Phenotypic plasticity is ubiquitous and generally regarded as a key mechanism for enabling organisms to survive in the face of environmental change. Because no organism is infinitely or ideally plastic, theory suggests that there must be limits (for example, the lack of ability to produce an optimal trait) to the evolution of phenotypic plasticity, or that plasticity may have inherent significant costs. Yet numerous experimental studies have not detected widespread costs. Explicitly differentiating plasticity costs from phenotype costs, we re-evaluate fundamental questions of the limits to the evolution of plasticity and of generalists vs specialists. We advocate for the view that relaxed selection and variable selection intensities are likely more important constraints to the evolution of plasticity than the costs of plasticity. Some forms of plasticity, such as learning, may be inherently costly. In addition, we examine opportunities to offset costs of phenotypes through ontogeny, amelioration of phenotypic costs across environments, and the condition-dependent hypothesis. We propose avenues of further inquiry in the limits of plasticity using new and classic methods of ecological parameterization, phylogenetics and omics in the context of answering questions on the constraints of plasticity. Given plasticity's key role in coping with environmental change, approaches spanning the spectrum from applied to basic will greatly enrich our understanding of the evolution of plasticity and resolve our understanding of limits.

Many organisms have complex life cycles with distinct life stages that experience different environmental conditions. How does the complexity of life cycles affect the ecological and evolutionary responses of organisms to climate change? We address this question by exploring several recent case studies and synthetic analyses of insects. First, different life stages may inhabit different microhabitats, and may differ in their thermal sensitivities and other traits that are important for responses to climate. For example, the life stages of Manduca experience different patterns of thermal and hydric variability, and differ in tolerance to high temperatures. Second, life stages may differ in their mechanisms for adaptation to local climatic conditions. For example, in Colias, larvae in different geographic populations and species adapt to local climate via differences in optimal and maximal temperatures for feeding and growth, whereas adults adapt via differences in melanin of the wings and in other morphological traits. Third, we extend a recent analysis of the temperature-dependence of insect population growth to demonstrate how changes in temperature can differently impact juvenile survival and adult reproduction. In both temperate and tropical regions, high rates of adult reproduction in a given environment may not be realized if occasional, high temperatures prevent survival to maturity. This suggests that considering the differing responses of multiple life stages is essential to understand the ecological and evolutionary consequences of climate change.

Understanding the evolution of reaction norms remains a major challenge in ecology and evolution. Investigating evolutionary divergence in reaction norm shapes between populations and closely related species is one approach to providing insights. Here we use a meta-analytic approach to compare divergence in reaction norms of closely related species or populations of animals and plants across types of traits and environments. We quantified mean-standardized differences in overall trait means (Offset) and reaction norm shape (including both Slope and Curvature). These analyses revealed that differences in shape (Slope and Curvature together) were generally greater than differences in Offset. Additionally, differences in Curvature were generally greater than differences in Slope. The type of taxon contrast (species vs. population), trait, organism, and the type and novelty of environments all contributed to the best-fitting models, especially for Offset, Curvature, and the total differences (Total) between reaction norms. Congeneric species had greater differences in reaction norms than populations, and novel environmental conditions increased the differences in reaction norms between populations or species. These results show that evolutionary divergence of curvature is common and should be considered an important aspect of plasticity, together with slope. Biological details about traits and environments, including cryptic variation expressed in novel environmental conditions, may be critical to understanding how reaction norms evolve in novel and rapidly changing environments.

Abstract. Thermal constraints on development are often invoked to predict insect distributions. These constraints tend to be characterized in species distribution models (SDMs) by calculating development time based on a constant lower development temperature (LDT). Here, we assessed whether species-specific estimates of LDT based on laboratory experiments can improve the ability of SDMs to predict the distribution shifts of six U.K. butterflies in response to recent climate warming. We find that species-specific and constant (58C) LDT degree-day models perform similarly at predicting distributions during the period of 1970-1982. However, when the models for the 1970-1982 period are projected to predict distributions in 1995-1999 and 2000-2004, species-specific LDT degree-day models modestly outperform constant LDT degree-day models. Our results suggest that, while including species-specific physiology in correlative models may enhance predictions of species' distribution responses to climate change, more detailed models may be needed to adequately account for interspecific physiological differences.

The thermal biology of ectotherms is often used to infer species' responses to changes in temperature. It is often proposed that temperate species are more cold-tolerant, less heat-tolerant, more plastic, have broader thermal performance curves (TPCs) and lower optimal temperatures when compared to tropical species. However, relatively little empirical work has provided support for this using large interspecific studies. In the present study, we measure thermal tolerance limits and thermal performance in 22 species of Drosophila that developed under common conditions. Specifically, we measure thermal tolerance (CT min and CT max ) as well as the fitness components viability, developmental speed and fecundity at seven temperatures to construct TPCs for each of these species. For 10 of the species, we also measure thermal tolerance and thermal performance following developmental acclimation to three additional temperatures. Using these data, we test several fundamental hypotheses about the evolution and plasticity of heat and cold resistance and thermal performance. We find that cold tolerance (CT min ) varied between the species according to the environmental temperature in the habitat from which they originated. These data support the idea that the evolution of cold tolerance has allowed species to persist in colder environments. However, contrary to expectation, we find that optimal temperature ( T opt ) and the breadth of thermal performance ( T breadth ) are similar in temperate, widespread and tropical species and we also find that the plasticity of TPCs was constrained. We suggest that the temperature range for optimal thermal performance is either fixed or under selection by the more similar temperatures that prevail during growing seasons. As a consequence, we find that T opt and T breadth are of limited value for predicting past, present and future distributions of species. This article is part of the theme issue ‘Physiological diversity, biodiversity patterns and global climate change: testing key hypotheses involving temperature and oxygen’.

Summary1. Rapid evolution of physiological traits in response to novel thermal environments has rarely been demonstrated in natural populations. 2. We studied the temperature dependence [thermal performance curves (TPCs)] of larval feeding rate for two populations each of Colias eurytheme and Colias eriphyle in North America that occur over a range of elevations and climates. Using historical data for two of the populations, we assessed changes over time in both air temperatures and TPCs for larval feeding. 3. Populations at lower elevations with longer growing seasons had broader TPCs for larval feeding. In contrast, higher elevation populations with shorter growing seasons had higher optimal and maximal temperatures for feeding. 4. Overall mean air temperatures during the growing season showed little change at the two sites, but the frequency of high air temperatures (>28°C) has increased markedly at both sites over the past 40 years. This climatic shift was associated with increased rates of larval feeding at higher temperatures (>28°C) in both populations. 5. These results suggest that recent climate warming has led to physiological shifts in the TPCs for larval feeding in this system, indicating that thermal adaptation can occur rapidly in response to changing thermal conditions.

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In many ectotherms, exposure to high temperatures can improve subsequent tolerance to higher temperatures. However, the differential effects of single, repeated or continuous exposure to high temperatures are less clear. We measured the effects of single heat shocks and of diurnally fluctuating or constant rearing temperatures on the critical thermal maximum (CT max ) for final instar larvae of Manduca sexta. Brief (2 h) heat shocks at temperatures of 35°C and above significantly increased CT max relative to control temperatures (25°C). Increasing mean temperatures (from 25 to 30°C) or greater diurnal fluctuations (from constant to ±10°C) during larval development also significantly increased CT max . Combining these data showed that repeated or continuous temperature exposure during development improved heat tolerance beyond the effects of a single exposure to the same maximum temperature. These results suggest that both acute and chronic temperature exposure can result in adaptive plasticity of upper thermal limits.