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Lenoir, Jonathan (Ed.)Abstract AimSquamate fitness is affected by body temperature, which in turn is influenced by environmental temperatures and, in many species, by exposure to solar radiation. The biophysical drivers of body temperature have been widely studied, but we lack an integrative synthesis of actual body temperatures experienced in the field, and their relationships to environmental temperatures, across phylogeny, behaviour and climate. LocationGlobal (25 countries on six continents). TaxaSquamates (210 species, representing 25 families). MethodsWe measured the body temperatures of 20,231 individuals of squamates in the field while they were active. We examined how body temperatures vary with substrate and air temperatures across taxa, climates and behaviours (basking and diel activity). ResultsHeliothermic lizards had the highest body temperatures. Their body temperatures were the most weakly correlated with substrate and air temperatures. Body temperatures of non‐heliothermic diurnal lizards were similar to heliotherms in relation to air temperature, but similar to nocturnal species in relation to substrate temperatures. The correlation of body temperature with air and substrate temperatures was stronger in diurnal snakes and non‐heliothermic lizards than in heliotherms. Body‐substrate and body‐air temperature correlations varied with mean annual temperatures in all diurnal squamates, especially in heliotherms. Thermal relations vary with behaviour (heliothermy, nocturnality) in cold climates but converge towards the same relation in warm climates. Non‐heliotherms and nocturnal species body temperatures are better explained by substrate temperature than by air temperature. Body temperature distributions become left‐skewed in warmer‐bodied species, especially in colder climates. Main ConclusionsSquamate body temperatures, their frequency distributions and their relation to environmental temperature, are globally influenced by behavioural and climatic factors. For all temperatures and climates, heliothermic species' body temperatures are consistently higher and more stable than in other species, but in regions with warmer climate these differences become less pronounced. A comparable variation was found in non‐heliotherms, but in not nocturnal species whose body temperatures were similar to air and substrate irrespective of the macroclimatic context.more » « less
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Abstract The Andes, with its diverse topography and climate, is a renowned cradle for adaptive radiation, particularly for vertebrate ectotherms such as lizards. Yet, the role of temperature in promoting physiological specialization in the Andes remains unclear. Aseasonality in the tropics should favour physiological specialization across elevation in lizards, but empirical data are limited and equivocal. Determining how thermal tolerances are geographically and phylogenetically structured is therefore a priority, particularly as environments continue to change rapidly. However, there is a gap in our knowledge of thermal limits of species from the Andes, one of the planet’s most biodiverse regions. Anoles, a diverse lizard group found across thousands of metres of elevation in the Andes, can offer insights into evolutionary adaptations to temperature. This study focused on 14 anole species from two clades (Dactyloa and Draconura) that independently diversified along elevational gradients in the Andes. We measured critical thermal limits (CTmin and CTmax) and found patterns of thermal tolerance specialization across elevation, both among and within species. Patterns of thermal specialization are similar among anole clades, indicating parallel responses to similar environmental pressures. Specifically, high-elevation anoles are more cold tolerant and less heat tolerant than their low-elevation counterparts, rendering thermal tolerance breadths stable across elevation (thermal specialization). Evolutionary rates of physiological traits were similar, reflecting parallel specialization in heat and cold tolerance across elevation. The adaptive radiation of anole lizards reflects physiological specialization across elevation, and the endemism such specialization favours, probably catalysed their remarkable diversity in the tropical Andes.more » « less
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Abstract Colour polymorphic species often exhibit variation in morphology, physiology, and behaviour among morphs. In particular, dominance status may be signalled by the interaction between behaviour and colour morph. Behavioural traits associated with dominance include boldness, exploration, and aggression, which influence access to preferred habitat, territorial defence, and mate acquisition. In ectotherms, the social structure associated with morphs may result in the exploitation of structural niches differing in thermal quality. Hence, social interactions among morphs may generate concordant variation in thermal preference and environmental temperature. However, few studies have assessed thermal preference variation in colour polymorphic species and its covariation with behaviour. Doing so can provide insight into niche specialization and the maintenance of colour polymorphism in populations. Here, we investigated the patterns of covariation in boldness behaviour, exploratory behaviour, and thermal preference in the tree lizard,Urosaurus ornatus. We assessed trait variation between territorial and non‐territorial male morphs and between orange and yellow female morphs. Boldness and exploratory behaviour were repeatable in maleU. ornatusand bolder individuals were significantly more likely to incur tail loss, a potential consequence of bold behaviour. Territorial male morphs were significantly bolder and more exploratory and preferred higher body temperatures with a narrowerTsetthan non‐territorial morphs. Female morphs did not vary in behavioural or thermal traits. This study highlights behavioural mechanisms that underly ecological niche segregation and variable habitat use between morphs in a colour polymorphic species.more » « less
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Synopsis Deserts have always amazed researchers due to their high diversity of habitats, where plant and animal species have been able to adapt and diversify, even when these areas impose several constraints on an organism’s activity patterns. In particular, deserts support several lizard species adapted to the thermal and water restrictions found in such biomes. Although several studies have attempted to understand how lizard species might respond to water deficits or droughts in deserts, few have addressed how these responses might vary along a latitudinal gradient. This raises the question of whether physiological buffering of the organism or the climatic environment affects water loss in lizards. Here, we used six populations of Urosaurus ornatus to test whether water loss is influenced more by the intrinsic physiology of the lizard or by the climatic niche. We found that water loss is primarily influenced by the climatic niche of the lizard. However, future studies should focus on how microclimatic variables can influence water loss in organisms found across large latitudinal gradients.more » « less
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Abstract Mounting evidence has shown that personality and behavioral syndromes have a substantial influence on interspecific interactions and individual fitness. However, the stability of covariation among multiple behavioral traits involved in antipredator responses has seldom been tested. Here, we investigate whether sex, gravidity, and parasite infestations influence the covariation between risk aversion (hiding time within a refuge) and escape response (immobility, escape distance) using a viviparous lizard,Zootoca vivipara, as a model system. Our results demonstrated a correlation between risk‐averse and escape behavior at the among‐individual level, but only in gravid females. We found no significant correlations in either males or neonates. A striking result was the loss of association in postparturition females. This suggests that the “risk‐averse – escape” syndrome is ephemeral and only emerges in response to constraints on locomotion driven by reproductive burden. Moreover, parasites have the potential to dissociate the correlations between risk aversion and escape response in gravid females, yet the causal chain requires further examination. Overall, our findings provide evidence of differences in the association between behaviors within the lifetime of an individual and indicate that individual states, sex, and life stages can together influence the stability of behavioral syndromes.more » « less
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Abstract Lizards engage in push-up displays to signal dominance and to secure access to important resources. The rate and patterns of push-up displays have been shown to vary based on both biotic and abiotic factors. We investigated push-up display rate in tree lizards,Urosaurus ornatus, to determine contributions from potentially conflicting factors including sex, throat colour, microhabitat usage, social context, and thermal traits. We found that display rate was best described by an interaction between microhabitat and body temperature (Tb). The relationship between display rate andTbwas significantly different between three microhabitats: sunny dead trees, the inner branches of trees, and tree trunks. We suggest that this variation in display rate is driven by shifts in microhabitat temperature over the course of the day and spatial and temporal adjustments being made depending on the probabilities of being detected by both conspecifics and predators.more » « less
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Abstract Anthropogenic global warming and deforestation are significant drivers of the global biodiversity crisis. Ectothermic and viviparous animals are especially vulnerable since high environmental temperatures can impair embryonic development, but we lack knowledge about these effects upon Neotropical organisms. Here, we estimate how much of the current area with suitable habitats overlaps with protected areas and model the combined effects of climate change and deforestation on the geographic distribution of the viviparous Neotropical lizardNotomabuya frenata(Scincidae). This species ranges in Brazil, Argentina, Paraguay, and Bolivia. We use environmental and physiological variables (locomotor performance and hours of activity) to predict suitable present and future areas, considering different scenarios of greenhouse gas emissions and deforestation. The most critical predictors of habitat suitability were isothermality (i.e., the ratio between mean diurnal temperature range and annual temperature range), precipitation during winter, and hours of activity under lower thermal extremes. Still, our models predict a contraction of suitable habitats in all future scenarios and the displacement of these areas towards eastern South America. In addition, protected areas are not enough to ensure suitable habitats for this species. Our findings highlight the vulnerability of tropical and viviparous ectotherms and suggest that even widely distributed species, such asN. frenata, may have their conservation compromised shortly due to the low representativeness of their suitable habitats in protected areas combined with the synergistic effects of climate change and deforestation. We stress the need for decision‐makers to consider the impact of range shifts in creating protected areas and managing endangered species.more » « less
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ABSTRACT Nocturnal temperatures are increasing at a pace exceeding diurnal temperatures in most parts of the world. The role of warmer nocturnal temperatures in animal ecology has received scant attention and most studies focus on diurnal or daily descriptors of thermal environments' temporal trends. Yet, available evidence from plant and insect studies suggests that organisms can exhibit contrasting physiological responses to diurnal and nocturnal warming. Limiting studies to diurnal trends can thus result in incomplete and misleading interpretations of the ability of species to cope with global warming. Although they are expected to be impacted by warmer nocturnal temperatures, insufficient data are available regarding the night‐time ecology of vertebrate ectotherms. Here, we illustrate the complex effects of nocturnal warming on squamate reptiles, a keystone group of vertebrate ectotherms. Our review includes discussion of diurnal and nocturnal ectotherms, but we mainly focus on diurnal species for which nocturnal warming affects a period dedicated to physiological recovery, and thus may perturb activity patterns and energy balance. We first summarise the physical consequences of nocturnal warming on habitats used by squamate reptiles. Second, we describe how such changes can alter the energy balance of diurnal species. We illustrate this with empirical data from the asp viper (Vipera aspis) and common wall lizard (Podarcis muralis), two diurnal species found throughout western Europe. Third, we make use of a mechanistic approach based on an energy‐balance model to draw general conclusions about the effects of nocturnal temperatures. Fourth, we examine how warmer nights may affect squamates over their lifetime, with potential consequences on individual fitness and population dynamics. We review quantitative evidence for such lifetime effects using recent data derived from a range of studies on the European common lizard (Zootoca vivipara). Finally, we consider the broader eco‐evolutionary ramifications of nocturnal warming and highlight several research questions that require future attention. Our work emphasises the importance of considering the joint influence of diurnal and nocturnal warming on the responses of vertebrate ectotherms to climate warming.more » « less
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Abstract Climate‐modulated parasitism is driven by a range of factors, yet the spatial and temporal variability of this relationship has received scant attention in wild vertebrate hosts. Moreover, most prior studies overlooked the intraspecific differences across host morphotypes, which impedes a full understanding of the climate–parasitism relationship. In the common lizard (Zootoca vivipara), females exhibit three colour morphs: yellow (Y‐females), orange (O‐females) and mixed (mixture of yellow and orange, M‐females).Zootoca viviparais also infested with an ectoparasite (Ophionyssusmites). We therefore used this model system to examine the intraspecific response of hosts to parasitism under climate change. We found infestation probability to differ across colour morphs at both spatial (10 sites) and temporal (20 years) scales: M‐females had lower parasite infestations than Y‐ and O‐females at lower temperatures, but became more susceptible to parasites as temperature increased. The advantage of M‐females at low temperatures was counterbalanced by their higher mortality rates thereafter, which suggests a morph‐dependent trade‐off between resistance to parasites and host survival. Furthermore, significant interactions between colour morphs and temperature indicate that the relationship between parasite infestations and climate warming was contingent on host morphotypes. Parasite infestations increased with temperature for most morphs, but displayed morph‐specific rates. Finally, infested M‐females had higher reductions in survival rates than infested Y‐ or O‐females, which implies a potential loss of intraspecific diversity within populations as parasitism and temperatures rise. Overall, we found parasitism increases with warming temperatures, but this relationship is modulated by host morphotypes and an interaction with temperature. We suggest that epidemiological models incorporate intraspecific diversity within species for better understanding the dynamics of wildlife diseases under climate warming.more » « less
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Abstract 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 lizardsZootoca viviparafrom 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.more » « less
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