Abstract. Species in which ambient temperatures directly determine offspring sex may be at particular risk as global climates change. Whether or not climate change affects sex ratio depends upon the effectiveness of buffering mechanisms that link ambient regimes to actual nest temperatures. For example, females may simply lay nests earlier in the season, or in more shaded areas, such that incubation thermal regimes are unchanged despite massive ambient fluctuation. Based on eight years of monitoring nests over a 10-year period in the field at an alpine site in southeastern Australia, we show that, even though lizards (Bassiana duperreyi, Scincidae) have adjusted both nest depth and seasonal timing of oviposition in response to rising ambient temperatures, they have been unable to compensate entirely for climate change. That inability stems from the fact that the seasonal progression of soil temperatures, and thus, the degree to which thermal regimes at the time of laying predict subsequent conditions during incubation, also has shifted with climate change. As a result, mean incubation temperatures in natural nests now have crossed the thermal threshold at which incubation temperature directly affects offspring sex in this population.
Research on the thermal ecology and physiology of free‐living organisms is accelerating as scientists and managers recognize the urgency of the global biodiversity crisis brought on by climate change. As ectotherms, temperature fundamentally affects most aspects of the lives of amphibians and reptiles, making them excellent models for studying how animals are impacted by changing temperatures. As research on this group of organisms accelerates, it is essential to maintain consistent and optimal methodology so that results can be compared across groups and over time. This review addresses the utility of reptiles and amphibians as model organisms for thermal studies by reviewing the best practices for research on their thermal ecology and physiology, and by highlighting key studies that have advanced the field with new and improved methods. We end by presenting several areas where reptiles and amphibians show great promise for further advancing our understanding of how temperature relations between organisms and their environments are impacted by global climate change.
Much recent theoretical and empirical work has sought to describe the physiological mechanisms underlying thermal tolerance in animals. Leading hypotheses can be broadly divided into two categories that primarily differ in organizational scale: 1) high temperature directly reduces the function of subcellular machinery, such as enzymes and cell membranes, or 2) high temperature disrupts system-level interactions, such as mismatches in the supply and demand of oxygen, prior to having any direct negative effect on the subcellular machinery. Nonetheless, a general framework describing the contexts under which either subcellular component or organ system failure limits organisms at high temperatures remains elusive. With this commentary, we leverage decades of research on the physiology of ectothermic tetrapods (amphibians and non-avian reptiles) to address these hypotheses. Available data suggest both mechanisms are important. Thus, we expand previous work and propose the Hierarchical Mechanisms of Thermal Limitation (HMTL) hypothesis, which explains how subcellular and organ system failures interact to limit performance and set tolerance limits at high temperatures. We further integrate this framework with the thermal performance curve paradigm commonly used to predict the effects of thermal environments on performance and fitness. The HMTL framework appears to successfully explain diverse observations in reptiles and amphibians and makes numerous predictions that remain untested. We hope that this framework spurs further research in diverse taxa and facilitates mechanistic forecasts of biological responses to climate change.
Over the last few decades, biologists have made substantial progress in understanding relationships between changing climates and organism performance. Much of this work has focused on temperature because it is the best kept of climatic records, in many locations it is predicted to keep rising into the future, and it has profound effects on the physiology, performance, and ecology of organisms, especially ectothermic organisms which make up the vast majority of life on Earth. Nevertheless, much of the existing literature on temperature-organism interactions relies on mean temperatures. In reality, most organisms do not directly experience mean temperatures; rather, they experience variation in temperature over many time scales, from seconds to years. We propose to shift the focus more directly on patterns of temperature variation, rather than on means per se, and present a framework both for analyzing temporal patterns of temperature variation and for incorporating those patterns into predictions about organismal biology. In particular, we advocate using the Fourier transform to decompose temperature time series into their component sinusoids, thus allowing transformations between the time and frequency domains. This approach provides (1) standardized ways of visualizing the contributions that different frequencies make to total temporal variation; (2) the ability to assess how patterns of temperature variation have changed over the past half century and may change into the future; and (3) clear approaches to manipulating temporal time series to ask "what if" questions about the potential effects of future climates. We first summarize global patterns of change in temperature variation over the past 40 years; we find meaningful changes in variation at the half day to yearly times scales. We then demonstrate the utility of the Fourier framework by exploring how power added to different frequencies alters the overall incidence of long-term waves of high and low temperatures, and find that power added to the lowest frequencies greatly increases the probability of long-term heat and cold waves. Finally, we review what is known about the time scales over which organismal thermal performance curves change in response to variation in the thermal environment. We conclude that integrating information characterizing both the frequency spectra of temperature time series and the time scales of resulting physiological change offers a powerful new avenue for relating climate, and climate change, to the future performance of ectothermic organisms.
By altering phenology, organisms have the potential to match life-history events with suitable environmental conditions. Because of this, phenological plasticity has been proposed as a mechanism whereby populations might buffer themselves from climate change. We examine the potential buffering power of advancing one aspect of phenology, nesting date, on sex ratio in painted turtles (Chrysemys picta), a species with temperature-dependent sex determination. We developed a modified constant temperature equivalent model that accounts for the effect of the interaction among climate change, oviposition date, and seasonal thermal pattern on temperature during sexual differentiation and thus on offspring sex ratio. Our results suggest that females will not be able to buffer their progeny from the negative consequences of climate change by adjusting nesting date alone. Not only are offspring sex ratios predicted to become 100% female, but our model suggests that many nests will fail. Because the seasonal thermal trends that we consider are experienced by most temperate species, our result that adjusting spring phenology alone will be insufficient to counter the effects of directional climate change may be broadly applicable.
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