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1. Current studies indicate that estimates of thermal tolerance limits in ectotherms depend on the experimental protocol used, with slower and presumably more ecologically relevant rates of warming negatively affecting the upper thermal limits (CTmax). Recent empirical evidence also gives credence to earlier speculations suggesting that estimates of heritability could drop with slower heating rates. 2. Using published data from the fruit fly Drosophila melanogaster, we show that empirical patterns can be explained if flies' physical condition decreases with experimental time in thermal tolerance assays. This problem could even overshadow potential benefits of thermal acclimation, also suggesting that a drop in CTmax with slower heating rates does not necessarily rule out beneficial acclimatory responses. 3. Numerical results from a simple illustrative model show that no clear conclusions can be obtained on how the phenotypic variance in CTmax will be affected with different rates of thermal change. Conversely, the genetic variance and estimated heritabilities are expected to drop with slower heating rates, hence ramping rates in experiments aiming to study the evolutionary potential of thermal tolerance to respond to global warming should be as fast as possible (within the range in which measurement accuracy and physical condition are not affected). 4. Measurements under ecologically realistic warming rates should also consider the impact of other physiological and behavioural strategies that might partly compensate the negative effects of slow heating rates. However, there are situations in which slow heating rates closely mimic natural conditions, as those encountered by some aquatic ectotherms. These heating rates may be an issue of major concern in these species, given its negative impact on CTmax and its adaptive potential.

The climate variability hypothesis posits that increased environmental thermal variation should select for thermal generalists, while stable environments should favor thermal specialists. This hypothesis has been tested on large spatial scales, such as latitude and elevation, but less so on smaller scales reflective of the experienced microclimate. Here, we estimated thermal tolerance limits of 75 species of amphibian tadpoles from an aseasonal tropical mountain range of the Ecuadorian Andes, distributed along a 3500 m elevational range, to test the climatic variability hypothesis at a large (elevation) and a small (microhabitat) scale. We show how species from less variable thermal habitats, such as lowlands and those restricted to streams, exhibit narrower thermal tolerance breadths than highland and pond-dwelling species respectively. Interestingly, while broader thermal tolerance breadths at large scales are driven by higher cold tolerance variation (heat-invariant hypothesis), at local scales they are driven by higher heat tolerance variation. This contrasting pattern may result from divergent selection on both thermal limits to face environmental thermal extremes at different scales. Specifically, within the same elevational window, exposure to extreme maximum temperatures could be avoided through habitat shifts from temporary ponds to permanent ponds or streams, while minimum peak temperatures remained invariable between habitats but steadily decreased with elevation. Therefore an understanding of the effects of habitat conversion is crucial for future research on resilience to climate change.

Variation and population structure play key roles in the speciation process, but adaptive intraspecific genetic variation is commonly ignored when forecasting species niches. Amphibians serve as excellent models for testing how climate and local adaptations shape species distributions due to physiological and dispersal constraints and long generational times. In this study, we analysed the climatic factors driving the evolution of the genus Alytes at inter- and intraspecific levels that may limit realized niches. We tested for both differences among the five recognized species and among intraspecific clades for three of the species ( Alytes obstetricans , A. cisternasii , and A. dickhilleni ). We employed ecological niche models with an ordination approach to perform niche overlap analyses and test hypotheses of niche conservatism or divergence. Our results showed strong differences in the environmental variables affecting species climatic requirements. At the interspecific level, tests of equivalence and similarity revealed that sister species were non-identical in their environmental niches, although they neither were entirely dissimilar. This pattern was also consistent at the intraspecific level, with the exception of A. cisternasii, whose clades appeared to have experienced a lower degree of niche divergence than clades of the other species. In conclusion, our results support that Alytes toads, examined at both the intra- and interspecific levels, tend to occupy similar, if not identical, climatic environments.

Climate change and infectious disease by the chytrid fungus Batrachochytrium dendrobatidis (Bd) are major drivers of amphibian extinctions, but the potential interactions of these two factors are not fully understood. Temperature is known to influence (1) the infectivity, pathogenicity and virulence of Bd; (2) host-parasite dynamics, especially when both hosts and parasites are ectothermic organisms exhibiting thermal sensitivities that may or may not differ; and (3) amphibian vulnerability to extinction depending on their heat tolerance, which may decrease with infection. Thus, in a global warming scenario, with rising temperatures and more frequent and extreme weather events, amphibians infected by Bd could be expected to be more vulnerable if temperatures approach their critical thermal maximum (CTmax). However, it is also possible that predicted high temperatures could clear the Bd infection, thus enhancing amphibian survival. We tested these hypotheses by measuring CTmax values of Bd-infected and Bd-free aquatic tadpoles and terrestrial toadlets/juveniles of the common midwife toad (Alytes obstetricans) and examining whether exposure of A. obstetricans individuals to peak temperatures reaching their CTmax clears them from Bd infection. We show that (1) Bd has a wide thermal tolerance range; (2) Bd is capable of altering the thermal physiology of A. obstetricans, which is stage-dependent, lowering CTmax in tadpoles but not in toadlets; and (3) Bd infection is not cleared after exposure of tadpoles or toadlets to CTmax. Living under climatic change with rising temperatures, the effect of Bd infection might tip the balance and lead some already threatened amphibian communities towards extinction.

To forecast biological responses to changing environments, we need to understand how a species's physiology varies through space and time and assess how changes in physiological function due to environmental changes may interact with phenotypic changes caused by other types of environmental variation. Amphibian larvae are well known for expressing environmentally induced phenotypes, but relatively little is known about how these responses might interact with changing temperatures and their thermal physiology. To address this question, we studied the thermal physiology of grey treefrog tadpoles (Hyla versicolor) by determining whether exposures to predator cues and an herbicide (Roundup) can alter their critical maximum temperature (CTmax) and their swimming speed across a range of temperatures, which provides estimates of optimal temperature (Topt) for swimming speed and the shape of the thermal performance curve (TPC). We discovered that predator cues induced a 0.4°C higher CTmax value, whereas the herbicide had no effect. Tadpoles exposed to predator cues or the herbicide swam faster than control tadpoles and the increase in burst speed was higher near Topt. In regard to the shape of the TPC, exposure to predator cues increased Topt by 1.5°C, while exposure to the herbicide marginally lowered Topt by 0.4°C. Combining predator cues and the herbicide produced an intermediate Topt that was 0.5°C higher than the control. To our knowledge this is the first study to demonstrate a predator altering the thermal physiology of amphibian larvae (prey) by increasing CTmax, increasing the optimum temperature, and producing changes in the thermal performance curves. Furthermore, these plastic responses of CTmax and TPC to different inducing environments should be considered when forecasting biological responses to global warming.

Spanish Ministry of Science and Innovation (MICINN) [CGL2012-40246-C02-01, CGL2012-40246-C02-02, BES-2010-032912]; Spanish Ministry of Economy, Industry and Competitiveness (MINECO) [CGL2017-86924-P]; Spanish Ministry of Education and Science (MEC FPU fellowship) [AP2005-0143]

Anurans breed in a variety of aquatic habitats with contrasting levels of desiccation risk, which may result in selection for faster development during larval stages. Previous studies suggest that species in ephemeral ponds reduce their developmental times to minimize desiccation risks, although it is not clear how variation in desiccation risk affects developmental strategies in different species. Employing a comparative phylogenetic approach including data from published and unpublished studies encompassing 62 observations across 30 species, we tested if species breeding in ephemeral ponds (High risk) develop faster than those from permanent ponds (Low risk) and/or show increased developmental plasticity in response to drying conditions. Our analyses support shorter developmental times in High risk, primarily by decreasing body mass at metamorphosis. Plasticity in developmental times was small and did not differ between groups. However, accelerated development in High risk species generally resulted in reduced sizes at metamorphosis, while some Low risk species were able compensate this effect by increasing mean growth rates. Taken together, our results suggest that plastic responses in species breeding in ephemeral ponds are constrained by a general trade‐off between development and growth rates. Anurans breed in a variety of aquatic habitats with contrasting levels of desiccation risk, which may result in selection for faster development during larval stages. Employing a comparative phylogenetic approach including data from published and unpublished studies encompassing 62 observations across 30 species, we tested if species breeding in ephemeral ponds (high risk) develop faster than those from permanent ponds (low risk) and/or show increased developmental plasticity in response to drying conditions.

Variation and population structure play key roles in the speciation process, but adaptive intraspecific genetic variation is commonly ignored when forecasting species niches. Amphibians serve as excellent models for testing how climate and local adaptations shape species distributions due to physiological and dispersal constraints and long generational times. In this study, we analysed the climatic factors driving the evolution of the genus Alytes at inter- and intraspecific levels that may limit realized niches. We tested for both differences among the five recognized species and among intraspecific clades for three of the species (Alytes obstetricans, A. cisternasii, and A. dickhilleni). We employed ecological niche models with an ordination approach to perform niche overlap analyses and test hypotheses of niche conservatism or divergence. Our results showed strong differences in the environmental variables affecting species climatic requirements. At the interspecific level, tests of equivalence and similarity revealed that sister species were non-identical in their environmental niches, although they neither were entirely dissimilar. This pattern was also consistent at the intraspecific level, with the exception of A. cisternasii, whose clades appeared to have experienced a lower degree of niche divergence than clades of the other species. In conclusion, our results support that Alytes toads, examined at both the intra- and interspecific levels, tend to occupy similar, if not identical, climatic environments.

Several amphibian lineages epitomize the faunal biodiversity crises, with numerous reports of population declines and extinctions worldwide. Predicting how such lineages will cope with environmental changes is an urgent challenge for biologists. A promising framework for this involves mechanistic modeling, which integrates organismal ecophysiological features and ecological models as a means to establish causal and consequential relationships of species with their physical environment. Solid frameworks built for other tetrapods (e.g., lizards) have proved successful in this context, but its extension to amphibians requires care. First, the natural history of amphibians is distinct within tetrapods, for it includes a biphasic life cycle that undergoes major habitat transitions and changes in sensitivity to environmental factors. Second, the accumulated data on amphibian ecophysiology is not nearly as expressive, is heavily biased towards adult lifeforms of few non-tropical lineages, and overlook the importance of hydrothermal relationships. Thus, we argue that critical usage and improvement in the available data is essential for enhancing the power of mechanistic modeling from the physiological ecology of amphibians. We highlight the complexity of ecophysiological variables and the need for understanding the natural history of the group under study and indicate directions deemed crucial to attaining steady progress in this field.

Captive-bred organisms are widely used in ecology, evolution and conservation research, especially in scenarios where natural populations are scarce or at risk of extinction. Yet, it is still unclear whether captivity may alter thermal tolerances, crucial traits to predict species resilience to global warming. Here, we study whether captive-bred tadpoles of the gliding treefrog (Agalychnis spurrelli) show different thermal tolerances than wild-caught individuals. Our results show that there are no differences between critical thermal limits (CTmax and CTmin) of captive-bred and wild-caught tadpoles exposed to three-day acclimatization at 20 °C. Therefore, we suggest that the use of captive-bred amphibians is valid and may be appropriate in experimental comparisons to thermal physiological studies of wild populations.