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Suboptimal conditions during development can shorten telomeres, the protective DNA caps on the end of chromosomes. Shorter early-life telomere length (TL) can indicate reduced somatic maintenance, leading to lower survival and shorter lifespan. However, despite some clear evidence, not all studies show a relationship between early-life TL and survival or lifespan, which may be due to differences in biology or study design (e.g., survival period measured). In superb fairy-wrens (Malurus cyaneus), we assessed whether early-life TL predicts mortality across different life-history stages (fledgling, juvenile, adult). However, in contrast to a similar study on a congener, early-life TL did not predict mortality across any life stage in this species. We then performed a meta-analysis including 32 effect sizes from 23 studies (15 birds and three mammals) to quantify the effect of early-life TL on mortality whilst taking into consideration potential sources of biological and methodological variation. Overall, the effect of early-life TL on mortality was significant, corresponding to a 15% reduction in mortality risk with each standard deviation increase in TL. However, the effect became weaker when correcting for publication bias. Contrary to our predictions, there was no evidence that effects of early-life TL on mortality varied with species lifespan or the period over which survival was measured. However, negative effects of early-life TL on mortality risk were pervasive throughout life. These results imply that effects of early-life TL on mortality are more likely to be context-dependent than age-dependent, although substantial power and publication bias issues highlight the need for more research.

enThis link goes to a English sectionfrThis link goes to a French section 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 highligh

Thermo-hydroregulation strategies involve concurrent changes in functional traits related to energy, water balance and thermoregulation and play a key role in determining life-history traits and population demography of terrestrial ectotherms. Local thermal and hydric conditions should be important drivers of the geographical variation of thermo-hydroregulation strategies, but we lack studies that examine these changes across climatic gradients in different habitat types. Here, we investigated intraspecific variation of morphology and thermo-hydroregulation traits in the widespread European common lizard (Zootoca vivipara louislantzi) across a multidimensional environmental gradient involving independent variation in air temperature and rainfall and differences in habitat features (access to free-standing water and forest cover). We sampled adult males for morphology, resting metabolic rate, total and cutaneous evaporative water loss and thermal preferences in 15 populations from the rear to the leading edge of the distribution across an elevational gradient ranging from sea level to 1750 m. Besides a decrease in adult body size with increasing environmental temperatures, we found little effect of thermal conditions on thermo-hydroregulation strategies. In particular, relict lowland populations from the warm rear edge showed no specific ecophysiological adaptations. Instead, body mass, body condition and resting metabolic rate were positively associated with a rainfall gradient, while forest cover and water access in the habitat throughout the season also influenced cutaneous evaporative water loss. Our study emphasizes the importance of rainfall and habitat features rather than thermal conditions for geographical variation in lizard morphology and physiology.

Thermoregulation is critical for ectotherms as it allows them to maintain their body temperature close to an optimum for ecological performance. Thermoregulation includes a range of behaviors that aim at regulating body temperature within a range centered around the thermal preference. Thermal preference is typically measured in a thermal gradient in fully-hydrated and post-absorptive animals. Short-term effects of the hydric environment on thermal preferences in such set-ups have been rarely quantified in dry-skinned ectotherms, despite accumulating evidence that dehydration might trade-off with behavioral thermoregulation. Using experiments performed under controlled conditions in climatic chambers, we demonstrate that thermal preferences of a ground-dwelling, actively foraging lizard (Zootoca vivipara) are weakly decreased by a daily restriction in free-standing water availability (less than 0.5 degrees C contrast). The influence of air humidity during the day on thermal preferences depends on time of the day and sex of the lizard, and is generally weaker than those of of free-standing water (less than 1 degrees C contrast). This shows that short-term dehydration can influence, albeit weakly, thermal preferences under some circumstances in this species. Environmental humidity conditions are important methodological factors to consider in the analysis of thermal preferences.

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 degrees 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.

Motivation The understanding of physiological adaptations, of evolutionary radiations and of ecological responses to global change urges for global, comprehensive databases of the functional traits of extant organisms. The ability to maintain an adequate water balance is a critical functional property influencing the resilience of animal species to climate variation. In terrestrial or semi-terrestrial organisms, total water loss includes a significant contribution from evaporative water loss (EWL). The analysis of geographic and phylogenetic variation in EWL rates must however account for differences in methods and potential confounding factors, which influence standard measures of whole-organism water loss. We compiled the global and standardized SquamEWL database of total, respiratory and cutaneous EWL for 325 species and subspecies of squamate reptiles (793 samples and 2,536 estimates) from across the globe. An extensive set of companion data and annotations associated with the EWL measurements of potential value for future investigation, including metabolic rate data, is provided. We present preliminary descriptive statistics for the compiled data, discuss gaps and biases, and identify promising avenues to update, expand and explore this database. Main types of variables contained Standard water loss rates, geographic data, metabolic rates. Spatial location Global. Time period Data were obtained from extant species and were collected between 1945 and 2020. Major taxa Reptilia, Squamata including lizards, snakes and amphisbaenians. Level of measurements Individual samples of animals from the same species, locality, age class and sex category. Software format csv.

Parent-offspring conflicts are widespread given that resources are often limited. Recent evidence has shown that availability of water can trigger such conflict during pregnancy in viviparous squamate species (lizards and snakes) and thus questions the role of water in the evolution of reproductive modes. Here, we examined the impact of water restriction during gravidity in the oviparous form of the bimodal common lizard (Zootoca vivipara), using a protocol previously used on the viviparous form. Females were captured in early gravidity from six populations along a 600 m altitudinal gradient to investigate whether environmental conditions (altitude, water access and temperature) exacerbate responses to water restriction. Females were significantly dehydrated after water restriction, irrespective of their reproductive status (gravid vs. non-reproductive), relative reproductive effort (relative clutch mass), and treatment timing (embryonic development stage). Female dehydration, together with reproductive performance, varied with altitude, probably due to long term acclimation or local adaptation. This moderate water-based intergenerational conflict in gravid females contrasts sharply with previous findings for the viviparous form, with implications to the evolutionary reversion from viviparity to oviparity. It is likely that oviparity constitutes a water-saving reproductive mode which might help mitigate intensive temperature-driven population extinctions at low altitudes.

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 postparturition 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.

Regulation of body temperature is crucial for optimizing physiological performance in ectotherms but imposes constraints in time and energy. Time and energy spent thermoregulating can be reduced through behavioral (e.g., basking adjustments) or biophysical (e.g., heating rate physiology) means. In a heterogeneous environment, we expect thermoregulation costs to vary according to local, climatic conditions and therefore to drive the evolution of both behavioral and biophysical thermoregulation. To date, there are limited data showing that thermal physiological adjustments have a direct relationship to climatic conditions. In this study, we explored the effect of environmental conditions on heating rates in the common lizard (Zootoca vivipara). We sampled lizards from 10 populations in the Massif Central Mountain range of France and measured whether differences in heating rates of individuals correlated with phenotypic traits (i.e., body condition and dorsal darkness) or abiotic factors (temperature and rainfall). Our results show that heat gain is faster for lizards with a higher body condition, but also for individuals from habitats with higher amount of precipitation. Altogether, they demonstrate that environmentally induced constraints can shape biophysical aspects of thermoregulation.

Reproduction involves considerable reorganization in an organism's physiology that incurs potential toxicity for cells (e.g., oxidative stress) and decrease in fitness. This framework has been the cornerstone of the so-called 'oxidative cost of reproduction', a theory that remains controversial and relatively overlooked in non-model ectotherms. Here, we used two complementary approaches in natural and controlled conditions to test whether altered access to climate conditions (water and temperature resources) alters oxidative status and mediates reproductive trade-offs in viviparous populations of the common lizard (Zootoca vivipara). First, we examined whether access to free-standing water and differences in ambient temperature across 12 natural populations could be related to variation in oxidative status, reproductive effort and reproductive success. Second, we determined whether an experimental restriction to water triggers higher oxidative cost of reproduction and correlates with fitness measures (reproductive success, future survival rate and probability of future reproduction). Pregnant females exhibited higher sensitivity than males to natural or experimental limitations in temperature and water access. That is, in restricted environments, pregnant females with higher reproductive effort exhibited stronger oxidative damage despite enhanced non-enzymatic antioxidant capacity. Enhanced antioxidant defensive capacity in pregnant females was positively correlated with higher reproductive success, whereas elevated oxidative damage negatively correlated with offspring annual survival. Altogether, our results revealed a context-dependent oxidative cost of reproduction that was concomitant with a conflict in water demand from offspring. These new insights should be critical for understanding ectotherm responses to heat waves and summer droughts that are increasing in frequency and duration. A free Plain Language Summary can be found within the Supporting Information

Chronic changes in climate conditions may select for acclimation responses in terrestrial animals living in fluctuating environments, and beneficial acclimation responses may be key to the resilience of these species to global changes. Despite evidence that climate warming induces changes in water availability, acclimation responses to water restriction are understudied compared with thermal acclimation. In addition, acclimation responses may involve different modes, paces, and trade-offs between physiological and behavioral traits. Here, we tested the dynamical acclimation responses of a dry-skinned terrestrial ectotherm to chronic water restriction. Yearling common lizards (Zootoca vivipara) were exposed to sublethal water restriction during 2 mo of the summer season in laboratory conditions, then released in outdoor conditions for 10 additional months. Candidate behavioral (exploration, basking, and thermal preferences) and physiological (metabolism at rest and standard water loss rate) traits potentially involved in the acclimation response were measured repeatedly during and after water restriction. We observed a sequential acclimation response in water-restricted animals (yearlings spent less time basking during the first weeks of water deprivation) that was followed by delayed sex-specific physiological consequences of the water restriction during the following months (thermal depression in males and lower standard evaporative water loss rates in females). Despite short-term negative effects of water restriction on body growth, annual growth, survival, and reproduction were not significantly different between water-restricted and control yearlings. This demonstrates that beneficial acclimation responses to water restriction involve both short-term flexible behavioral responses and delayed changes in thermal and water biology traits.

Increased global temperatures have opened previously inhospitable habitats, such as at higher elevations. However, the reduction of oxygen partial pressure with increase in elevation represents an important physiological constraint that may limit colonization of such habitats, even if the thermal niche is appropriate. To test the mechanisms underlying the response to ecologically relevant levels of hypoxia, we performed a translocation experiment with the common wall lizard (Podarcis muralis), a widespread European lizard amenable to establishing populations outside its natural range. We investigated the impacts of hypoxia on the oxygen physiology and reproductive output of gravid common wall lizards and the subsequent development and morphology of their offspring. Lowland females transplanted to high elevations increased their haematocrit and haemoglobin concentration within days and maintained routine metabolism compared with lizards kept at native elevations. However, transplanted lizards suffered from increased reactive oxygen metabolite production near the oviposition date, suggesting a cost of reproduction at high elevation. Transplanted females and females native to different elevations did not differ in reproductive output (clutch size, egg mass, relative clutch mass or embryonic stage at oviposition) or in post-oviposition body condition. Developing embryos reduced heart rates and prolonged incubation times at high elevations within the native range and at extreme high elevations beyond the current range, but this reduced oxygen availability did not affect metabolic rate, hatching success or hatchling size. These results suggest that this opportunistic colonizer is capable of successfully responding to novel environmental constraints in these important life-history stages.

The regulation of body temperature (thermoregulation) and of water balance (defined here as hydroregulation) are key processes underlying ecological and evolutionary responses to climate fluctuations in wild animal populations. In terrestrial (or semiterrestrial) ectotherms, thermoregulation and hydroregulation closely interact and combined temperature and water constraints should directly influence individual performances. Although comparative physiologists traditionally investigate jointly water and temperature regulation, the ecological and evolutionary implications of these coupled processes have so far mostly been studied independently. Here, we revisit the concept of thermo-hydroregulation to address the functional integration of body temperature and water balance regulation in terrestrial ectotherms. We demonstrate how thermo-hydroregulation provides a framework to investigate functional adaptations to joint environmental variation in temperature and water availability, and potential physiological and/or behavioral conflicts between thermoregulation and hydroregulation. We extend the classical cost-benefit model of thermoregulation in ectotherms to highlight the adaptive evolution of optimal thermo-hydroregulation strategies. Critical gaps in the parameterization of this conceptual optimality model and guidelines for future empirical research are discussed. We show that studies of thermo-hydroregulation refine our mechanistic understanding of physiological and behavioral plasticity, and of the fundamental niche of the species. This is illustrated with relevant and recent examples of space use and dispersal, resource-based trade-offs, and life-history tactics in insects, amphibians, and nonavian reptiles.

The evolution of sex determination is complex and yet crucial in our understanding of population stability. In ectotherms, sex determination involves a variety of mechanisms including genetic determination (GSD), environment determination (ESD), but also interactions between the two via sex reversal. In this study, we investigated whether water deprivation during pregnancy could override GSD in two heterogametic squamate reptiles. We demonstrated that water restriction in early gestation induced a male-biased secondary sex ratio in both species, which could be explained by water sex reversal as the more likely mechanism. We further monitored some long-term fitness estimates of offspring, which suggested that water sex determination (WSD) represented a compensatory strategy producing the rarest sex according to Fisher's assumptions of frequency-dependent selection models. This study provides new insights into sex determination modes and calls for a general investigation of mechanisms behind WSD and to examine the evolutionary implications.

High levels of anthropogenic noise produced in urban areas are known to negatively affect wildlife. Although most research has been focused on the disturbances of communication systems, chronic noise exposure can also lead to physiological and behavioural changes that have strong consequences for fitness. For instance, behavioural changes mediated by anthropogenic noise (e.g. quality of parental care) may alter development and could influence nestling phenotype. We tested if nestling metabolism was influence by traffic noise in an urban exploiter, the house sparrow Passer domesticus. We experimentally exposed breeding house sparrows from a rural area to a playback of traffic noise and we examined the impacts of this experimental procedure on metabolic rates and morphology of nestlings. We did not find an effect of traffic noise on the morphology of nestlings. Surprisingly, we found that disturbed nestlings had overall lower metabolic rates and mass-adjusted metabolic rates than undisturbed birds. Our results suggest a specific effect of noise exposure per se, rather than an indirect effect of anthropogenic noise through the quality of parental care. Both the proximate mechanisms and the ultimate consequences of such metabolic changes on nestlings remain unknown and deserve future experimental studies.

The physiological mechanisms underlying the 'cost of reproduction' remain under debate, though oxidative stress has emerged as a potential candidate. The 'oxidative cost of reproduction' has received considerable attention with regards to food and antioxidant availability; however, the limitation of water availability has thus far been neglected. In this study, we experimentally examined the combined effect of pregnancy and water deprivation on oxidative status in a viviparous snake (Vipera aspis), a species naturally exposed to periods of water and food deprivation. We predicted a cumulative effect of pregnancy and dehydration on oxidative stress levels. Our results support the occurrence of an oxidative cost of reproduction as we found higher oxidative damage levels in pregnant females than in non-reproductive individuals, despite an upregulation of antioxidant defences. Surprisingly, water deprivation was associated with an up-regulation of antioxidant defences, and did not increase oxidative damage, either alone or in combination with reproduction.

Circulating glucocorticoid (GC) levels may increase as a result of reproductive effort or in response to unpredictable events. However, GC secretion can vary with the availability of vital trophic resources such as energy. While water represents another critical resource, the impact of water deprivation on GC secretion during reproduction has not yet been thoroughly investigated. Here, we examined the effects of water deprivation on plasma corticosterone (CORT) concentrations of female aspic vipers (Vipera aspis), and determined the impacts of water deprivation on offspring traits. We exposed both pregnant and non-reproductive females to a 20-day water deprivation and compared their pre- and post-deprivation CORT levels with those of control females. At the end of the treatment, only water-deprived pregnant females showed a significant increase in CORT levels. In pregnant females, changes in baseline CORT level were correlated with changes in female hydration state. Changes in baseline CORT levels were also negatively influenced by maternal reproductive effort in pregnant control females, while such a relationship was not apparent in pregnant water-deprived females. Finally, we found that offspring from water-deprived females had higher growth rates than offspring from control females. Offspring growth was also positively correlated with changes in both maternal osmolality and baseline CORT levels. Together, our results suggest that dehydration increases maternal CORT levels, which may subsequently influence offspring development. Further long-term field studies are therefore required to assess whether there is an adaptive significance of this response.