Evaluating the fitness consequences of plasticity in tolerance to pesticides

In a rapidly changing world, phenotypic plasticity may be a critical mechanism allowing populations to rapidly acclimate when faced with novel anthropogenic stressors. Theory predicts that if exposure to anthropogenic stress is heterogeneous, plasticity should be maintained as it allows organisms to avoid unnecessary expression of costly traits (i.e., phenotypic costs) when stressors are absent. Conversely, if exposure to stressors becomes constant, costs or limits of plasticity may lead to evolutionary trait canalization (i.e., genetic assimilation). While these concepts are well‐established in theory, few studies have examined whether these factors explain patterns of plasticity in natural populations facing anthropogenic stress. Using wild populations of wood frogs that vary in plasticity in tolerance to pesticides, the goal of this study was to evaluate the environmental conditions under which plasticity is expected to be advantageous or detrimental. We found that when pesticides were absent, more plastic populations exhibited lower pesticide tolerance and were more fit than less plastic populations, likely avoiding the cost of expressing high tolerance when it was not necessary. Contrary to our predictions, when pesticides were present, more plastic populations were as fit as less plastic populations, showing no signs of costs or limits of plasticity. Amidst unprecedented global change, understanding the factors shaping the evolution of plasticity will become increasingly important.     

Stronger compensatory growth in a permanent-pond Lestes damselfly relative to temporary-pond Lestes

Compensatory growth where animals compensate for time stress or transient nutritional or thermal stress by accelerating their growth rate is widespread. We know, however, relatively little about the evolution and ecological correlates of compensatory growth. For this we need studies on congeneric species with known phylogenetic relationships that also focus on the associated largely understudied costs. Here we tested for compensatory growth and associated costs in response to time stress (manipulated by photoperiod) and a transient period of starvation or cooling in larvae of the permanent-pond damselfly Lestes eurinus , and compare the results with former studies on temporary-pond Lestes . Larvae showed full compensation in body mass at emergence for all combinations of time stress and starvation or cooling. Unexpectedly, compensatory growth to starvation or cooling was not stronger under time stress. Instead, males under time stress delayed emergence after these transient stressors. In line with a stronger compensatory growth response to time stress than to the other stressors, physiological costs in terms of a reduced investment in immune response (measured as phenoloxidase activity) and energy storage (measured as fat content) were detected only under time stress. Compared to temporary-pond Lestes , L. eurinus showed stronger compensatory growth to time stress. We hypothesize that the stronger compensatory (growth) response in permanent-pond Lestes co-evolved with their derived slower lifestyle when they invaded permanent ponds.     

Collateral damage: rapid exposure-induced evolution of pesticide resistance leads to increased susceptibility to parasites

Although natural populations may evolve resistance to anthropogenic stressors such as pollutants, this evolved resistance may carry costs. Using an experimental evolution approach, we exposed different Daphnia magna populations in outdoor containers to the carbamate pesticide carbaryl and control conditions, and assessed the resulting populations for both their resistance to carbaryl as well as their susceptibility to infection by the widespread bacterial microparasite Pasteuria ramosa. Our results show that carbaryl selection led to rapid evolution of carbaryl resistance with seemingly no cost when assessed in a benign environment. However, carbaryl-resistant populations were more susceptible to parasite infection than control populations. Exposure to both stressors reveals a synergistic effect on sterilization rate by P. ramosa, but this synergism did not evolve under pesticide selection. Assessing costs of rapid adaptive evolution to anthropogenic stress in a semi-natural context may be crucial to avoid too optimistic predictions for the fitness of the evolving populations.     

Evaluation of physiological stress in free-ranging bears: current knowledge and future directions

Stress responses, which are mediated by the neurogenic system (NS) and hypothalamic–pituitary–adrenal (HPA) axis help vertebrates maintain physiological homeostasis. Fight-or-flight responses are activated by the NS, which releases norepinephrine/noradrenaline and epinephrine/adrenaline in response to immediate stressors, whilst the HPA axis releases glucocorticoid hormones (e.g. cortisol and corticosterone) to help mitigate allostatic load. There have been many studies on stress responses of captive animals, but they are not truly reflective of typical ranges or the types of stressors encountered by free-ranging wildlife, such as responses and adaptation to environmental change, which are particularly important from a conservation perspective. As stress can influence the composition of age and sex classes of free-ranging populations both directly and indirectly, ecological research must be prioritised towards more vulnerable taxa. Generally, large predators tend to be particularly at risk of anthropogenically driven population declines because they exhibit reduced behavioural plasticity required to adapt to changing landscapes and exist in reduced geographic ranges, have small population sizes, low fecundity rates, large spatial requirements and occupy high trophic positions. As a keystone species with a long history of coexistence with humans in highly anthropogenic landscapes, there has been growing concern about how humans influence bear behaviour and physiology, via numerous short- and long-term stressors. In this review, we synthesise research on the stress response in free-ranging bear populations and evaluate the effectiveness and limitations of current methodology in measuring stress in bears to identify the most effective metrics for future research. Particularly, we integrate research that utilised haematological variables, cardiac monitors and Global Positioning System (GPS) collars, serum/plasma and faecal glucocorticoid concentrations, hair cortisol levels, and morphological metrics (primarily skulls) to investigate the stress response in ursids in both short- and long-term contexts. We found that in free-ranging bears, food availability and consumption have the greatest influence on individual stress, with mixed responses to anthropogenic influences. Effects of sex and age on stress are also mixed, likely attributable to inconsistent methods. We recommend that methodology across all stress indicators used in free-ranging bears should be standardised to improve interpretation of results and that a wider range of species should be incorporated in future studies.     

Potential local adaptation in populations of invasive reed canary grass (Phalaris arundinacea) across an urbanization gradient

Urban stressors represent strong selective gradients that can elicit evolutionary change, especially in non‐native species that may harbor substantial within‐population variability. To test whether urban stressors drive phenotypic differentiation and influence local adaptation, we compared stress responses of populations of a ubiquitous invader, reed canary grass (Phalaris arundinacea). Specifically, we quantified responses to salt, copper, and zinc additions by reed canary grass collected from four populations spanning an urbanization gradient (natural, rural, moderate urban, and intense urban). We measured ten phenotypic traits and trait plasticities, because reed canary grass is known to be highly plastic and because plasticity may enhance invasion success. We tested the following hypotheses: (a) Source populations vary systematically in their stress response, with the intense urban population least sensitive and the natural population most sensitive, and (b) plastic responses are adaptive under stressful conditions. We found clear trait variation among populations, with the greatest divergence in traits and trait plasticities between the natural and intense urban populations. The intense urban population showed stress tolerator characteristics for resource acquisition traits including leaf dry matter content and specific root length. Trait plasticity varied among populations for over half the traits measured, highlighting that plasticity differences were as common as trait differences. Plasticity in root mass ratio and specific root length were adaptive in some contexts, suggesting that natural selection by anthropogenic stressors may have contributed to root trait differences. Reed canary grass populations in highly urbanized wetlands may therefore be evolving enhanced tolerance to urban stressors, suggesting a mechanism by which invasive species may proliferate across urban wetland systems generally.     

植物抗污染进化及其遗传生态学代价

植物抗污染进化及其遗传生态学代价熊治廷（武汉大学环境科学系,４３００７２）ＰｏｌｕｔｉｏｎＲｅｓｉｓｔａｎｔＥｖｏｌｕｔｉｏｎｉｎＰｌａｎｔｓａｎｄＩｔｓＧｅｎｅｃｏｌｏｇｉｃａｌＣｏｓｔｓ．ＸｉｏｎｇＺｈｉｔｉｎｇ（ＤｅｐａｒｔｍｅｎｔｏｆＥｎｖ...     

Limitations of cross‐ and multigenerational plasticity for marine invertebrates faced with global climate change

Although cross generation (CGP) and multigenerational (MGP) plasticity have been identified as mechanisms of acclimation to global change, the weight of evidence indicates that parental conditioning over generations is not a panacea to rescue stress sensitivity in offspring. For many species, there were no benefits of parental conditioning. Even when improved performance was observed, this waned over time within a generation or across generations and fitness declined. CGP and MGP studies identified resilient species with stress tolerant genotypes in wild populations and selected family lines. Several bivalves possess favourable stress tolerance and phenotypically plastic traits potentially associated with genetic adaptation to life in habitats where they routinely experience temperature and/or acidification stress. These traits will be important to help ‘climate proof’ shellfish ventures. Species that are naturally stress tolerant and those that naturally experience a broad range of environmental conditions are good candidates to provide insights into the physiological and molecular mechanisms involved in CGP and MGP. It is challenging to conduct ecologically relevant global change experiments over the long times commensurate with the pace of changing climate. As a result, many studies present stressors in a shock‐type exposure at rates much faster than projected scenarios. With more gradual stressor introduction over longer experimental durations and in context with conditions species are currently acclimatized and/or adapted to, the outcomes for sensitive species might differ. We highlight the importance to understand primordial germ cell development and the timing of gametogenesis with respect to stressor exposure. Although multigenerational exposure to global change stressors currently appears limited as a universal tool to rescue species in the face of changing climate, natural proxies of future conditions (upwelling zones, CO₂ vents, naturally warm habitats) show that phenotypic adjustment and/or beneficial genetic selection is possible for some species, indicating complex plasticity–adaptation interactions.     

Variability in responses to thermal stress in parasitoids

Abstract 1. To study phenotypic effects of stress, a stress is applied to cohorts of organisms with an increasing intensity. In the absence of mortality the response of traits will be a decreasing function of stress intensity because of increasing physiological costs. We call such decreasing functions type A responses. 2. However, when stress caused mortality, some studies have found that for high stress intensities, survivors performed as well as control individuals (type B responses). We proposed that type A responses are caused by the physiological cost of stress whereas type B responses are caused by a mixture of physiological costs and selection. 3. The present study exposed Aphidius picipes wasps to an increasing duration of cold storage (cold stress), and obtained variable responses as predicted when both physiological costs and selection of resistant individuals determine the outcome. 4. When cold storage of parasitoids for biological control is desirable, research should be carried out to find (i) the temperature regime and duration of storage and (ii) the least sensitive stage for storage to minimise losses from mortality and reduction of fitness of survivors. 5. Selection by cold stress as observed in the present study could result in rapid adaptation of populations exposed to such stress.     

Long-term forecast of thermal mortality with climate warming in riverine amphipods

Forecasting long-term consequences of global warming requires knowledge on thermal mortality and how heat stress interacts with other environmental stressors on different timescales. Here, we describe a flexible analytical framework to forecast mortality risks by combining laboratory measurements on tolerance and field temperature records. Our framework incorporates physiological acclimation effects, temporal scale differences and the ecological reality of fluctuations in temperature, and other factors such as oxygen. As a proof of concept, we investigated the heat tolerance of amphipods Dikerogammarus villosus and Echinogammarus trichiatus in the river Waal, the Netherlands. These organisms were acclimated to different temperatures and oxygen levels. By integrating experimental data with high-resolution field data, we derived the daily heat mortality probabilities for each species under different oxygen levels, considering current temperatures as well as 1 and 2°C warming scenarios. By expressing heat stress as a mortality probability rather than a upper critical temperature, these can be used to calculate cumulative annual mortality, allowing the scaling up from individuals to populations. Our findings indicate a substantial increase in annual mortality over the coming decades, driven by projected increases in summer temperatures. Thermal acclimation and adequate oxygenation improved heat tolerance and their effects were magnified on longer timescales. Consequently, acclimation effects appear to be more effective than previously recognized and crucial for persistence under current temperatures. However, even in the best-case scenario, mortality of D. villosus is expected to approach 100% by 2100, while E. trichiatus appears to be less vulnerable with mortality increasing to 60%. Similarly, mortality risks vary spatially: In southern, warmer rivers, riverine animals will need to shift from the main channel toward the cooler head waters to avoid thermal mortality. Overall, this framework generates high-resolution forecasts on how rising temperatures, in combination with other environmental stressors such as hypoxia, impact ecological communities.     

Plasticity and cross‐tolerance to heterogeneous environments: divergent stress responses co‐evolved in an African fruit fly

Plastic adjustments of physiological tolerance to a particular stressor can result in fitness benefits for resistance that might manifest not only in that same environment but also be advantageous when faced with alternative environmental stressors, a phenomenon termed ‘cross‐tolerance’. The nature and magnitude of cross‐tolerance responses can provide important insights into the underlying genetic architecture, potential constraints on or versatility of an organism's stress responses. In this study, we tested for cross‐tolerance to a suite of abiotic factors that likely contribute to setting insect population dynamics and geographic range limits: heat, cold, desiccation and starvation resistance in adult Ceratitis rosa following acclimation to all these isolated individual conditions prior to stress assays. Traits of stress resistance scored included critical thermal (activity) limits, chill coma recovery time (CCRT), heat knockdown time (HKDT), desiccation and starvation resistance. In agreement with other studies, we found that acclimation to one stress typically increased resistance for that same stress experienced later in life. A more novel outcome, however, is that here we also found substantial evidence for cross‐tolerance. For example, we found an improvement in heat tolerance (critical thermal maxima, CT_max) following starvation or desiccation hardening and improved desiccation resistance following cold acclimation, indicating pronounced cross‐tolerance to these environmental stressors for the traits examined. We also found that two different traits of the same stress resistance differed in their responsiveness to the same stress conditions (e.g. HKDT was less cross‐resistant than CT_max). The results of this study have two major implications that are of broader importance: (i) that these traits likely co‐evolved to cope with diverse or simultaneous stressors, and (ii) that a set of common underlying physiological mechanisms might exist between apparently divergent stress responses in this species. This species may prove to be a valuable model for future work on the evolutionary and mechanistic basis of cross‐tolerance.     

South African ecotoxicology — present status and future prognosis

Toxicology studies the interactions of a chemical substance with individual organisms, whereas ecotoxicology is a multidisciplinary approach incorporating ecology and other disciplines, e.g. chemistry, microbiology, etc., to determine responses of individuals, populations and whole ecosystems to stressors such as chemicals. We present here the current status of toxicity testing in South Africa and propose a future prognosis for such tests. We propose a path forward for the development of ecotoxicology in South Africa and also globally. Toxicity testing issues dealt with are the use of surrogate species as opposed to indigenous species, their comparative tolerances, and the selection of relevant endpoints as measures of toxicity. Ecotoxicological considerations need to address the following key ecological realities: tolerance (both physiological acclimation and genetic adaptation), trophic redundancies, resilience, compensation (e.g. density dependence), evolution, and recovery. We believe that predictive ecotoxicology will play a major role in the future management of ecosystems that are constantly changing. We also believe that such management must be proactive to the point of intervention to create desired change, specifically the maintenance of ecosystem services.     

Cold shock and fish

Rapid decreases in water temperature may result in a number of physiological, behavioural and fitness consequences for fishes termed ‘cold shock’. Cold‐shock stress occurs when a fish has been acclimated to a specific water temperature or range of temperatures and is subsequently exposed to a rapid decrease in temperature, resulting in a cascade of physiological and behavioural responses and, in some cases, death. Rapid temperature decreases may occur from either natural (e.g. thermocline temperature variation, seiches and storm events) or anthropogenic sources (e.g. varied thermal effluents from power generation and production industries). The magnitude, duration and frequency of the temperature change as well as the initial acclimation temperatures of individuals can influence the extent of the consequences of cold shock on fishes. Early research on cold shock focused on documenting mortality events associated with cold shock. However, in recent years, a shift in research has occurred where the focus of cold‐shock studies now involves characterizing the sublethal effects of cold shock in terms of the stress response in fishes. This shift has revealed that cold shock can actually be used as a tool for fisheries science (e.g. to induce polyploidy). The cold‐shock stress response offers opportunities to develop many exciting research questions, yet to date, cold‐shock research has been largely unfocused. Few studies attempt to link laboratory physiology experiments with ecologically relevant field data on behaviour, growth, bioenergetics and fitness. Additional research will allow for the development of more focused and robust management policies and conservation initiatives. This review synthesizes the sublethal physiological and behavioural consequences of cold‐shock stress on fishes, identifies natural and anthropogenic sources of cold shock, discusses the benefits of cold shock to fisheries science and describes mitigation and management efforts. Existing knowledge gaps and opportunities for future cold‐shock research are presented.     

Fecal glucocorticoids reflect socio-ecological and anthropogenic stressors in the lives of wild spotted hyenas

Our aim was to identify natural and anthropogenic influences on the stress physiology of large African carnivores, using wild spotted hyenas (Crocuta crocuta) as model animals. With both longitudinal data from a single social group, and cross-sectional data from multiple groups, we used fecal glucocorticoids (fGC) to examine potential stressors among spotted hyenas. Longitudinal data from adult members of a group living on the edge of the Masai Mara National Reserve, Kenya, revealed that fGC concentrations were elevated during two periods of social upheaval among adults, especially among younger females; however, prey availability, rainfall, and presence of lions did not influence fGC concentrations among hyenas. Our results suggested that anthropogenic disturbance in the form of pastoralist activity, but not tourism, influenced fGC concentrations among adult male hyenas; rising concentrations of fGC among males over 12 years were significantly correlated with increasing human population density along the edge of the group's home range. As hyenas from this social group were frequently exposed to anthropogenic disturbance, we compared fGC concentrations among these hyenas with those obtained concurrently from hyenas living in three other groups undisturbed by pastoralist activity. We found that fGC concentrations from the undisturbed groups were significantly lower than those in the disturbed group, and we were able to rule out tourism and ecological stressors as sources of variation in fGC among the populations. Thus it appears that both social instability and anthropogenic disturbance, but not the ecological variables examined, elevate fGC concentrations and represent stressors for wild spotted hyenas. Further work will be necessary to determine whether interpopulation variation in stress physiology predicts population decline in groups exposed to intensive anthropogenic disturbance.     

Behavioral and respiratory responses to stressors in multiple populations of three-spined sticklebacks that differ in predation pressure

Individual animals of the same species inhabiting environments which differ in the frequency and magnitude of stressors often exhibit different physiological and behavioral responses to stressors. Here, we compare the respiratory response to confinement stress, and behavioral responses to ecologically relevant challenges among sticklebacks from 11 different populations varying in predation pressure. We found that sticklebacks from high predation populations breathed faster in response to confinement stress and were, on an average, more behaviorally responsive to a pike behind glass compared with sticklebacks from low predation populations. These patterns differ from the results of studies on other species, highlighting the need for a conceptual framework to understand the proximate and ultimate factors shaping variable responses to stressors over developmental and evolutionary time. Moreover, physiological and behavioral responses were integrated with each other, both at the individual and population levels. In general, fish that were more aggressive and bold in the presence of a predator breathed faster, independent of body size. These results are consistent with the growing body of evidence that individuals differ in a suite of physiological and behavioral mechanisms for coping with challenges in the environment.     

Elevational variation in adult body size and growth rate but not in metabolic rate in the tree weta Hemideina crassidens

Populations of the same species inhabiting distinct localities experience different ecological and climatic pressures that might result in differentiation in traits, particularly those related to temperature. We compared metabolic rate (and its thermal sensitivity), growth rate, and body size among nine high- and low-elevation populations of the Wellington tree weta, Hemideina crassidens, distributed from 9 to 1171 m a.s.l across New Zealand. Our results did not indicate elevational compensation in metabolic rates (metabolic cold adaptation). Cold acclimation decreased metabolic rate compared to warm-acclimated individuals from both high- and low-elevation populations. However, we did find countergradient variation in growth rates, with individuals from high-elevation populations growing faster and to a larger final size than individuals from low-elevation populations. Females grew faster to a larger size than males, although as adults their metabolic rates did not differ significantly. The combined physiological and morphological data suggest that high-elevation individuals grow quickly and achieve larger size while maintaining metabolic rates at levels not significantly different from low-elevation individuals. Thus, morphological differentiation among tree weta populations, in concert with genetic variation, might provide the material required for adaptation to changing conditions.     

Telomere length reveals cumulative individual and transgenerational inbreeding effects in a passerine bird

Inbreeding results in more homozygous offspring that should suffer reduced fitness, but it can be difficult to quantify these costs for several reasons. First, inbreeding depression may vary with ecological or physiological stress and only be detectable over long time periods. Second, parental homozygosity may indirectly affect offspring fitness, thus confounding analyses that consider offspring homozygosity alone. Finally, measurement of inbreeding coefficients, survival and reproductive success may often be too crude to detect inbreeding costs in wild populations. Telomere length provides a more precise measure of somatic costs, predicts survival in many species and should reflect differences in somatic condition that result from varying ability to cope with environmental stressors. We studied relative telomere length in a wild population of Seychelles warblers (Acrocephalus sechellensis) to assess the lifelong relationship between individual homozygosity, which reflects genome‐wide inbreeding in this species, and telomere length. In juveniles, individual homozygosity was negatively associated with telomere length in poor seasons. In adults, individual homozygosity was consistently negatively related to telomere length, suggesting the accumulation of inbreeding depression during life. Maternal homozygosity also negatively predicted offspring telomere length. Our results show that somatic inbreeding costs are environmentally dependent at certain life stages but may accumulate throughout life.     

Does the Cost of Adaptation to Extremely Stressful Environments Diminish Over Time? A Literature Synthesis on How Plants Adapt to Heavy Metals and Pesticides

Populations adapted to locally stressful environmental conditions are predicted to carry costs in performance and fitness, particularly when compared to non-stress adapted populations in the absence of stress. However, empirical observations found fitness costs incurred by stress-resistant genotypes are often ambiguous or absent. Compensatory evolution may purge genotypes with relatively high costs over time, resulting in the recovery of fitness in a stress-resistant population. We assessed the magnitude of adaptation costs over time to test for a reduction in negative genetic effects by compiling published data on measures of fitness from plant populations inhabiting mine tailings and populations adapted to herbicides. Heavy metal contaminated sites represent a stress that is immediate and unchanging; herbicides represent a stress that changes over time with dosage or the type of herbicide as treated populations become more resistant. To quantify costs, for each comparison we recorded the performance of plants from stress and non-stress environments grown under benign conditions. Time since the initiation of the stress was determined to test whether costs change over time. Costs were overall constant through time. The magnitude of cost were consistent with trade-offs for heavy metal resistance and certain herbicide mechanisms (triazine and resistance via P450 enzyme), but not for other herbicides where costs were inconsistent and appear to be low if not absent. Superior stress-resistant populations with higher performance than non-stress populations were found from both herbicide and metal stress, with some extreme cases early from time since initiation. There was an increasing benefit to cost ratio over time for herbicide resistant populations. We found that adaptation to stressful environments is generally costly except in herbicide resistance, and that costs are not diminished over time. Stress-resistant populations without costs also arise infrequently, though these populations may often be restricted from spreading.     

Evaluating methods to quantify anthropogenic stressors on wild animals

Humans have a variety of direct and indirect impacts on wildlife and a number of methods have been proposed to identify and quantify anthropogenic stressors that negatively impact wildlife. The ideal method would ultimately help predict the presence, absence, or population viability of animals living with a particular stressor. We critically review seven methods that have been used, or are potentially useful, to identify anthropogenic stressors on animals. We rank them from fitness indicators to disturbance indicators: breeding success, mate choice, fluctuating asymmetry, flight initiation distance, immunocompetence, glucocorticoids, and cardiac response. We describe each method's ease of use, precision in quantifying the stressor, accuracy in predicting the presence, absence, or population viability of a species experiencing a given stressor, and the repeatability of the results across populations and species. From this analysis, we conclude that there is no single optimal method to quantify anthropogenic stressors; method selection will depend on precise goals and fiscal constraints.     

A warmer environment can reduce sociability in an ectotherm

The costs and benefits of being social vary with environmental conditions, so individuals must weigh the balance between these trade-offs in response to changes in the environment. Temperature is a salient environmental factor that may play a key role in altering the costs and benefits of sociality through its effects on food availability, predator abundance, and other ecological parameters. In ectotherms, changes in temperature also have direct effects on physiological traits linked to social behaviour, such as metabolic rate and locomotor performance. In light of climate change, it is therefore important to understand the potential effects of temperature on sociality. Here, we took the advantage of a ‘natural experiment’ of threespine sticklebacks from contrasting thermal environments in Iceland: geothermally warmed water bodies (warm habitats) and adjacent ambient-temperature water bodies (cold habitats) that were either linked (sympatric) or physically distinct (allopatric). We first measured the sociability of wild-caught adult fish from warm and cold habitats after acclimation to a low and a high temperature. At both acclimation temperatures, fish from the allopatric warm habitat were less social than those from the allopatric cold habitat, whereas fish from sympatric warm and cold habitats showed no differences in sociability. To determine whether differences in sociability between thermal habitats in the allopatric population were heritable, we used a common garden breeding design where individuals from the warm and the cold habitat were reared at a low or high temperature for two generations. We found that sociability was indeed heritable but also influenced by rearing temperature, suggesting that thermal conditions during early life can play an important role in influencing social behaviour in adulthood. By providing the first evidence for a causal effect of rearing temperature on social behaviour, our study provides novel insights into how a warming world may influence sociality in animal populations.