The relationships among species' physiological capacities and the geographical variation of ambient climate are of key importance to understanding the distribution of life on the Earth. Furthermore, predictions of how species will respond to climate change will profit from the explicit consideration of their physiological tolerances. The climatic variability hypothesis, which predicts that climatic tolerances are broader in more variable climates, provides an analytical framework for studying these relationships between physiology and biogeography. However, direct empirical support for the hypothesis is mostly lacking for endotherms, and few studies have tried to integrate physiological data into assessments of species' climatic vulnerability at the global scale. Here, we test the climatic variability hypothesis for endotherms, with a comprehensive dataset on thermal tolerances derived from physiological experiments, and use these data to assess the vulnerability of species to projected climate change. We find the expected relationship between thermal tolerance and ambient climatic variability in birds, but not in mammals-a contrast possibly resulting from different adaptation strategies to ambient climate via behaviour, morphology or physiology. We show that currently most of the species are experiencing ambient temperatures well within their tolerance limits and that in the future many species may be able to tolerate projected temperature increases across significant proportions of their distributions. However, our findings also underline the high vulnerability of tropical regions to changes in temperature and other threats of anthropogenic global changes. Our study demonstrates that a better understanding of the interplay among species' physiology and the geography of climate change will advance assessments of species' vulnerability to climate change.

The extent to which different kinds of organisms have adapted to environmental temperature regimes is central to understanding how they respond to climate change. The Scholander-Irving (S-I) model of heat transfer lays the foundation for explaining how endothermic birds and mammals maintain their high, relatively constant body temperatures in the face of wide variation in environmental temperature. The S-I model shows how body temperature is regulated by balancing the rates of heat production and heat loss. Both rates scale with body size, suggesting that larger animals should be better adapted to cold environments than smaller animals, and vice versa. However, the global distributions of ∼9,000 species of terrestrial birds and mammals show that the entire range of body sizes occurs in nearly all climatic regimes. Using physiological and environmental temperature data for 211 bird and 178 mammal species, we test for mass-independent adaptive changes in two key parameters of the S-I model: basal metabolic rate (BMR) and thermal conductance. We derive an axis of thermal adaptation that is independent of body size, extends the S-I model, and highlights interactions among physiological and morphological traits that allow endotherms to persist in a wide range of temperatures. Our macrophysiological and macroecological analyses support our predictions that shifts in BMR and thermal conductance confer important adaptations to environmental temperature in both birds and mammals.macrophysiology | Bergmann's rule | body size | metabolic rate | thermal conductance A fundamental problem in ecology and biogeography is to elucidate the physiological processes that determine the environmental tolerances and influence the distributions of species. Across their nearly worldwide distributions, endothermic birds and mammals maintain near-constant body temperatures in the face of extreme and fluctuating environmental temperatures. Elucidating the morphological and physiological adaptations that allow species to inhabit such a wide spectrum of thermal environments is important for understanding the distribution of biodiversity and for predicting responses of species to climate change (1, 2).In a seminal paper, Scholander et al. (3) showed how endotherms balance rates of heat production and heat loss so as to maintain a constant body temperature in the face of varying environmental temperatures. The essence of the Scholander-Irving (S-I) model is the equation:where T b is body temperature, T a is ambient temperature, B is the rate of metabolic heat production, and C is the rate of heat loss or thermal conductance (4). For a resting animal, which has minimized heat loss by maximizing insulation and optimizing body posture, C = minimum thermal conductance (C MIN ); B = basal metabolic rate (BMR); and T a = T lc , where T lc is the lower critical temperature or the lower limit of the thermal neutral zone (TNZ).The TNZ is ecologically important because it is the range of environmental temperatures where energy expenditure is minimal; out...

Aim Physiological traits that approximate the fundamental climatic niche – the climatic conditions where a species can survive – are the outcome of adaptation to the environment under historical and current environmental constraints. If a large amount of the variation in physiological traits among species can be explained by their phylogeny rather than by contemporary environmental conditions, this would indicate phylogenetic conservatism in physiological traits, i.e. the tendency of species to retain their ancestral physiology over time. Here, we evaluate the relative contributions of phylogeny and environment to explain the variation in physiological traits of birds and mammals at the global level, as well as separately for tropical versus temperate species. Location Global. Methods We compiled a large data set from the literature, on the thermal traits and basal metabolic rates of 552 endotherms (255 bird and 297 mammal species) as measured in physiological experiments, along with phylogenetic, geographical and climatic data. Our analyses, which were performed separately for birds and mammals, partitioned the variation in comparative physiological data into the relative contributions of phylogenetic and environmental distance matrices. Results Overall, the current environment explained a larger amount of variation in thermal traits among species than the phylogeny. However, we found that phylogeny was much more important than current environment for explaining the variation in physiological traits in the tropics, whereas environment was more important than phylogeny in temperate species. Main conclusions While evidence for phylogenetic conservatism in physiological traits at the global level was weak, results for tropical species suggest phylogenetic conservatism in their physiological traits. These results indicate a stronger tendency in tropical species to retain their ancestral thermal traits, which might in turn imply a lower physiological adaptability of tropical species to ongoing and future climate change.

Aim To understand how climatic conditions influence the geographical distributions of species and their potential responses to climate change, we investigated the relationships between the thermal tolerances of species and the size and limits of their distributions. We tested two hypotheses for endotherms: the climatic variability hypothesis, which predicts increases in range size with increasing breadth of thermal tolerance, and the climatic extreme hypothesis, which predicts that range limits are related to thermal tolerance limits. Furthermore, we tested whether these relationships differ between temperate and tropical areas. Location Global. Time period Present. Major taxa studied Birds and mammals. Methods We compiled data on thermal tolerances that had been measured in physiological experiments for 453 endothermic species, along with information on geographical ranges and climatic conditions. We applied phylogenetic generalized least square regressions to test for relationships between thermal tolerance and (a) range size or limits and (b) breadth and extremes of the climatic conditions that each species experiences across its distribution. Results We found that range size was not related to the breadth thermal tolerance for endotherms. However, the range limits at high latitudes as well as the minimum temperatures experienced by species were closely related to the physiological cold tolerances of species. These relationships were particularly strong in temperate regions, but these patterns were not found in the tropics. Main conclusions Our results are inconsistent with the predictions of the climatic variability hypothesis, but are in line with the predictions of the climatic extreme hypothesis. Furthermore, the factors determining species distributions do not appear to be the same in tropical and temperate regions. Our study emphasizes the need to combine spatially explicit distribution models with information from physiological experiments in order to capture regional differences and improve predictions of the responses of species to climate change.

Mechanistic approaches for predicting the ranges of endotherms are needed to forecast their responses to environmental change. We test whether physiological constraints on maximum metabolic rate and the factor by which endotherms can elevate their metabolism (metabolic expansibility) influence cold range limits for mammal and bird species. We examine metabolic expansibility at the cold range boundary (MECRB) and whether species’ traits can predict variability in MECRB and then use MECRB as an initial approach to project range shifts for 210 mammal and 61 bird species. We find evidence for metabolic constraints: the distributions of metabolic expansibility at the cold range boundary peak at similar values for birds (2.7) and mammals (3.2). The right skewed distributions suggest some species have adapted to elevate or evade metabolic constraints. Mammals exhibit greater skew than birds, consistent with their diverse thermoregulatory adaptations and behaviors. Mammal and bird species that are small and occupy low trophic levels exhibit high levels of MECRB. Mammals with high MECRB tend to hibernate or use torpor. Predicted metabolic rates at the cold range boundaries represent large energetic expenditures (>50% of maximum metabolic rates). We project species to shift their cold range boundaries poleward by an average of 3.9° latitude by 2070 if metabolic constraints remain constant. Our analysis suggests that metabolic constraints provide a viable mechanism for initial projections of the cold range boundaries for endotherms. However, errors and approximations in estimating metabolic constraints (e.g., acclimation responses) and evasion of these constraints (e.g., torpor/hibernation, microclimate selection) highlight the need for more detailed, taxa‐specific mechanistic models. Even coarse considerations of metabolism will likely lead to improved predictions over exclusively considering thermal tolerance for endotherms.

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Active species reintroduction is an important conservation tool when aiming for the restoration of biological communities and ecosystems. The effective monitoring of reintroduction success is a crucial factor in this process. Here, we used a combination of environmental DNA (eDNA) techniques and species distribution models (SDMs) to evaluate the success of recent reintroductions of the freshwater fish Alburnoides bipunctatus in central Germany. We built SDMs without and with eDNA presence data to locate further suitable reintroduction sites and potentially overlooked populations of the species. We successfully detected eDNA of A. bipunctatus at all reintroduction sites, as well as several adjacent sites mostly in downstream direction, which supports the success of reintroduction efforts. eDNA‐based species detection considerably improved SDMs for A. bipunctatus, which allowed to identify species presence in previously unknown localities. Our results confirm the usefulness of eDNA techniques as standard tool to monitor reintroduced fish populations. We propose that combining eDNA with SDMs is a highly effective approach for long‐term monitoring of reintroduction success in aquatic species.

To study the potential effects of climate change on species, one of the most popular approaches are species distribution models (SDMs). However, they usually fail to consider important species‐specific biological traits, such as species’ physiological capacities or dispersal ability. Furthermore, there is consensus that climate change does not influence species distributions in isolation, but together with other anthropogenic impacts such as land‐use change, even though studies investigating the relative impacts of different threats on species and their geographic ranges are still rare. Here we propose a novel integrative approach which produces refined future range projections by combining SDMs based on distribution, climate, and physiological tolerance data with empirical data on dispersal ability as well as current and future land‐use. Range projections based on different combinations of these factors show strong variation in projected range size for our study species Emberiza hortulana. Using climate and physiological data alone, strong range gains are projected. However, when we account for land‐use change and dispersal ability, future range‐gain may even turn into a future range loss. Our study highlights the importance of accounting for biological traits and processes in species distribution models and of considering the additive effects of climate and land‐use change to achieve more reliable range projections. Furthermore, with our approach we present a new tool to assess species’ vulnerability to climate change which can be easily applied to multiple species.