Lizards from warm and declining populations are born with extremely short telomeres.

Aging is the price to pay for acquiring and processing energy through cellular activity and life history productivity. Climate warming can exacerbate the inherent pace of aging, as illustrated by a faster erosion of protective telomere DNA sequences. This biomarker integrates individual pace of life and parental effects through the germline, but whether intra- and intergenerational telomere dynamics underlies population trends remains an open question. Here, we investigated the covariation between life history, telomere length (TL), and extinction risk among three age classes in a cold-adapted ectotherm (Zootoca vivipara) facing warming-induced extirpations in its distribution limits. TL followed the same threshold relationships with population extinction risk at birth, maturity, and adulthood, suggesting intergenerational accumulation of accelerated aging rate in declining populations. In dwindling populations, most neonates inherited already short telomeres, suggesting they were born physiologically old and unlikely to reach recruitment. At adulthood, TL further explained females' reproductive performance, switching from an index of individual quality in stable populations to a biomarker of reproductive costs in those close to extirpation. We compiled these results to propose the aging loop hypothesis and conceptualize how climate-driven telomere shortening in ectotherms may accumulate across generations and generate tipping points before local extirpation.

Breeding phenology drives variation in reproductive output, reproductive costs, and offspring fitness in a viviparous ectotherm.

Phenological advances are a widespread response to global warming and can contribute to determine the climate vulnerability of organisms, particularly in ectothermic species, which are highly dependent on ambient temperatures to complete their life cycle. Yet, the relative contribution of breeding dates and temperature conditions during gestation on fitness of females and their offspring is poorly documented in reptiles. Here, we exposed females of the common lizard Zootoca vivipara to contrasting thermal scenarios (cold vs. hot treatment) during gestation and quantified effects of parturition dates and thermal treatment on life-history traits of females and their offspring for 1 year. Overall, our results suggest that parturition date has a greater impact than thermal conditions during gestation on life history strategies. In particular, we found positive effects of an earlier parturition date on juvenile survival, growth, and recruitment suggesting that environmental-dependent selection and/or differences in parental quality between early and late breeders underlie seasonal changes in offspring fitness. Yet, an earlier parturition date compromised the energetic condition of gravid females, which suggests the existence of a mother-offspring conflict regarding the optimization of parturition dates. While numerous studies focused on the direct effects of alterations in incubation temperatures on reptile life-history traits, our results highlight the importance of considering the role of breeding phenology in assessing the short- and long-term effects of thermal developmental plasticity.

Prey consumption does not restore hydration state but mitigates the energetic costs of water deprivation in an insectivorous lizard.

To cope with limited availability of drinking water in their environment, terrestrial animals have developed numerous behavioral and physiological strategies including maintaining an optimal hydration state through dietary water intake. Recent studies performed in snakes, which are generalist carnivorous reptiles, suggest that the benefits of dietary water intake are negated by hydric costs of digestion. Most lizards are generalist insectivores that can shift their prey types, but firm experimental demonstration of dietary water intake is currently missing in these organisms. Here, we performed an experimental study in the common lizard Zootoca vivipara, a keystone mesopredator from temperate climates exhibiting a great diversity of prey in its mesic habitats, in order to investigate the effects of food consumption and prey type on physiological responses to water deprivation. Our results indicate that common lizards cannot improve their hydration state through prey consumption, irrespective of prey type, suggesting that they are primarily dependent upon drinking water. Yet, high-quality prey consumption reduced the energetic costs of water deprivation, potentially helping lizards to conserve a better body condition during periods of limited water availability. These findings have important implications for understanding the physiological responses of ectotherms to water stress, and highlight the complex interactions between hydration status, energy metabolism and feeding behavior in insectivorous lizards.

Behavioural hydroregulation protects against acute effects of drought in a dry-skinned ectotherm.

During extreme climate events, behavioural thermoregulation may buffer ectotherms from thermal stress and overheating. However, heatwaves are also combined with dry spells and limited water availability, and how much individuals can behaviourally mitigate dehydration risks through microclimate selection remains largely unknown. Herein, we investigated the behavioural and physiological responses to changes in air and microhabitat humidity in a terrestrial ectotherm, the asp viper (Vipera aspis). We exposed individuals to a simulated heatwave together with water deprivation for 3 weeks, and manipulated air water vapour density (wet air vs. dry air) and microclimate (wet shelter vs. dry shelter) in a two-by-two factorial design. Dry air conditions led to substantial physiological dehydration and muscle wasting. Vipers exposed to dry air used more often a shelter that offered a moist microclimate, which reduced dehydration and muscle wasting at the individual level. These results provide the first experimental evidence that active behavioural hydroregulation can mitigate specific physiological stress responses caused by a dry spell in an ectotherm. Future studies investigating organismal responses to climate change should consider moisture gradient in the habitat and integrate both hydroregulation and thermoregulation behaviours.

Two stressors are worse than one: combined heatwave and drought affect hydration state and glucocorticoid levels in a temperate ectotherm.

Heatwaves and droughts are becoming more intense and frequent with climate change. These extreme weather events often occur simultaneously and may alter organismal physiology, yet their combined impacts remain largely unknown. Here, we experimentally investigated physiological responses of a temperate ectotherm, the asp viper (Vipera aspis), to a simulated heatwave and drought. We applied a two-by-two factorial design by manipulating the daily temperature cycle (control versus heatwave) and the water availability (water available versus water deprived) over a month followed by exposure to standard thermal conditions with ad libitum access to water. Simulated heatwave and water deprivation additively increased mass loss, while water deprivation led to greater plasma osmolality (dehydration). Mass gain from drinking after the treatment period was higher in vipers from the heatwave and water-deprived group, suggesting that thirst was synergistically influenced by thermal and water constraints. Heatwave conditions and water deprivation also additively increased baseline corticosterone levels but did not influence basal metabolic rate and plasma markers of oxidative stress. Our results demonstrate that a short-term exposure to combined heatwave and drought can exacerbate physiological stress through additive effects, and interactively impact behavioral responses to dehydration. Considering combined effects of temperature and water availability is thus crucial to assess organismal responses to climate change.

Environmental conditions and male quality traits simultaneously explain variation of multiple colour signals in male lizards.

Male lizards often display multiple pigment-based and structural colour signals which may reflect various quality traits (e.g. performance, parasitism), with testosterone (T) often mediating these relationships. Furthermore, environmental conditions can explain colour signal variation by affecting processes such as signal efficacy, thermoregulation and camouflage. The relationships between colour signals, male quality traits and environmental factors have often been analysed in isolation, but simultaneous analyses are rare. Thus, the response of multiple colour signals to variation in all these factors in an integrative analysis remains to be investigated. Here, we investigated how multiple colour signals relate to their information content, examined the role of T as a potential mediator of these relationships and how environmental factors explain colour signal variation. We performed an integrative study to examine the covariation between three colour signals (melanin-based black, carotenoid-based yellow-orange and structural UV), physiological performance, parasitism, T levels and environmental factors (microclimate, forest cover) in male common lizards Zootoca vivipara from 13 populations. We found that the three colour signals conveyed information on different aspects of male condition, supporting a multiple message hypothesis. T influenced only parasitism, suggesting that T does not directly mediate the relationships between colour signals and their information content. Moreover, colour signals became more saturated in forested habitats, suggesting an adaptation to degraded light conditions, and became generally brighter in mesic conditions, in contradiction with the thermal melanism hypothesis. We show that distinct individual quality traits and environmental factors simultaneously explain variations of multiple colour signals with different production modes. Our study therefore highlights the complexity of colour signal evolution, involving various sets of selective pressures acting at the same time, but in different ways depending on colour production mechanism.

Les lézards mâles arborent souvent plusieurs signaux colorés de nature pigmentaire et structurale qui reflètent de multiples traits de qualité (e.g. performance, parasitisme), et la testostérone (T) joue souvent un rôle de médiateur dans ces relations. En outre, les conditions environnementales peuvent également expliquer les variations des signaux colorés en influençant des aspects tels que l'efficacité des signaux, la thermorégulation ou le camouflage. Les relations entre signaux colorés, traits de qualité individuelle et facteurs environnementaux ont souvent été analysées séparément, mais rarement de manière simultanée. Ainsi, la réponse de ces multiples signaux colorés aux variations de tous ces facteurs reste à explorer dans le contexte d'une étude intégrative. Ici, nous explorons la relation entre ces multiples signaux colorés et leur contenu informatif, nous examinons le rôle de T comme médiateur potentiel de ces relations et nous recherchons si les conditions environnementales expliquent la variation de ces signaux colorés. Nous avons mené une étude intégrative afin d'examiner la covariation entre trois types de signaux colorés (noir produit par la mélanine, jaune-orange produit par les caroténoïdes et UV produit par des éléments structuraux), la performance physiologique, le parasitisme, les niveaux de T et les conditions environnementales (e.g. microclimat, couverture forestière) chez des mâles du lézard vivipare (Zootoca vivipara) provenant de 13 populations. Nos résultats indiquent que les trois signaux colorés transmettent des informations sur différents aspects de la condition des mâles, en accord avec l'hypothèse de « messages multiples ¼. T influence uniquement le parasitisme, suggérant que T n'agit pas en tant que médiateur des relations entre ces signaux colorés et leur contenu informatif. De plus, les signaux colorés sont plus saturés dans les habitats les plus forestiers, ce qui suggère une adaptation à des conditions lumineuses dégradées. Enfin, les signaux colorés sont plus intenses lorsque les conditions sont mésiques, en contradiction avec l'hypothèse du mélanisme thermal. Nous démontrons que différents traits de qualité individuelle et facteurs environnementaux expliquent de manière simultanée les variations de multiples signaux colorés impliquant différents modes de production. Notre étude souligne ainsi la complexité de l'évolution des signaux colorés, qui implique plusieurs types de pressions de sélection agissant en même temps mais dans des directions différentes selon le mode de production.

Ecotrons: Powerful and versatile ecosystem analysers for ecology, agronomy and environmental science.

Ecosystems integrity and services are threatened by anthropogenic global changes. Mitigating and adapting to these changes require knowledge of ecosystem functioning in the expected novel environments, informed in large part through experimentation and modelling. This paper describes 13 advanced controlled environment facilities for experimental ecosystem studies, herein termed ecotrons, open to the international community. Ecotrons enable simulation of a wide range of natural environmental conditions in replicated and independent experimental units while measuring various ecosystem processes. This capacity to realistically control ecosystem environments is used to emulate a variety of climatic scenarios and soil conditions, in natural sunlight or through broad-spectrum lighting. The use of large ecosystem samples, intact or reconstructed, minimizes border effects and increases biological and physical complexity. Measurements of concentrations of greenhouse trace gases as well as their net exchange between the ecosystem and the atmosphere are performed in most ecotrons, often quasi continuously. The flow of matter is often tracked with the use of stable isotope tracers of carbon and other elements. Equipment is available for measurements of soil water status as well as root and canopy growth. The experiments ran so far emphasize the diversity of the hosted research. Half of them concern global changes, often with a manipulation of more than one driver. About a quarter deal with the impact of biodiversity loss on ecosystem functioning and one quarter with ecosystem or plant physiology. We discuss how the methodology for environmental simulation and process measurements, especially in soil, can be improved and stress the need to establish stronger links with modelling in future projects. These developments will enable further improvements in mechanistic understanding and predictive capacity of ecotron research which will play, in complementarity with field experimentation and monitoring, a crucial role in exploring the ecosystem consequences of environmental changes.

Intense nocturnal warming alters growth strategies, colouration and parasite load in a diurnal lizard.

In the past decades, nocturnal temperatures have been playing a disproportionate role in the global warming of the planet. Yet, they remain a neglected factor in studies assessing the impact of global warming on natural populations. Here, we question whether an intense augmentation of nocturnal temperatures is beneficial or deleterious to ectotherms. Physiological performance is influenced by thermal conditions in ectotherms and an increase in temperature by only 2°C is sufficient to induce a disproportionate increase in metabolic expenditure. Warmer nights may expand ectotherms' species thermal niche and open new opportunities for prolonged activities and improve foraging efficiency. However, increased activity may also have deleterious effects on energy balance if exposure to warmer nights reduces resting periods and elevates resting metabolic rate. We assessed whether warmer nights affected an individual's growth, dorsal skin colouration, thermoregulation behaviour, oxidative stress status and parasite load by exposing yearling common lizards (Zootoca vivipara) from four populations to either ambient or high nocturnal temperatures for approximately 5 weeks. Warmer nocturnal temperatures increased the prevalence of ectoparasitic infestation and altered allocation of resources towards structural growth rather than storage. We found no change in markers for oxidative stress. The thermal treatment did not influence thermal preferences, but influenced dorsal skin brightness and luminance, in line with a predicted acclimation response in colder environments to enhance heat gain from solar radiation. Altogether, our results highlight the importance of considering nocturnal warming as an independent factor affecting ectotherms' life history in the context of global climate change. ​.

Additive effects of temperature and water availability on pregnancy in a viviparous lizard.

One of the greatest current threats to biodiversity is climate change. However, understanding of organismal responses to fluctuations in temperature and water availability is currently lacking, especially during fundamental life-history stages such as reproduction. To further explore how temperature and water availability impact maternal physiology and reproductive output, we used the viviparous form of the European common lizard (Zootoca vivipara) in a two-by-two factorial design manipulating both hydric and thermal conditions, for the first time. We collected blood samples and morphological measurements during early pregnancy and post-parturition to investigate how water availability, temperature and a combination of the two influence maternal phenology, morphology, physiology and reproductive output. We observed that dehydration during gestation negatively affects maternal physiological condition (lower mass gain, higher tail reserve mobilization) but has little effect on reproductive output. These effects are mainly additive to temperature regimes, with a proportional increase in maternal costs in warmer environments. Our study demonstrates the importance of considering combined effects of water and temperature when investigating organismal responses to climate changes, especially during periods crucial for species survival such as reproduction.

Micro-geographic shift between negligible and actuarial senescence in a wild snake.

While it has long been known that species have contrasted life expectancy (pace of mortality) and generation time (pace of reproduction), recent studies have also uncovered that the shape of adult age trajectories of mortality and reproduction can vary remarkably among species along a continuum of senescence ranging from strong deterioration (senescence), insignificant deterioration (negligible senescence) to improvement with advancing age (negative senescence). As for many long-lived ectotherms with asymptotic growth and increasing reproductive output with age, snakes are good candidates for negligible senescence to occur. Yet, intraspecific variation in the pace and shape of actuarial and reproductive senescence across wild populations of these species remains to be explored. Here, we used 37 years of mark-recapture data in two nearby habitats inside a meadow viper Vipera ursinii population to quantify life expectancies, generation times and the shape of actuarial and reproductive senescence. Female vipers maintained stable reproductive performances at old ages, even when accounting for the predicted increase of fertility with body size, providing evidence for negligible reproductive senescence in both habitats. Males had a higher adult mortality and a shorter life expectancy on average than females and actuarial senescence shifted from negligible senescence in the optimal habitat to strong senescence in the sub-optimal habitat. Overall, these results demonstrate that micro-geographic environmental variation can generate qualitative shifts in actuarial senescence patterns. This highlights that taking into account the within-species plasticity of age-dependent trajectories could prove useful in better understanding what determines the evolution of life-history age trajectories.

La longévité et la manière de vieillir varient grandement entre espèces. Chez certaines espèces, la survie et la reproduction déclinent avec l'âge : les individus souffrent de sénescence. Chez d'autres, les vieux individus maintiennent des performances reproductives et de survie similaires à celles des jeunes : la sénescence est alors dite négligeable. Enfin, la sénescence peut être négative quand la survie des adultes ou leur reproduction s'améliorent avec l'âge. Les formes des trajectoires de mortalité et de reproduction avec l'âge sont encore mal connues chez les serpents, mais la théorie évolutive prédit une sénescence négligeable chez les espèces qui, comme les serpents, continuent à grandir à l'âge adulte et dont les femelles produisent des portées dont l'effectif augmente avec la taille corporelle de la mère. Nous avons exploité ici 37 années de données de capture-marquage-recapture de vipères d'Orsini, Vipera ursinii ursinii, dans deux habitats voisins d'une petite population isolée du Sud-Est de la France pour quantifier l'espérance de vie, les temps de génération et les formes de la sénescence de survie et de la sénescence reproductrice. Nous montrons que les vipères femelles parviennent à maintenir de bonnes performances reproductives même à des âges avancés, illustrant une sénescence reproductrice négligeable dans cette population. Nous montrons aussi que la forme de la sénescence de survie varie fortement à une très petite échelle géographique : dans l'habitat optimal, la survie adulte des vipères demeure stable quel que soit leur âge alors qu'à quelques centaines de mètres, dans un habitat moins favorable, les vipères subissent une forte sénescence de survie. Ces résultats démontrent que de forts changements qualitatifs dans les profils de sénescence peuvent être observés à une échelle spatiale réduite. Cette plasticité intra-spécifique de la sénescence offre de nouvelles perspectives pour mieux comprendre ce qui détermine l'évolution de la sénescence.

Short-term change in water availability influences thermoregulation behaviours in a dry-skinned ectotherm.

Mechanistic models of terrestrial ectotherms predict that climate warming will induce activity restriction due to heat stress and loss of shade, leading to the extinction of numerous populations. Such models rely on the assumption that activity patterns are dictated by simple temperature thresholds independent of changes in water availability. However, changes in water availability may further influence thermoregulation behaviour of ectotherms through dehydration risk perception, changes in water balance or changes in microclimatic conditions. Here, we experimentally assess the interactive effects of thermal conditions and water availability on activity patterns, shade selection and thermoregulation efficiency in a model ectothermic species. Thermoregulation behaviour of adult common lizards Zootoca vivipara was monitored in outdoor mesocosms as we manipulated water availability, providing water as mist in the morning and free-standing water during the daytime. We recorded operative temperatures and micro-meteorological conditions to infer thermal constraints and dehydration risk. Activity and shade selection were better predicted by continuous changes in thermal conditions and dehydration risk, respectively, than by threshold functions. In addition, water supplementation increased activity in males and reduced shade selection in both sexes, most probably as a behavioural response to the perception of a stronger dehydration risk. Water supplementation also influenced the thermal quality of the environment, which in turn altered daily activity patterns and thermoregulation statistics. This demonstrates that dual effects of heat and water stress on activity patterns may lead to stronger activity restriction as a result of climate change than currently predicted.

Chronic water restriction triggers sex-specific oxidative stress and telomere shortening in lizards.

Animals use a variety of strategies to avoid acute dehydration and death. Yet, how chronic exposure to sub-lethal dehydration may entail physiological and fitness costs remains elusive. In this study, we experimentally tested if water restriction causes increased oxidative stress (OS) and telomere length (TL) shortening, two well-described mediators of environment-fitness relationships. We exposed 100 yearling female and male common lizards (Zootoca vivipara) either to a 51-day period of water restriction or to water ad libitum, followed by 45 days in common garden outdoor conditions. We measured the kinetic changes in OS and TL and found that water-restricted males had enhanced antioxidant defences and decreased oxidative damage at day 36, whereas females did not immediately respond. A month and a half after water restriction, both sexes experienced a drop in antioxidant capacity but only males exhibited significant TL shortening. In the following 3 years, we found that lizards with longer initial TL and those who maintained stronger antioxidant defences experienced higher longevity, irrespective of sex and water restriction. Together, these results unravelled sex-specific responses to water restriction, with potential applications in better understanding the physiological costs of increasing summer droughts as a result of global climate change.

Chronic elevation of glucorticoids late in life generates long lasting changes in physiological state without a life history switch.

Chronic stressors have profound impacts on phenotypes and life history strategies on the short term, but delayed effects of stress experienced late in life remain poorly investigated in wild populations. Here, we used a combined laboratory and field experiment to test if chronic stress late in life has immediate and delayed effects on physiological and demographic traits in the common lizard, Zootoca vivipara. We increased plasma corticosterone levels in adults and yearlings during three weeks of the post-reproductive season. We quantified immediate responses in the laboratory, delayed intra-generational effects in field enclosures one month and one year later during the next reproductive season, and delayed inter-generational effects in the first generation of offspring. Our phenotypic assays included metabolism, immune capacities, lipid metabolism and oxidative stress. Relative to placebos, lizards treated with corticosterone had higher body condition and lower oxidative damages but an increased skin swelling response directly after the manipulation. Delayed responses in field enclosures were of three types. First, we found catch-up growth for body mass such the placebos had similar body conditions one month after the laboratory manipulation. Second, we found persistent differences in oxidative damages during one month but not one year later. Third, during the next reproductive season, corticosterone-treated females had higher levels of plasma triglycerides, whereas corticosterone-treated individuals had a higher skin swelling response. We found no delayed inter-generational effects on demographic traits of offspring. Our study demonstrates the potential for long-lasting physiological consequences of chronic corticosterone enhancement despite no obvious changes in life history.

Reduction in baseline corticosterone secretion correlates with climate warming and drying across wild lizard populations.

Climate change should lead to massive loss of biodiversity in most taxa, but the detailed physiological mechanisms underlying population extinction remain largely elusive so far. In vertebrates, baseline levels of hormones such as glucocorticoids (GCs) may be indicators of population state as their secretion to chronic stress can impair survival and reproduction. However, the relationship between GC secretion, climate change and population extinction risk remains unclear. In this study, we investigated whether levels of baseline corticosterone (the main GCs in reptiles) correlate with environmental conditions and associated extinction risk across wild populations of the common lizard Zootoca vivipara. First, we performed a cross-sectional comparison of baseline corticosterone levels along an altitudinal gradient among 14 populations. Then, we used a longitudinal study in eight populations to examine the changes in corticosterone levels following the exposure to a heatwave period. Unexpectedly, baseline corticosterone decreased with increasing thermal conditions at rest in females and was not correlated with extinction risk. In addition, baseline corticosterone levels decreased after exposure to an extreme heatwave period. This seasonal corticosterone decrease was more pronounced in populations without access to standing water. We suggest that low basal secretion of corticosterone may entail downregulating activity levels and limit exposure to adverse climatic conditions, especially to reduce water loss. These new insights suggest that rapid population decline might be preceded by a downregulation of the corticosterone secretion.

Habitat degradation increases stress-hormone levels during the breeding season, and decreases survival and reproduction in adult common lizards.

The allostatic load model describes how individuals maintain homeostasis in challenging environment and posits that costs induced by a chronic perturbation (i.e., allostatic load) are correlated to the secretion of glucocorticoids, such as corticosterone. Habitat perturbations from anthropogenic activities are multiple and functional responses to those are still unclear. Here, we manipulated the habitat quality in 24 semi-natural populations of the common lizard during 1 year. We tested the predictions of the allostatic load model that habitat degradation should increase baseline corticosterone levels, and should induce concomitant physiological changes, such as lipid mobilization and lower immunocompetence, and demographic changes, such as lower body growth, survival and/or reproductive performances. Our results highlight stage-dependent effects of habitat degradation on physiological traits during the breeding season: adult lizards had higher baseline corticosterone levels and yearling lizards had a lower inflammatory response than adults, whereas juveniles had higher circulating lipid levels than yearlings and adults without concomitant change in corticosterone levels. In addition, habitat degradation reduced the performances of adults but not of juveniles: in low habitat quality populations, adult males had a lower survival and females had a smaller fecundity. These results are in accordance with the allostatic load model given that allostatic load was detected only during the season and in life stages of maximal energy expenditure. This underlines the importance to account for individual energy requirements to better understand demographic responses to habitat perturbation.

Chronic stress, energy transduction, and free-radical production in a reptile.

Stress hormones, such as corticosterone, play a crucial role in orchestrating physiological reaction patterns shaping adapted responses to stressful environments. Concepts aiming at predicting individual and population responses to environmental stress typically consider that stress hormones and their effects on metabolic rate provide appropriate proxies for the energy budget. However, uncoupling between the biochemical processes of respiration, ATP production, and free-radical production in mitochondria may play a fundamental role in the stress response and associated life histories. In this study, we aim at dissecting sub-cellular mechanisms that link these three processes by investigating both whole-organism metabolism, liver mitochondrial oxidative phosphorylation processes (O2 consumption and ATP production) and ROS emission in Zootoca vivipara individuals exposed 21 days to corticosterone relative to a placebo. Corticosterone enhancement had no effect on mitochondrial activity and efficiency. In parallel, the corticosterone treatment increased liver mass and mitochondrial protein content suggesting a higher liver ATP production. We also found a negative correlation between mitochondrial ROS emission and plasma corticosterone level. These results provide a proximal explanation for enhanced survival after chronic exposure to corticosterone in this species. Importantly, none of these modifications affected resting whole-body metabolic rate. Oxygen consumption, ATP, and ROS emission were thus independently affected in responses to corticosterone increase suggesting that concepts and models aiming at linking environmental stress and individual responses may misestimate energy allocation possibilities.

Water availability and environmental temperature correlate with geographic variation in water balance in common lizards.

Water conservation strategies are well documented in species living in water-limited environments, but physiological adaptations to water availability in temperate climate environments are still relatively overlooked. Yet, temperate species are facing more frequent and intense droughts as a result of climate change. Here, we examined variation in field hydration state (plasma osmolality) and standardized evaporative water loss rate (SEWL) of adult male and pregnant female common lizards (Zootoca vivipara) from 13 natural populations with contrasting air temperature, air humidity, and access to water. We found different patterns of geographic variation between sexes. Overall, males were more dehydrated (i.e. higher osmolality) than pregnant females, which likely comes from differences in field behaviour and water intake since the rate of SEWL was similar between sexes. Plasma osmolality and SEWL rate were positively correlated with environmental temperature in males, while plasma osmolality in pregnant females did not correlate with environmental conditions, reproductive stage or reproductive effort. The SEWL rate was significantly lower in populations without access to free standing water, suggesting that lizards can adapt or adjust physiology to cope with habitat dryness. Environmental humidity did not explain variation in water balance. We suggest that geographic variation in water balance physiology and behaviour should be taken account to better understand species range limits and sensitivity to climate change.

Geographic variation and acclimation effects on thermoregulation behavior in the widespread lizard Liolaemus pictus.

Populations at the warm range margins of the species distribution may be at the greatest risks of extinction from global warming unless they can tolerate extreme environmental conditions. Yet, some studies suggest that the thermal behavior of some lizard species is evolutionarily rigid. During two successive years, we compared the thermal biology of two populations of Liolaemus pictus living at the northern (warmer) and one population living at the southern (colder) range limits, thus spanning an 800km latitudinal distance. Populations at the two range margins belong to two deeply divergent evolutionary clades. We quantified field body temperatures (Tb), laboratory preferred body temperatures (PBT), and used operative temperature data (Te) to calculate the effectiveness of thermoregulation (E). During one year in all populations, we further exposed half of the lizards to a cold or a hot acclimation treatment to test for plasticity in the thermal behavior. The environment at the southern range limit was characterized by cooler weather and lower Te. Despite that, females had higher Tb and both males and females had higher PBT in the southernmost population (or clade) than in the northernmost populations. Acclimation to cold conditions led to higher PBT in all populations suggesting that plastic responses to thermal conditions, instead of evolutionary history, may contribute to geographic variation. Lizards regulated moderately well their body temperature (E≈0.7): they avoided warm microhabitats in the northern range but capitalized on warm microhabitats in the southern range. We review literature data to show that Liolaemus species increase their thermoregulation efficiency in thermally challenging environments. Altogether, this indicates that habitats of low thermal quality generally select against thermoconformity in these lizards.

Climate and habitat interact to shape the thermal reaction norms of breeding phenology across lizard populations.

Substantial plastic variation in phenology in response to environmental heterogeneity through time in the same population has been uncovered in many species. However, our understanding of differences in reaction norms of phenology among populations from a given species remains limited. As the plasticity of phenological traits is often influenced by local thermal conditions, we expect local temperature to generate variation in the reaction norms between populations. Here, we explored temporal variation in parturition date across 11 populations of the common lizard (Zootoca vivipara) from four mountain chains as a function of air temperatures during mid-gestation. We characterized among-population variation to assess how local weather conditions (mean and variance of ambient temperatures during mid-gestation) and habitat openness (an index of anthropogenic disturbance) influence the thermal reaction norms of the parturition date. Our results provide evidence of interactive effects of anthropogenic disturbance and thermal conditions, with earlier parturition dates in warmer years on average especially in closed habitats. Variation in the reaction norms for parturition date was correlated with mean local thermal conditions at a broad geographical scale. However, populations exposed to variable thermal conditions had flatter thermal reaction norms. Assessing whether environmental heterogeneity drives differentiation among reaction norms is crucial to estimate the capacity of different populations to contend with projected climatic and anthropogenic challenges.

Population viability analysis of plant and animal populations with stochastic integral projection models.

Integral projection models (IPM) make it possible to study populations structured by continuous traits. Recently, Vindenes et al. (Ecology 92:1146-1156, 2011) proposed an extended IPM to analyse the dynamics of small populations in stochastic environments, but this model has not yet been used to conduct population viability analyses. Here, we used the extended IPM to analyse the stochastic dynamics of IPM of small size-structured populations in one plant and one animal species (evening primrose and common lizard) including demographic stochasticity in both cases and environmental stochasticity in the lizard model. We also tested the accuracy of a diffusion approximation of the IPM for the two empirical systems. In both species, the elasticity for λ was higher with respect to parameters linked to body growth and size-dependent reproduction rather than survival. An analytical approach made it possible to quantify demographic and environmental variance to calculate the average stochastic growth rate. Demographic variance was further decomposed to gain insights into the most important size classes and demographic components. A diffusion approximation provided a remarkable fit to the stochastic dynamics and cumulative extinction risk, except for very small populations where stochastic growth rate was biased upward or downward depending on the model. These results confirm that the extended IPM provides a powerful tool to assess the conservation status and compare the stochastic demography of size-structured species, but should be complemented with individual based models to obtain unbiased estimates for very small populations of conservation concern.