life-history traits such as age or body condition can be major drivers of telomere dynamics for this species, whereas environmental conditions apparently had scarce or no effects on lizard telomeres. Our findings emphasize the relevance of understanding species' life histories for fully disentangling the causes and consequences of differences in ageing in ectotherms.

Multiple color patches and parasites in Sceloporus occidentalis: differential relationships by sex and infection.

Parasites generally have a negative influence on the color expression of their hosts. Sexual selection theory predicts resistant high-quality individuals should show intense coloration, whereas susceptible low-quality individuals would show poor coloration. However, intensely colored males of different species of Old and New World lizards were more often infected by hemoparasites. These results suggest that high-quality males, with intense coloration, would suffer higher susceptibility to hemoparasites. This hypothesis remains poorly understood and contradicts general theories on sexual selection. We surveyed a population of Sceloporus occidentalis for parasites and found infections by the parasite genera Lankesterella and Acroeimeria. In this population, both males and females express ventral blue and yellow color patches. Lankesterella was almost exclusively infecting males. The body size of the males significantly predicted the coloration of both blue and yellow patches. Larger males showed darker (lower lightness) blue ventral patches and more saturated yellow patches that were also orange-skewed. Moreover, these males were more often infected by Lankesterella than smaller males. The intestinal parasite Acroeimeria infected both males and females. The infection by intestinal parasites of the genus Acroeimeria was the best predictor for the chroma in the blue patch of the males and for hue in the yellow patch of the females. Those males infected by Acroeimeria expressed blue patches with significantly lower chroma than the uninfected males. However, the hue of the yellow patch was not significantly different between infected and uninfected females. These results suggest a different effect of Lankesterella and Acroeimeria on the lizards. On the one hand, the intense coloration of male lizards infected by Lankesterella suggested high-quality male lizards may tolerate it. On the other hand, the low chroma of the blue coloration of the infected males suggested that this coloration could honestly express the infection by Acroeimeria.

Opposed elevational variation in prevalence and intensity of endoparasites and their vectors in a lizard.

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.

Prey availability, prey selection, and trophic niche width in the lizard Psammodromus algirus along an elevational gradient.

Mountains imply enormous environmental variation, with alpine habitats entailing harsh environments, especially for ectotherms such as lizards. This environmental variability also may imply variation in prey availability. However, little is known about how lizard trophic ecology varies with elevation. In this study, we analyze diet, prey availability, prey selection, and trophic niche width in the lacertid lizard Psammodromus algirus along a 2,200-m elevational gradient in the Sierra Nevada (SE Spain). The analysis of fecal samples has shown that Orthoptera, Formicidae, Hemiptera, Coleoptera, and Araneae are the main prey, although, according to their abundance in pitfall traps, Formicidae and Coleoptera are rejected by the lizard whereas Orthoptera, Hemiptera, and Araneae are preferred. Prey abundance and diversity increase with elevation and diet subtly varies along with the elevational gradient. The consumption of Coleoptera increases with elevation probably as a consequence of the lizard foraging more in open areas while basking. The electivity for Araneae increases with elevation. Araneae are rejected in the lowlands-where they are relatively abundant-whereas, at high elevation, this lizard positively selects them, despite they being less abundant. The lizard trophic niche width expands with elevation due to concomitant greater prey diversity and hence this lizard feeds on more prey types in highlands. Although no sex difference in diet has been found, the trophic niche is broader in females than males. As a whole, alpine lizards show a trophic niche similar to that found at lower elevations, suggesting that P. algirus is well adapted to the harsh environment found in alpine areas.

Variations in chemical sexual signals of Psammodromus algirus lizards along an elevation gradient may reflect altitudinal variation in microclimatic conditions.

Chemical signals used in intraspecific communication are expected to evolve or to show phenotipic plasticity to maximize efficacy in the climatic conditions of a given environment. Elevational environmental gradients in mountains provide a good opportunity to test this hypothesis by examining variation in characteristics of signals in species found across different elevations with different climatic conditions. We analyzed by gas chromatography-mass spectrometry (GC-MS) the lipophilic fraction of the femoral gland secretions of male lizards Psammodromus algirus (Fam. Lacertidae) from six localities located along a 2200 m elevational gradient at Sierra Nevada Mountains (SE Spain). There was elevational clinal variation in climatic variables, number of femoral pores and in the relative proportions of some classes of compounds (i.e., ethyl esters of fatty acids, waxy esters, and aldehydes) but not others. We discuss how this variation would result in different physicochemical properties of the entire femoral secretion, which might help optimize the efficacy of chemical signals under the particular microclimatic conditions at each elevation.

Elevational variation in body-temperature response to immune challenge in a lizard.

Immunocompetence benefits animal fitness by combating pathogens, but also entails some costs. One of its main components is fever, which in ectotherms involves two main types of costs: energy expenditure and predation risk. Whenever those costs of fever outweigh its benefits, ectotherms are expected not to develop fever, or even to show hypothermia, reducing costs of thermoregulation and diverting the energy saved to other components of the immune system. Environmental thermal quality, and therefore the thermoregulation cost/benefit balance, varies geographically. Hence, we hypothesize that, in alpine habitats, immune-challenged ectotherms should show no thermal response, given that (1) hypothermia would be very costly, as the temporal window for reproduction is extremely small, and (2) fever would have a prohibitive cost, as heat acquisition is limited in such habitat. However, in temperate habitats, immune-challenged ectotherms might show a febrile response, due to lower cost/benefit balance as a consequence of a more suitable thermal environment. We tested this hypothesis in Psammodromus algirus lizards from Sierra Nevada (SE Spain), by testing body temperature preferred by alpine and non-alpine lizards, before and after activating their immune system with a typical innocuous pyrogen. Surprisingly, non-alpine lizards responded to immune challenge by decreasing preferential body-temperature, presumably allowing them to save energy and reduce exposure to predators. On the contrary, as predicted, immune-challenged alpine lizards maintained their body-temperature preferences. These results match with increased costs of no thermoregulation with elevation, due to the reduced window of time for reproduction in alpine environment.

Thermoregulation in the lizard Psammodromus algirus along a 2200-m elevational gradient in Sierra Nevada (Spain).

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.

Thermal dependence of sprint performance in the lizard Psammodromus algirus along a 2200-meter elevational gradient: Cold-habitat lizards do not perform better at low temperatures.

Sprint speed has a capital relevance in most animals' fitness, mainly for fleeing from predators. Sprint performance is maximal within a certain range of body temperatures in ectotherms, whose thermal upkeep relies on exogenous thermal sources. Ectotherms can respond to diverse thermal environments either by shifting their thermal preferences or maintaining them through different adaptive mechanisms. Here, we tested whether maximum sprint speed of a lizard that shows conservative thermal ecology along a 2200-meter elevational gradient differs with body temperature in lizards from different elevations. Lizards ran faster at optimum than at suboptimum body temperature. Notably, high-elevation lizards were not faster than mid- and low-elevation lizards at suboptimum body temperature, despite their low-quality thermal environment. This result suggests that both preferred body temperature and thermal dependence of speed performance are co-adapted along the elevational gradient. High-elevation lizards display a number of thermoregulatory strategies that allow them to achieve high optimum body temperatures in a low thermal-quality habitat and thus maximize speed performance. As for reproductive condition, we did not find any effect of it on sprint speed, or any significant interaction with elevation or body temperature. However, strikingly, gravid females were significantly slower than males and non-gravid females at suboptimum temperature, but performed similarly well at optimal temperature.

Ultraviolet radiation does not increase oxidative stress in the lizard Psammodromus algirus along an elevational gradient.

Lizards, as ectotherms, spend much time basking for thermoregulating exposed to solar radiation. Consequently, they are subjected to ultraviolet radiation (UVR), which is the most harmful component of solar radiation spectrum. UVR can provoke damages, from the molecular to tissue level, even cause death. Photooxidation triggered by UVR produces reactive oxidative species (ROS). When antioxidant machinery cannot combat the ROS concentration, oxidative stress occurs in the organisms. Given that UVR increases with elevation, we hypothesised that lizards from high elevations should be better adapted against UVR than lizards from lower elevations. In this work, we test this hypothesis in Psammodromus algirus along an elevation gradient (three elevational belts, from 300 to 2500 m above sea level). We ran an experiment in which lizards from each elevation belt were exposed to 5-hour doses of UVR (UV-light bulb, experimental group) or photosynthetically active radiation (white-light bulb, control group) and, 24 h after the exposure, we took tissue samples from the tail. We measured oxidative damage (lipid and protein peroxidation) and antioxidant capacity as oxidative-stress biomarkers. We found no differences in oxidative stress between treatments. However, consistent with a previous work, less oxidative damage appeared in lizards from the highlands. We conclude that UVR is not a stressor agent for P. algirus; however, our findings suggest that the lowland environment is more oxidative for lizards. Therefore, P. algirus is well adapted to inhabit a large elevation range, and this would favour the lizard in case it ascends in response to global climate change.

Oxidative stress decreases with elevation in the lizard Psammodromus algirus.

Oxidative stress is considered one of the main ecological and evolutionary forces. Several environmental stressors vary geographically and thus organisms inhabiting different sites face different oxidant environments. Nevertheless, there is scarce information about how oxidative damage and antioxidant defences vary geographically in animals. Here we study how oxidative stress varies from lowlands (300-700 m asl) to highlands (2200-2500 m asl) in the lizard Psammodromus algirus. To accomplish this, antioxidant enzymatic activity (catalase, superoxide dismutase, glutathione peroxidase, glutathione reductase, glutathione transferase, DT-diaphorase) and lipid peroxidation were assayed in tissue samples from the lizards' tail. Lipid peroxidation was higher in individuals from lowlands than from highlands, indicating higher oxidative stress in lowland lizards. These results suggest that environmental conditions are less oxidant at high elevations with respect to low ones. Therefore, our study shows that oxidative stress varies geographically, which should have important consequences for our understanding of geographic variation in physiology and life-history of organisms.