Thermal Quality Of Habitat Research Articles

Alpine viper in changing climate: thermal ecology and prospects of a cold-adapted reptile in the warming Mediterranean

In a rapidly changing thermal environment, reptiles are primarily dependent on in situ adaptation because of their limited ability to disperse and the restricted opportunity to shift their ranges. However, the rapid pace of climate change may surpass these adaptation capabilities or elevate energy expenditures. Therefore, understanding the variability in thermal traits at both individual and population scales is crucial, offering insights into reptiles' vulnerability to climate change. We studied the thermal ecology of the endangered Greek meadow viper (Viperagraeca), an endemic venomous snake of fragmented alpine-subalpine meadows above 1600 m of the Pindos mountain range in Greece and Albania, to assess its susceptibility to anticipated changes in the alpine thermal environment. We measured preferred body temperature in artificial thermal gradient, field body temperatures of 74 individuals in five populations encompassing the entire geographic range of the species, and collected data on the available of temperatures for thermoregulation. We found that the preferred body temperature (Tp) differed only between the northernmost and the southernmost populations and increased with female body size but did not depend on sex or the gravidity status of females. Tp increased with latitude but was unaffected by the phylogenetic position of the populations. We also found high accuracy of thermoregulation in V.graeca populations and variation in the thermal quality of habitats throughout the range. The overall effectiveness of thermoregulation was high, indicating that V.graeca successfully achieves its target temperatures and exploits the thermal landscape. Current climatic conditions limit the activity period by an estimated 1278 h per year, which is expected to increase considerably under future climate scenarios. Restricted time available for thermoregulation, foraging and reproduction will represent a serious threat to the fitness of individuals and the persistence of populations in addition to habitat loss due to mining, tourism or skiing and habitat degradation due to overgrazing in the shrinking mountaintop habitats of V.graeca.

Unoccupied aerial vehicles as a tool to map lizard operative temperature in tropical environments

AbstractTo understand how ectotherms will respond to warming temperatures, we require information on thermal habitat quality at spatial resolutions and extents relevant to the organism. Measuring thermal habitat quality is either limited to small spatial extents, such as with ground‐based 3D operative temperature (Te) replicas, representing the temperature of the animal at equilibrium with its environment, or is based on microclimate derived from physical models that use land cover variables and downscale coarse climate data. We draw on aspects of both these approaches and test the ability of unoccupied aerial vehicle (UAV) data (optical RGB) to predict fine‐scale heterogeneity in sub‐canopy lizard (Anolis bicaorum) Te in tropical forest using random forest models. Anolis bicaorum is an endemic, critically endangered, species, facing significant threats of habitat loss and degradation, and work was conducted as part of a larger project. Our findings indicate that a model incorporating solely air temperature, measured at the centre of the 20 × 20 m plot, and ground‐based leaf area index (LAI) measurements, measured at directly above the 3D replica, predicted Te well. However, a model with air temperature and UAV‐derived canopy metrics performed slightly better with the added advantage of enabling the mapping of Te with continuous spatial extent at high spatial resolutions, across the whole of the UAV orthomosaic, allowing us to capture and map Te across the whole of the survey plot, rather than purely at 3D replica locations. Our work provides a feasible workflow to map sub‐canopy lizard Te in tropical environments at spatial scales relevant to the organism, and across continuous areas. This can be applied to other species and can represent species within the same community that have evolved a similar thermal niche. Such methods will be imperative in risk modelling of such species to anthropogenic land cover and climate change.

Living on the edge: thermoregulation at the niche margin in the Bedriaga’s rock lizard

Abstract Thermoregulation is essential for ectotherms but its relative cost, especially under ongoing climate change, depends on the thermal quality of habitats. Populations at the warm margin of a thermal niche could be negatively affected by environmental temperatures that approach the limits of a species’ thermoregulation capacity. This study aims to define the thermal niche of the Bedriaga’s rock lizard, Archaeolacerta bedriagae, a rock-dwelling species endemic to the Corsica and Sardinia islands (western Mediterranean Sea), and to investigate its thermoregulation effectiveness at the warm edge of its niche. We collected data on climate, body temperature, and microhabitat temperature throughout the species’ range to characterize its thermal niche. We found that A. bedriagae does not occupy the entire climatic space available across its distribution range; rather, it selects temperate climates. Remarkably, thermoregulation effort increases when the habitat thermal quality decreases towards warmer sites. Populations at the warm edge of the thermal niche show the best thermoregulation effectiveness, but they are also more sensitive to the effects of climate change as they may already be at (or beyond) the species’ maximum thermal capacity under the current conditions. We observed such a pattern at the extreme hot side of the thermal niche. This study provides key information on the thermoregulatory response of A. bedriagae to ongoing climate change that can be useful to identify populations facing a higher extinction risk either currently or in the near future.

Small Island Effects on the Thermal Biology of the Endemic Mediterranean Lizard Podarcis gaigeae.

Ectotherms are vastly affected by climatic conditions as they rely on external sources of heat to regulate their body temperature, and changes in their habitat thermal quality could seriously affect their overall biology. To overcome the problems of a thermally unfavorable habitat, lizards need to either adjust their thermoregulatory behavior or respond to directional selection and shift their preferred body temperatures. To assess the impact of habitat thermal quality on the thermoregulatory profile, we studied multiple islet and 'mainland' populations of the Skyros wall lizard Podarcis gaigeae, an endemic lacertid to Skyros Archipelago, Greece. We evaluated the effectiveness of thermoregulation (E) using the three main thermal parameters: body (Tb), operative (Te), and preferred (Tpref) temperatures. We first hypothesized that the spatial homogeneity, the scarcity of thermal shelters, and the exposure to higher winds on islets would result in more demanding climate conditions. Second, we anticipated that islet lizards would achieve higher E in response to the lower thermal quality therein. As hypothesized, thermal parameters differed between populations but not in the expected manner. Skyros 'mainland' habitats reached higher temperatures, had more intense fluctuations, and were of lower thermal quality. As a result, lizards showed higher accuracy, precision, and effectiveness of thermoregulation. Noteworthy, we found that lizards from different populations have shifted their thermal profile and preferred body temperatures to cope with the particular conditions prevailing in their habitats. The latter supports the labile view on the evolution of thermoregulation.

Negative effects of the spatial clumping of thermal resources on lizard thermoregulation in a fragmented habitat

In ecosystems threatened by the expansion of croplands, habitat fragmentation and climate change, two of the main extinction drivers, may have thermoregulation-mediated interacting effects on demographic trends of terrestrial ectotherms. We studied the thermal biology of a metapopulation of the widespread Mediterranean lacertid Psammodromus algirus in ten fragments of evergreen or deciduous oak forests interspersed among cereal fields. We obtained thermoregulation statistics (selected temperature range, body and operative temperatures, thermal quality of the habitat, and precision, accuracy, and effectiveness of thermoregulation) that could be compared among fragments and with conspecific populations living in unfragmented habitat. We also measured the selection (use vs. availability) and spatial distribution of sunlit and shaded patches used for behavioral thermoregulation in fragments, and we estimated operative temperatures and thermal habitat quality in the agricultural matrix surrounding the fragments. Variation of the thermal environment was much larger within fragments than among them, and thermoregulation was accurate, precise, and efficient throughout the fragmented landscape; its effectiveness was similar to that of previously studied unfragmented populations. The average distance between sunlit and shaded patches was shorter in deciduous than in evergreen fragments, producing a more clumped distribution of the mosaic of thermal resources. Consequently, in evergreen habitat the cost of thermoregulation was higher, because lizards were more selective in their choice of sunlit sites (i.e. they used sunlit patches closer to shade and refuge than expected at random, and the extent of such selection was larger than at deciduous habitat). Temperatures available in croplands were too high to allow lizard dispersal, at least in the post-breeding season. This result confirms the role of croplands as a thermal barrier that promotes inbreeding and associated fitness losses in isolated fragments, and it forecasts a dark future for populations of forest lizards in agricultural landscapes under the combined effects of habitat fragmentation and global warming.

Preliminary Antecedents on the Selected Temperature (Tsel) of Rhinella dpinulosa Wiegmann 1834 (Anura: Bufonidae) from the Andean Environments of Chile

The ambient temperature is a decisive factor in many activities of poikilothermic organisms, since it influences the correct development of their vital processes, which implies the existence of an optimal temperature range. The present study describes selected temperatures (Tsel) of R. spinulosa in adult individuals. For this purpose, adults of this species were acclimatized at 10°C and then subjected to a thermal gradient was used in the laboratory. The mean Tsel in the studied individuals was 23.38±2.65°C, with a range from 17.73°C to 28.93°C. Future surveys are suggested to determine the habitat thermal quality and thermoregulatory precision.

Seasonal and elevational variation in thermal ecology of the crevice-dwelling knob-scaled lizard Xenosaurus fractus from central-eastern Mexico

There is strong covariation between the thermal physiology of ectothermic animals and their thermal environment. Spatial and temporal differences in the thermal environment across a species' range may result in changes in thermal preferences between populations of that species. Alternatively, thermoregulatory-based microhabitat selection can allow individuals to maintain similar body temperatures across a broad thermal gradient. Which strategy a species adopts is often dependent on taxon-specific levels of physiological conservatism or ecological context. Identifying which strategies species use in response to spatial and temporal variation in environmental temperatures requires empirical evidence, which then can support predictions as to how a species might respond to a changing climate. Here we present findings of our analyses of the thermal quality, thermoregulatory accuracy and efficiency for the lizard, Xenosaurus fractus, across an elevation-thermal gradient and over the temporal thermal variation associated with seasonal changes. Xenosaurus fractus is a strict crevice-dweller, a habitat that can buffer this lizard from extreme temperatures and is a thermal conformer (body temperatures reflect air and substrate temperatures). We found populations of this species differed in their thermal preferences along an elevation gradient and between seasons. Specifically, we found that habitat thermal quality, thermoregulatory accuracy and efficiency (all measures of how well the lizards' body temperatures compared to their preferred body temperatures) varied along thermal gradients and with season. Our findings indicate that this species has adapted to local conditions and shows seasonal flexibility in those spatial adaptations. Along with their strict crevice-dwelling habitat, these adaptations may provide some protection against a warming climate.

Habitat thermal quality for Gopherus evgoodei in tropical deciduous forest and consequences of habitat modification by buffelgrass

Tortoises of the genus Gopherus evolved in North America and have survived major environmental challenges in the past 40 million years. However, this genus now faces multiple anthropogenic threats, such as the introduction of invasive plant species. Buffelgrass (Cenchrus ciliaris) is considered one of the greatest threats to arid and tropical ecosystems, where gopher tortoises inhabit, because the grass displaces native flora and fauna. Modification of the environment as a result of this invasive plant portends an alteration of the available thermal landscape. The aim of this paper is twofold: 1) to evaluate the thermal quality of the primary habitat of Gopherus evgoodei (tropical deciduous forest [TDF], and 2) determine the potential thermal changes due to habitat modification by buffelgrass. First, we obtained data on body temperature of active tortoises in semi-captivity. Second, we measured the operative environmental temperature during 5 years at three sites south of Sonora, Mexico that support G. evgoodei: a) a pristine TDF (Conserved-TDF); b) a forest patch surrounded by introduced buffelgrass pasture (Partial-TDF); and c) an introduced buffelgrass pasture area (Buffel-Pasture). Our results demonstrate that the intact microhabitats within the TDF provide G. evgoodei with high thermal quality at both spatial and temporal scales. However modified habitat by buffelgrass had higher operative temperatures for G. evgoodei than TDF. The thermal quality of the sites disturbed with buffelgrass can exceed the thermal requirements of G. evgoodei by up to 25 °C. Finally, we discussed potential collateral effects of habitat modification by invasion by buffelgrass.

Riparian Land Cover, Water Temperature Variability, and Thermal Stress for Aquatic Species in Urban Streams

Thermal regime warming and increased variability can result in human developed watersheds due to runoff over impervious surfaces and influence of stormwater pipes. This study quantified relationships between tree canopy, impervious surface, and water temperature in stream sites with 4 to 62% impervious land cover in their “loggersheds” to predict water temperature metrics relevant to aquatic species thermal stress thresholds. This study identified significant (≥0.7, p < 0.05) negative correlations between water temperature and percent tree canopy in the 5 m riparian area and positive correlations between water temperature and total length of stormwater pipe in the loggershed. Mixed-effects models predicted that tree canopy cover in the 5 m riparian area would reduce water temperatures 0.01 to 6 °C and total length of stormwater pipes in the loggershed would increase water temperatures 0.01 to 2.6 °C. To our knowledge, this is the first time that the relationship between stormwater pipes and water temperature metrics has been explored to better understand thermal dynamics in urban watersheds. The results highlight important aspects of thermal habitat quality and water temperature variability for aquatic species living in urban streams based on thermal thresholds relevant to species metabolism, growth, and life history.

Thermal ecology of a thermophilic lizard Callisaurus draconoides through a latitudinal gradient

Desert squamates need to regulate their body temperature within a specific range in response to extreme temperatures. Variation of thermal resources through altitudinal and latitudinal gradients influences the thermal physiological traits and thermoregulation of organisms. In some species, the physiological requirements between populations can vary correlated with environmental conditions. In other species, the physiological parameters may remain similar across geographic gradients. In this study, we compared the thermal requirements and thermoregulatory strategies of a lizard (Callisaurus draconoides) that inhabits contrasting desert ecoregions across a latitudinal gradient. To assess thermoregulation of C. draconoides, we measured their field body temperature during periods of activity, the preferred body temperature in a laboratory thermal gradient and the environmental operative temperature in three contrasting sites. We also analyzed and compared three thermoregulatory indices (accuracy and effectiveness of thermoregulation and habitat thermal quality) for each population. Our results suggest that C. draconoides maintains similar thermal requirements and tolerances across its distribution range. Furthermore, their thermoregulation was accurate and effective in low thermal quality environments. Therefore, the thermoregulation in C. draconoides is essential to buffer extreme environmental conditions and avoid the deadly consequences of overheating in each ecoregion along the Baja California peninsula.

Disentangling controls on animal abundance: Prey availability, thermal habitat, and microhabitat structure.

The question of what controls animal abundance has always been fundamental to ecology, but given rapid environmental change, understanding the drivers and mechanisms governing abundance is more important than ever. Here, we determine how multidimensional environments and niches interact to determine population abundance along a tropical habitat gradient. Focusing on the endemic lizard Anolis bicaorum on the island of Utila (Honduras), we evaluate direct and indirect effects of three interacting niche axes on abundance: thermal habitat quality, structural habitat quality, and prey availability. We measured A. bicaorum abundance across a series of thirteen plots and used N‐mixture models and path analysis to disentangle direct and indirect effects of these factors. Results showed that thermal habitat quality and prey biomass both had positive direct effects on anole abundance. However, thermal habitat quality also influenced prey biomass, leading to a strong indirect effect on abundance. Thermal habitat quality was primarily a function of canopy density, measured as leaf area index (LAI). Despite having little direct effect on abundance, LAI had a strong overall effect mediated by thermal quality and prey biomass. Our results demonstrate the role of multidimensional environments and niche interactions in determining animal abundance and highlight the need to consider interactions between thermal niches and trophic interactions to understand variation in abundance, rather than focusing solely on changes in the physical environment.

Thermoregulatory strategies of three reclusive lizards (genus Xantusia) from the Baja California peninsula, Mexico, under current and future microenvironmental temperatures.

The thermal quality of the habitat is key for the regulation of body temperature in terrestrial ectotherms and, therefore, permits them to carry out their fundamental biological activities. In thermally heterogeneous environments, ectotherms might follow different behavioral or physiological strategies to maintain their body temperature within biologically adequate boundaries, for which they depend on microhabitat selection. These aspects are, thus, relevant in the context of habitat degradation and land-use change. In this study, we characterized the thermal ecology of three lizard species (genus Xantusia) that differ in microhabitat use along the Baja California peninsula, Mexico. We made three predictions: (1) the three species will follow different thermoregulatory strategies according to habitat thermal quality; (2) the thermal requirements and tolerances of these species will match the environmental or microenvironmental thermal conditions; and (3) due to their habitat and range restriction, the species studied will be highly vulnerable to climate change. Our results indicate the existence of thermoregulatory mechanisms in Xantusia to face thermal heterogeneity, including behavioral thermoregulation by choosing different microhabitats, shifts in activity periods, and adaptation to particular high thermal quality microhabitats. Furthermore, despite their association to specific microhabitats and specialized physiology, the studied species will not be adversely affected by climate change, as the increased microenvironmental temperatures will lead to a higher habitat thermal quality and lower costs of thermoregulation. However, we do not discard other indirect adverse effects of climate change not considered in this study.

Thermoregulation in a saxicolous population of the lizard Urosaurus ornatus from the northern Chihuahuan Desert, Mexico

Abstract Environmental temperatures often regulate the activity and physiological processes of ectotherms. Because environmental temperatures can vary significantly among seasons, lizards exposed to different thermal conditions in different months could differ in how they thermoregulate or behave. Here, we present a combination of field, laboratory, and modeling approaches to examine thermoregulation, habitat thermal quality, and hours of thermal restriction on activity in a saxicolous population of Urosaurus ornatus in two thermally contrasting months (June and October) in a micro-insular mountain system in the northern Chihuahuan Desert in Mexico. Both active and preferred body temperatures did not vary between months. In this population, U. ornatus is an active thermoregulator with highly accurate and efficient thermoregulation despite the thermal quality in both months. However, during the breeding season (June) activity is restricted (i.e., high number of hours of restriction) compared to the non-breeding season (October). Therefore, our results suggest that this saxicolous population of U. ornatus could be threatened by global climate change and it is essential to determine a conservation strategy for this population.

Tree Removal Likely Improves Thermal Quality of Basking Sites for an Imperiled Timber Rattlesnake Population at the Northern Edge of Its Range

Lack of natural disturbance in many forests in northeastern North America has resulted in increased canopy cover and a reduction of open-habitat patches. Crotalus horridus (Timber Rattlesnake) in this region depends upon open basking and gestation sites, and its populations have declined markedly. We evaluated the utility of mechanical tree removal to restore potential basking habitat for a highly imperiled Timber Rattlesnake population in New Hampshire. We used a before–after, control–impact study to test for the effects of tree removal on environmental temperature and Timber Rattlesnake and other wildlife use of 3 potential basking sites. Mean temperatures were significantly higher and more variable post-treatment, while minimum temperatures were relatively unaffected. The number of temperature readings within the span of selected body temperatures of gravid female Timber Rattlesnakes (25.2–31.7 °C) increased post-treatment. Using time-lapse cameras, we detected rattlesnakes during 5 days at post-treatment sites but on only 1 day at a control site. Detections of potential rattlesnake prey species did not change following treatment, and few potential rattlesnake predators were detected pre- or post-treatment. Our results suggest that tree removal around potential basking sites likely improves thermal habitat quality for Timber Rattlesnakes.

Impact of natural resource extraction on thermal properties of wood turtle (Glyptemys insculpta) habitat.

One of the main threats to freshwater turtle populations is habitat destruction, which occurs as a result of human activities such as infrastructure development, forestry, aggregate extraction, and agriculture. However, the impacts of these activities on thermoregulatory opportunities for turtles are not well understood. We examined the impacts of forestry and aggregate extraction on thermal characteristics of wood turtle (Glyptemys insculpta) habitat in the Sudbury District, Ontario, by measuring the differences in absolute temperature, thermal landscape structure, and habitat thermal quality among relatively undisturbed sites (N = 2), harvested forestry sites (N = 2), and aggregate pits (N = 2) in 2015. We also tested the potential use of the thermal landscape concept as a predictor of habitat thermal quality. Undisturbed habitats were of higher thermal quality than impacted sites in terms of temperatures experienced in situ by wild wood turtles in their natural home ranges, and never reached temperatures that would be injurious to turtles (i.e., below CTMIN or above CTMAX). The undisturbed sites were of lower quality in terms of optimal temperatures (i.e., Tset), and were generally cooler and their temperatures less variable than those in impacted habitats. The thermal landscape concept provided a useful predictor of habitat thermal quality when the influence of time of day was factored into the predictive model. Our data are important to the conservation and management of wood turtles because they provide a preliminary quantification of the thermal impacts of natural resource extraction on the habitat of an endangered species, and can guide the development of mitigation and rehabilitation plans by providing measures of, and targets for, thermal habitat quality.

Broad seasonal changes in thermoregulation of Podarcis lilfordi (Squamata, Lacertidae) at Binicodrell islet (Menorca, Spain)

Most lizards maintain quite constant body temperatures by behavioural means. Seasonal variations of environmental factors, such as temperature, sunlight exposure and wind intensity, influence lizard thermoregulatory abilities. Understanding how seasonal environmental shifts influence lizards’ thermoregulation helps us to know how they deal behaviourally with environmental changes, in general. We examined seasonal shifts (spring vs. summer) in behavioural thermoregulation in Podarcislilfordi from Binicodrell islet (Menorca, Spain). Operative temperatures varied between microhabitats and seasons, being lower in spring than in summer, regardless of sunlight exposure. Lizard body temperatures were also lower in spring than in summer. Lizards used sunny microhabitats more frequently in spring and shaded areas in summer. Habitat thermal quality was similar during both seasons, but lizards thermoregulated less accurately in spring than in summer. Thermoregulatory effectiveness was low in spring (0.28) and moderate in summer (0.76). In comparison with previously published results, our findings showed the marked seasonal variation in the effectiveness of thermoregulation amongst island populations, which should be considered in future comparative studies.

Rethinking the scale and formulation of indices assessing organism vulnerability to warmer habitats

Ecologists often use indices or proxies to communicate complex ecological entities. Indices commonly known as thermal safety margin, habitat thermal quality and hours of restriction describe species’ vulnerability to climate change by comparing organisms’ thermal limits or preferences to available habitat temperatures. Ready access to temperature data, from global gridded datasets or limited in situ measurements, has made these indices popular for vulnerability assessments across taxonomic groups and regions. However, such coarse descriptions of thermal landscape mask the spatio‐temporal heterogeneity that organisms experience, compromising the value of these indices. Full understanding of how scale affects index estimates is lacking, leaving ecologists and conservation managers with little guidance for applying or interpreting indices. Here, we show that incomplete temperature sampling, in space or time, provides erroneous assessments of vulnerability. Gradually sub‐sampling a long‐term, fine‐scale dataset of operative environmental temperature altered the index estimates for a lizard. Uncertainty associated with the selection of data increased with coarser scales, often leading to contrasting interpretations about the species’ vulnerability to climate change when different data subsets were used. Compressing the environmental temperature data into central or extreme tendencies, as traditionally done to compute these indices, further masked the thermal variation that animals exploit to buffer warming. We suggest the use of improved index formulations that better describe temperature availability at scales that are appropriate to the study organism.

Habitat management alters thermal opportunity

AbstractEctotherms engage in behavioral thermoregulation to optimize body temperatures, however, thermoregulatory effort varies across species, over time, and among habitats. Classic cost‐benefit models of ectothermic thermoregulation postulate that ectotherms should increase thermoregulatory effort when the benefits of thermoregulatory behavior outweigh the costs (Huey and Slatkin, 1976). However, alteration of habitat via land management may lead to unforeseen shifts in microclimate that change associated costs and benefits. In light of anthropogenic impacts on natural habitats, thermoregulatory effort should be examined in cases where land management results in environmental temperatures that deviate from the preferred temperature range of a given species.We examined how habitat management alters microclimates and influences activity time, thermoregulatory behavior, and thermal physiology in a lizard (Sceloporus woodi) whose range has been significantly reduced due to habitat loss. We compared populations from two ecologically distinct habitats: longleaf pine and sand pine scrub, which have very different management histories. Daily surveys were conducted in each habitat type while environmental temperatures were recorded simultaneously with biophysical models.Thermal performance curves differed between populations such that physiological performance was optimized in each environment. Longleaf pine habitat had greater thermal opportunity because cooler microclimates were available on arboreal perches. In longleaf pine habitat, lizards were able to stay active longer and thermoregulate more effectively. Regardless of habitat, thermoregulatory effort ofS. woodiincreased when thermal quality of habitat was low. However, when unfavorably high temperatures were experienced, the thermoregulatory effort increased significantly.Our results demonstrate that ectotherms can afford to imprecisely thermoregulate when habitat thermal quality is poor due to cooler environmental temperatures. However under unfavorably hot environmental temperatures, lizards must precisely regulate body temperature in spite of any costs. These results further demonstrate that habitat management has major implications on species‐specific thermal opportunity, thermoregulatory behavior, and thermal physiology, such that shifts in thermoregulatory behavior may lead to higher energetic costs, enhanced predation risk, and constrained activity times.Aplain language summaryis available for this article.

Thermoregulatory challenges in the habitat of the world's smallest tortoise, Chersobius signatus

Ectotherms have various means of dealing with low environmental temperatures, but relatively few species have been rigorously investigated. Consequently, we have little information to predict how ectotherm populations might respond to global temperature changes. Tortoises from temperate and subtropical regions often overcome periodically cool conditions by hibernation, but speckled dwarf tortoises (Chersobius signatus) need to remain active to exploit ephemeral resources in their arid winter-rainfall habitat. This study investigated how dwarf tortoises cope with low temperatures in winter and spring, by measuring thermal habitat quality and thermoregulation based on differently-sized operative temperature models in sun, shade, and in deep crevices. Investigations continued in summer and autumn to obtain a year-round picture of thermoregulatory challenges. Although large models (i.e., larger than dwarf tortoises) were expected to have lower operative temperatures than smaller models, due to the former's larger thermal inertia, all model sizes had similar temperatures. Hence, the species’ small body size does not appear constrained by obtainable body temperatures in cool seasons. Nevertheless, low operative temperatures in winter posed a challenge for the tortoises, which reached their field-preferred body temperature for an average of only 0.8−0.9h per day. Moreover, a low thermoregulation effectiveness suggested that tortoises traded-off physiological benefits of favourable body temperatures against predation risk. Spring and autumn provided higher temperatures, but summer caused the greatest thermoregulatory challenge. Although summer body temperatures were closer to field-preferred body temperature than in any other season, tortoises required rock crevices to avoid overheating. The small size of dwarf tortoises might help them utilise crevices. In summer, maximum operative temperatures in crevices were similar to field-preferred body temperature, indicating that an increase in environmental temperatures might be detrimental to dwarf tortoises. In light of projected temperature rises, future studies should assess if dwarf tortoises can cope with higher environmental temperatures in summer.

Natural selection on thermal preference, critical thermal maxima and locomotor performance.

Climate change is resulting in a radical transformation of the thermal quality of habitats across the globe. Whereas species have altered their distributions to cope with changing environments, the evidence for adaptation in response to rising temperatures is limited. However, to determine the potential of adaptation in response to thermal variation, we need estimates of the magnitude and direction of natural selection on traits that are assumed to increase persistence in warmer environments. Most inferences regarding physiological adaptation are based on interspecific analyses, and those of selection on thermal traits are scarce. Here, we estimate natural selection on major thermal traits used to assess the vulnerability of ectothermic organisms to altered thermal niches. We detected significant directional selection favouring lizards with higher thermal preferences and faster sprint performance at their optimal temperature. Our analyses also revealed correlational selection between thermal preference and critical thermal maxima, where individuals that preferred warmer body temperatures with cooler critical thermal maxima were favoured by selection. Recent published estimates of heritability for thermal traits suggest that, in concert with the strong selective pressures we demonstrate here, evolutionary adaptation may promote long-term persistence of ectotherms in altered thermal environments.