The majority of ectotherms grow slower but mature at a larger body size in colder environments. This phenomenon has puzzled biologists because classic theories of life-history evolution predict smaller sizes at maturity in environments that retard growth. During the last decade, intensive theoretical and empirical research has generated some plausible explanations based on nonadaptive or adaptive plasticity. Nonadaptive plasticity of body size is hypothesized to result from thermal constraints on cellular growth that cause smaller cells at higher temperatures, but the generality of this theory is poorly supported. Adaptive plasticity is hypothesized to result from greater benefits or lesser costs of delayed maturation in colder environments. These theories seem to apply well to some species but not others. Thus, no single theory has been able to explain the generality of temperature-size relationships in ectotherms. We recommend a multivariate theory that focuses on the coevolution of thermal reaction norms for growth rate and size at maturity. Such a theory should incorporate functional constraints on thermal reaction norms, as well as the natural covariation between temperature and other environmental variables.
Two major approaches address the need to predict species distributions in response to environmental changes. Correlative models estimate parameters phenomenologically by relating current distributions to environmental conditions. By contrast, mechanistic models incorporate explicit relationships between environmental conditions and organismal performance, estimated independently of current distributions. Mechanistic approaches include models that translate environmental conditions into biologically relevant metrics (e.g. potential duration of activity), models that capture environmental sensitivities of survivorship and fecundity, and models that use energetics to link environmental conditions and demography. We compared how two correlative and three mechanistic models predicted the ranges of two species: a skipper butterfly (Atalopedes campestris) and a fence lizard (Sceloporus undulatus). Correlative and mechanistic models performed similarly in predicting current distributions, but mechanistic models predicted larger range shifts in response to climate change. Although mechanistic models theoretically should provide more accurate distribution predictions, there is much potential for improving their flexibility and performance.
We review the evidence for the role of climate change in triggering disease outbreaks of chytridiomycosis, an emerging infectious disease of amphibians. Both climatic anomalies and disease-related extirpations are recent phenomena, and effects of both are especially noticeable at high elevations in tropical areas, making it difficult to determine whether they are operating separately or synergistically. We compiled reports of amphibian declines from Lower Central America and Andean South America to create maps and statistical models to test our hypothesis of spatiotemporal spread of the pathogen Batrachochytrium dendrobatidis (Bd), and to update the elevational patterns of decline in frogs belonging to the genus Atelopus. We evaluated claims of climate change influencing the spread of Bd by including error into estimates of the relationship between air temperature and last year observed. Available data support the hypothesis of multiple introductions of this invasive pathogen into South America and subsequent spread along the primary Andean cordilleras. Additional analyses found no evidence to support the hypothesis that climate change has been driving outbreaks of amphibian chytridiomycosis, as has been posited in the climate-linked epidemic hypothesis. Future studies should increase retrospective surveys of museum specimens from throughout the Andes and should study the landscape genetics of Bd to map fine-scale patterns of geographic spread to identify transmission routes and processes.
Although climates are rapidly changing on a global scale, these changes cannot easily be extrapolated to the local scales experienced by organisms. In fact, such generalizations might be quite problematic. For instance, models used to predict shifts in the ranges of species during climate change rarely incorporate data resolved to <1 km(2), although most organisms integrate climatic drivers at much smaller scales. Empirical studies alone suggest that the operative temperatures of many organisms vary by as much as 10-20 °C on a local scale, depending on vegetation, geology, and topography. Furthermore, this variation in abiotic factors ignores thermoregulatory behaviors that many animals use to balance heat loads. Through a set of simulations, we demonstrate how variability in elevational topography can attenuate the effects of warming climates. These simulations suggest that changing climates do not always impact organisms negatively. Importantly, these simulations involve well-known relationships in biophysical ecology that show how no two organisms experience the same climate in the same way. We suggest that, when coupled with thermoregulatory behavior, variation in topographic features can mask the acute effect of climate change in many cases.
Although most organisms thermoregulate behaviorally, biologists still cannot easily predict whether mobile animals will thermoregulate in natural environments. Current models fail because they ignore how the spatial distribution of thermal resources constrains thermoregulatory performance over space and time. To overcome this limitation, we modeled the spatially explicit movements of animals constrained by access to thermal resources. Our models predict that ectotherms thermoregulate more accurately when thermal resources are dispersed throughout space than when these resources are clumped. This prediction was supported by thermoregulatory behaviors of lizards in outdoor arenas with known distributions of environmental temperatures. Further, simulations showed how the spatial structure of the landscape qualitatively affects responses of animals to climate. Biologists will need spatially explicit models to predict impacts of climate change on local scales.behavioral thermoregulation | thermal heterogeneity | thermal ecology | spatial ecology | individual-based model T he rapid warming of many environments has generated great concern about the potential impacts on biodiversity (1). Genetic changes in response to anthropogenic warming seem rare (2) or limited (3), and many species have shifted habitats over space and time (4-7). Indeed, facultative behavioral strategies are the primary means by which many species cope with changing environments (8). In a warming world, behavioral thermoregulation could enable most organisms to maintain body temperatures that promote physiological performance (9-11). However, excessive warming constrains thermoregulation, potentially leading to extinction of populations. At local scales, recent warming apparently caused numerous extinctions by limiting the duration of foraging by lizards (12). According to mechanistic models, thermal constraints on activity will play a major role in biological invasions and local extinctions (13-16). Given constraints on thermoregulatory behaviors, some have predicted that global warming could eliminate more than 40% of lizard species by 2080 (12).Such projections, although dire, underestimate the impacts of climate change by failing to consider costs of thermoregulation that are imposed by environmental heterogeneity (10,17,18). Most models assume that an animal can access either unshaded michrohabitats or shaded microhabitats without using energy to search for and move between them (14, 19). As long as the animals prefers a body temperature within the range of operative environmental temperatures, an animal can thermoregulate by shuttling between microhabitats at no cost. Given this assumption, researchers combine meteorological data and biophysical equations to calculate the expected performance of an organism in specific climates. However, thermoregulatory behaviors impose costs such as energy loss, predation risk, and missed opportunities for foraging and breeding (20), which researchers have ignored when modeling the biological impacts of clim...
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