Climate drives among‐year variation in natural selection on flowering time

To predict long‐term responses to climate change, we need to understand how changes in temperature and precipitation elicit both immediate phenotypic responses and changes in natural selection. We used 22 years of data for the perennial herb Lathyrus vernus to examine how climate influences flowering phenology and phenotypic selection on phenology. Plants flowered earlier in springs with higher temperatures and higher precipitation. Early flowering was associated with a higher fitness in nearly all years, but selection for early flowering was significantly stronger in springs with higher temperatures and lower precipitation. Climate influenced selection through trait distributions, mean fitness and trait?fitness relationships, the latter accounting for most of the among‐year variation in selection. Our results show that climate both induces phenotypic responses and alters natural selection, and that the change in the optimal phenotype might be either weaker, as for spring temperature, or stronger, as for precipitation, than the optimal response.     

Dynamics of phenotypic change: wing length declines in a resident farmland passerine despite survival advantage of longer wings

In many taxa, environmental changes that alter resource availability and energetics, such as climate change and land use change, are associated with changes in body size. We use wing length as a proxy for overall structural body size to examine a paradoxical trend of declining wing length within a Yellowhammer Emberiza citrinella population sampled over 21 years, in which it has been previously shown that longer wings are associated with higher survival rates. Higher temperatures during the previous winter (prior to the moult determining current wing length) explained 23% of wing length decrease within our population, but changes may also be correlated with non‐climatic environmental variation such as changes in farming mechanisms linked to food availability. We found no evidence for within‐individual wing length shrinkage with age, but our data suggested a progressive decline in the sizes of immature birds recruiting to the population. This trend was weaker, although not significantly so, among adults, suggesting that the decline in the sizes of recruits was offset by higher subsequent survival of larger birds post‐recruitment. These data suggest that ecological processes can contribute more than selection to observed phenotypic trends and highlight the importance of long‐term studies for providing longitudinal insights into population processes.     

Rapid evolution of thermal plasticity in mountain lake Daphnia populations

Populations at risk of extinction due to climate change may be rescued by adaptive evolution or plasticity. Selective agents, such as introduced predators, may enhance or constrain plastic or adaptive responses to temperature. We tested responses of Daphnia to temperature by collecting populations from lakes across an elevational gradient in the presence and absence of fish predators (long‐term selection). We subsequently grew these populations at two elevations in field mesocosms over two years (short‐term selection), followed by a common‐garden experiment at two temperatures in the lab to measure life‐history traits. Both long‐term and short‐term selection affected traits, suggesting that genetic variation of plasticity within populations enabled individuals to rapidly evolve plasticity in response to high temperature. We found that short‐term selection by high temperature increased plasticity for growth rate in all populations. Fecundity was higher in populations from fishless lakes and body size showed greater plasticity in populations from warm lakes (long‐term selection). Neither body size nor fecundity were affected by short‐term thermal selection. These results demonstrate that plasticity is an important component of the life‐history response of Daphnia, and that genetic variation within populations enabled rapid evolution of plasticity in response to selection by temperature.     

Body size‐dependent responses of a marine fish assemblage to climate change and fishing over a century‐long scale

Commercial fishing and climate change have influenced the composition of marine fish assemblages worldwide, but we require a better understanding of their relative influence on long‐term changes in species abundance and body‐size distributions. In this study, we investigated long‐term (1911–2007) variability within a demersal fish assemblage in the western English Channel. The region has been subject to commercial fisheries throughout most of the past century, and has undergone interannual changes in sea temperature of over 2.0 °C. We focussed on a core 30 species that comprised 99% of total individuals sampled in the assemblage. Analyses showed that temporal trends in the abundance of smaller multispecies size classes followed thermal regime changes, but that there were persistent declines in abundance of larger size classes. Consistent with these results, larger‐growing individual species had the greatest declines in body size, and the most constant declines in abundance, while abundance changes of smaller‐growing species were more closely linked to preceding sea temperatures. Together these analyses are suggestive of dichotomous size‐dependent responses of species to long‐term climate change and commercial fishing over a century scale. Small species had rapid responses to the prevailing thermal environment, suggesting their life history traits predisposed populations to respond quickly to changing climates. Larger species declined in abundance and size, reflecting expectations from sustained size‐selective overharvesting. These results demonstrate the importance of considering species traits when developing indicators of human and climatic impacts on marine fauna.     

Genotype‐environment interactions rule the response of a widespread butterfly to temperature variation

Understanding how organisms adapt to complex environments is a central goal of evolutionary biology and ecology. This issue is of special interest in the current era of rapidly changing climatic conditions. Here, we investigate clinal variation and plastic responses in life history, morphology and physiology in the butterfly Pieris napi along a pan‐European gradient by exposing butterflies raised in captivity to different temperatures. We found clinal variation in body size, growth rates and concomitant development time, wing aspect ratio, wing melanization and heat tolerance. Individuals from warmer environments were more heat‐tolerant and had less melanised wings and a shorter development, but still they were larger than individuals from cooler environments. These findings suggest selection for rapid growth in the warmth and for wing melanization in the cold, and thus fine‐tuned genetic adaptation to local climates. Irrespective of the origin of butterflies, the effects of higher developmental temperature were largely as expected, speeding up development; reducing body size, potential metabolic activity and wing melanization; while increasing heat tolerance. At least in part, these patterns likely reflect adaptive phenotypic plasticity. In summary, our study revealed pronounced plastic and genetic responses, which may indicate high adaptive capacities in our study organism. Whether this may help such species, though, to deal with current climate change needs further investigation, as clinal patterns have typically evolved over long periods.     

Subtle but ubiquitous selection on body size in a natural population of collared flycatchers over 33 years

Understanding the magnitude and long‐term patterns of selection in natural populations is of importance, for example, when analysing the evolutionary impact of climate change. We estimated univariate and multivariate directional, quadratic and correlational selection on four morphological traits (adult wing, tarsus and tail length, body mass) over a time period of 33 years (≈ 19 000 observations) in a nest‐box breeding population of collared flycatchers (Ficedula albicollis). In general, selection was weak in both males and females over the years regardless of fitness measure (fledged young, recruits and survival) with only few cases with statistically significant selection. When data were analysed in a multivariate context and as time series, a number of patterns emerged; there was a consistent, but weak, selection for longer wings in both sexes, selection was stronger on females when the number of fledged young was used as a fitness measure, there were no indications of sexually antagonistic selection, and we found a negative correlation between selection on tarsus and wing length in both sexes but using different fitness measures. Uni‐ and multivariate selection gradients were correlated only for wing length and mass. Multivariate selection gradient vectors were longer than corresponding vector of univariate gradients and had more constrained direction. Correlational selection had little importance. Overall, the fitness surface was more or less flat with few cases of significant curvature, indicating that the adaptive peak with regard to body size in this species is broader than the phenotypic distribution, which has resulted in weak estimates of selection.     

Intrinsic determinants of a population trend in timing of breeding in the wandering albatross

Numerous studies of wild animal species have documented that population level responses to environmental change are underpinned by individual level phenotypic plasticity. However, where the relationship between an individual trait and a climate variable occurs when both show a trend over time, phenotypic plasticity may be confounded by ageing. We investigated between and within individual change in laying date in the wandering albatross Diomedea exulans, a long‐lived species experiencing a dramatic decline in population size. Laying date has advanced over the last three decades. A mean‐centering analysis demonstrated that this pattern was driven by within‐individual changes as opposed to appearance or disappearance of phenotypes. Furthermore, a lack of between individual effect suggested the change resulted from ageing as opposed to phenotypic plasticity. Females varied significantly in rate of advance, such that those with low past reproductive rates exhibited a negative temporal trend in laying date, whereas birds with moderate to high past reproductive performance showed little change. The population trend was therefore driven by a subset with low past breeding success. An analysis of effects of timing of breeding on breeding success revealed stabilizing selection for relative laying date. Furthermore, current breeding success was positively related to past success rate, which suggests that there may be indirect selection against plasticity in this population. Our results show that population trends can arise from individual level change unrelated to prevailing environmental conditions, thus demonstrating the importance of longitudinal analyses in the interpretation of climate change effects.     

How do disturbances and environmental heterogeneity affect the pace of forest distribution shifts under climate change?

Although it is widely predicted that the geographic distributions of tree species and forest types will undergo substantial shifts in future, modelling approaches used to date are largely unable to project the pace at which forest distributions will respond to environmental change. The expansion and contraction of forest distributions act against considerable demographic inertia in the present composition and size‐structure of forest stands as climate‐induced changes in growth, mortality, and recruitment alter population dynamics through time. We aimed to better understand how shifts in forest distributions reflect long‐term changes in tree demographic rates and population dynamics, and how such shifts are influenced by 1) disturbance from forest harvesting and 2) local environmental heterogeneity. Using a simple, data‐constrained gap model, we simulated regional forest dynamics in the eastern United States over the next 500 yr. We then compared the geographic distributions of five different forest types through time under present and altered climatic conditions, in scenarios that variously included and excluded forest harvesting and environmental heterogeneity. Although we held climate fixed after 100 yr, it took another 160 yr after this for these forest types to collectively experience 90% of their eventual climate‐related distribution gains and losses. Competition strongly affected the nature of responses to climate change. Harvesting accelerated and amplified gains by an early‐successional forest type at the expense of a late‐successional one, but these gains did not occur faster than those for other forest types. Environmental heterogeneity had little effect on distribution gains or losses through time. These findings indicate that forest distributions should respond quite slowly to climate change, with the leading and trailing edges of different forest types shifting over a span of centuries. Disturbances can expedite some transitions, but are unlikely to lead to wholesale changes in forest types in the coming decades.     

Temporal patterns of avian body size reflect linear size responses to broadscale environmental change over the last 50 years

Alongside well researched shifts in species' distributions and phenology, reduction in the body size of organisms has been suggested as a third universal response to contemporary climate change. Despite mounting evidence for declining body size, several recent reviews highlight studies reporting increases in body size or no change over time. This variability in response may derive from the geographic scale of contributing studies, masking species‐level responses to broad‐scale environmental change and instead reflecting local influences on single populations. Using museum specimens, we examine temporal patterns of body size of 24 Australian passerine species, sampling multiple populations across the geographic ranges of each species between 1960 and 2007. Generalised additive models indictated that the majority (67%) of species showed important inter‐annual body size variation, and there was striking cross‐species similarity in temporal size patterns. Most displayed near‐linear or linear, unidirectional size trends, suggesting a pervasive and directional change in environmental conditions, consistent with climate change. For species showing linear size responses, the absolute rate of size change ranged between 0.016 and 0.114% of body size (wing length) per year, consistent with studies on other continents. Overall, 38% (9/24) of species showed temporal declines in body size and 21% (5/24) showed increases, consistent with the variability and direction of size responses thus far documented among populations; declining body size is a pervasive response to climate change but it is not universal.     

Evolutionary potential of a widespread clonal grass under changing climate

Adaptive responses are probably the most effective long‐term responses of populations to climate change, but they require sufficient evolutionary potential upon which selection can act. This requires high genetic variance for the traits under selection and low antagonizing genetic covariances between the different traits. Evolutionary potential estimates are still scarce for long‐lived, clonal plants, although these species are predicted to dominate the landscape with climate change. We studied the evolutionary potential of a perennial grass, Festuca rubra, in western Norway, in two controlled environments corresponding to extreme environments in natural populations: cold–dry and warm–wet, the latter being consistent with the climatic predictions for the country. We estimated genetic variances, covariances, selection gradients and response to selection for a wide range of growth, resource acquisition and physiological traits, and compared their estimates between the environments. We showed that the evolutionary potential of F. rubra is high in both environments, and genetic covariances define one main direction along which selection can act with relatively few constraints to selection. The observed response to selection at present is not sufficient to produce genotypes adapted to the predicted climate change under a simple, space for time substitution model. However, the current populations contain genotypes which are pre‐adapted to the new climate, especially for growth and resource acquisition traits. Overall, these results suggest that the present populations of the long‐lived clonal plant may have sufficient evolutionary potential to withstand long‐term climate changes through adaptive responses.     

Response of marine communities to local temperature changes

As global climate change and variability drive shifts in species’ distributions, ecological communities are being reorganized. One approach to understand community change in response to climate change has been to characterize communities by a collective thermal preference, or community temperature index (CTI), and then to compare changes in CTI with changes in temperature. However, important questions remain about whether and how responsive communities are to changes in their local thermal environments. We used CTI to analyze changes in 160 marine assemblages (fish and invertebrates) across the rapidly‐changing Northeast U.S. Continental Shelf Large Marine Ecosystem and calculated expected community change based on historical relationships between species presence and temperature from a separate training dataset. We then compared interannual and long‐term temperature changes with expected community responses and observed community responses over both temporal scales. For these marine communities, we found that community composition as well as composition changes through time could be explained by species associations with bottom temperature. Individual species had non‐linear responses to changes in temperature, and these nonlinearities scaled up to a nonlinear relationship between CTI and temperature. On average, CTI increased by 0.36°C (95% CI: 0.34–0.38°C) for every 1°C increase in bottom temperature, but the relationship between CTI and temperature also depended on community composition. In addition, communities responded more strongly to interannual variation than to long‐term trends in temperature. We recommend that future research into climate‐driven community change accounts for nonlinear responses and examines ecological responses across a range of temporal and geographical scales.     

Climate change and nesting behaviour in vertebrates: a review of the ecological threats and potential for adaptive responses

Nest building is a taxonomically widespread and diverse trait that allows animals to alter local environments to create optimal conditions for offspring development. However, there is growing evidence that climate change is adversely affecting nest‐building in animals directly, for example via sea‐level rises that flood nests, reduced availability of building materials, and suboptimal sex allocation in species exhibiting temperature‐dependent sex determination. Climate change is also affecting nesting species indirectly, via range shifts into suboptimal nesting areas, reduced quality of nest‐building environments, and changes in interactions with nest predators and parasites. The ability of animals to adapt to sustained and rapid environmental change is crucial for the long‐term persistence of many species. Many animals are known to be capable of adjusting nesting behaviour adaptively across environmental gradients and in line with seasonal changes, and this existing plasticity potentially facilitates adaptation to anthropogenic climate change. However, whilst alterations in nesting phenology, site selection and design may facilitate short‐term adaptations, the ability of nest‐building animals to adapt over longer timescales is likely to be influenced by the heritable basis of such behaviour. We urgently need to understand how the behaviour and ecology of nest‐building in animals is affected by climate change, and particularly how altered patterns of nesting behaviour affect individual fitness and population persistence. We begin our review by summarising how predictable variation in environmental conditions influences nest‐building animals, before highlighting the ecological threats facing nest‐building animals experiencing anthropogenic climate change and examining the potential for changes in nest location and/or design to provide adaptive short‐ and long‐term responses to changing environmental conditions. We end by identifying areas that we believe warrant the most urgent attention for further research.     

Experimental evidence of gradual size‐dependent shifts in body size and growth of fish in response to warming

A challenge facing ecologists trying to predict responses to climate change is the few recent analogous conditions to use for comparison. For example, negative relationships between ectotherm body size and temperature are common both across natural thermal gradients and in small‐scale experiments. However, it is unknown if short‐term body size responses are representative of long‐term responses. Moreover, to understand population responses to warming, we must recognize that individual responses to temperature may vary over ontogeny. To enable predictions of how climate warming may affect natural populations, we therefore ask how body size and growth may shift in response to increased temperature over life history, and whether short‐ and long‐term growth responses differ. We addressed these questions using a unique setup with multidecadal artificial heating of an enclosed coastal bay in the Baltic Sea and an adjacent reference area (both with unexploited populations), using before‐after control‐impact paired time‐series analyses. We assembled individual growth trajectories of ~13,000 unique individuals of Eurasian perch and found that body growth increased substantially after warming, but the extent depended on body size: Only among small‐bodied perch did growth increase with temperature. Moreover, the strength of this response gradually increased over the 24 year warming period. Our study offers a unique example of how warming can affect fish populations over multiple generations, resulting in gradual changes in body growth, varying as organisms develop. Although increased juvenile growth rates are in line with predictions of the temperature–size rule, the fact that a larger body size at age was maintained over life history contrasts to that same rule. Because the artificially heated area is a contemporary system mimicking a warmer sea, our findings can aid predictions of fish responses to further warming, taking into account that growth responses may vary both over an individual's life history and over time.     

Grassland responses to increased rainfall depend on the timescale of forcing

Forecasting impacts of future climate change is an important challenge to biologists, both for understanding the consequences of different emissions trajectories and for developing adaptation measures that will minimize biodiversity loss. Existing variation provides a window into the effects of climate on species and ecosystems, but in many places does not encompass the levels or timeframes of forcing expected under directional climatic change. Experiments help us to fill in these uncertainties, simulating directional shifts to examine outcomes of new levels and sustained changes in conditions. Here, we explore the translation between short‐term responses to climate variability and longer‐term trajectories that emerge under directional climatic change. In a decade‐long experiment, we compare effects of short‐term and long‐term forcings across three trophic levels in grassland plots subjected to natural and experimental variation in precipitation. For some biological responses (plant productivity), responses to long‐term extension of the rainy season were consistent with short‐term responses, while for others (plant species richness, abundance of invertebrate herbivores and predators), there was pronounced divergence of long‐term trajectories from short‐term responses. These differences between biological responses mean that sustained directional changes in climate can restructure ecological relationships characterizing a system. Importantly, a positive relationship between plant diversity and productivity turned negative under one scenario of climate change, with a similar change in the relationship between plant productivity and consumer biomass. Inferences from experiments such as this form an important part of wider efforts to understand the complexities of climate change responses.     

Continent‐scale global change attribution in European birds ‐ combining annual and decadal time scales

Species attributes are commonly used to infer impacts of environmental change on multiyear species trends, e.g. decadal changes in population size. However, by themselves attributes are of limited value in global change attribution since they do not measure the changing environment. A broader foundation for attributing species responses to global change may be achieved by complementing an attributes‐based approach by one estimating the relationship between repeated measures of organismal and environmental changes over short time scales. To assess the benefit of this multiscale perspective, we investigate the recent impact of multiple environmental changes on European farmland birds, here focusing on climate change and land use change. We analyze more than 800 time series from 18 countries spanning the past two decades. Analysis of long‐term population growth rates documents simultaneous responses that can be attributed to both climate change and land‐use change, including long‐term increases in populations of hot‐dwelling species and declines in long‐distance migrants and farmland specialists. In contrast, analysis of annual growth rates yield novel insights into the potential mechanisms driving long‐term climate induced change. In particular, we find that birds are affected by winter, spring, and summer conditions depending on the distinct breeding phenology that corresponds to their migratory strategy. Birds in general benefit from higher temperatures or higher primary productivity early on or in the peak of the breeding season with the largest effect sizes observed in cooler parts of species' climatic ranges. Our results document the potential of combining time scales and integrating both species attributes and environmental variables for global change attribution. We suggest such an approach will be of general use when high‐resolution time series are available in large‐scale biodiversity surveys.     

EVOLUTION OF VARIATION AND VARIABILITY UNDER FLUCTUATING,STABILIZING, AND DISRUPTIVE SELECTION

How variation and variability (the capacity to vary) may respond to selection remain open questions. Indeed, effects of different selection regimes on variational properties, such as canalization and developmental stability are under debate. We analyzed the patterns of among‐ and within‐individual variation in two wing‐shape characters in populations of Drosophila melanogaster maintained under fluctuating, disruptive, and stabilizing selection for more than 20 generations. Patterns of variation in wing size, which was not a direct target of selection, were also analyzed. Disruptive selection dramatically increased phenotypic variation in the two shape characters, but left phenotypic variation in wing size unaltered. Fluctuating and stabilizing selection consistently decreased phenotypic variation in all traits. In contrast, within‐individual variation, measured by the level of fluctuating asymmetry, increased for all traits under all selection regimes. These results suggest that canalization and developmental stability are evolvable and presumably controlled by different underlying genetic mechanisms, but the evolutionary responses are not consistent with an adaptive response to selection on variation. Selection also affected patterns of directional asymmetry, although inconsistently across traits and treatments.     

Climate warming is associated with smaller body size and shorter lifespans in moose near their southern range limit

Despite the importance of body size for individual fitness, population dynamics and community dynamics, the influence of climate change on growth and body size is inadequately understood, particularly for long‐lived vertebrates. Although temporal trends in body size have been documented, it remains unclear whether these changes represent the adverse impact of climate change (environmental stress constraining phenotypes) or its mitigation (via phenotypic plasticity or evolution). Concerns have also been raised about whether climate change is indeed the causal agent of these phenotypic shifts, given the length of time‐series analysed and that studies often do not evaluate – and thereby sufficiently rule out – other potential causes. Here, we evaluate evidence for climate‐related changes in adult body size (indexed by skull size) over a 4–decade period for a population of moose (Alces alces) near the southern limit of their range whilst also considering changes in density, predation, and human activities. In particular, we document: (i) a trend of increasing winter temperatures and concurrent decline in skull size (decline of 19% for males and 13% for females) and (ii) evidence of a negative relationship between skull size and winter temperatures during the first year of life. These patterns could be plausibly interpreted as an adaptive phenotypic response to climate warming given that latitudinal/temperature clines are often accepted as evidence of adaptation to local climate. However, we also observed: (iii) that moose with smaller skulls had shorter lifespans, (iv) a reduction in lifespan over the 4‐decade study period, and (v) a negative relationship between lifespan and winter temperatures during the first year of life. Those observations indicate that this phenotypic change is not an adaptive response to climate change. However, this decline in lifespan was not accompanied by an obvious change in population dynamics, suggesting that climate change may affect population dynamics and life‐histories differently.     

Would climate change drive species out of reserves? An assessment of existing reserve‐selection methods

Concern for climate change has not yet been integrated in protocols for reserve selection. However if climate changes as projected, there is a possibility that current reserve‐selection methods might provide solutions that are inadequate to ensure species' long‐term persistence within reserves. We assessed, for the first time, the ability of existing reserve‐selection methods to secure species in a climate‐change context. Six methods using a different combination of criteria (representation, suitability and reserve clustering) are compared. The assessment is carried out using European distributions of 1200 plant species and considering two extreme scenarios of response to climate change: no dispersal and universal dispersal. With our data, 6–11% of species modelled would be potentially lost from selected reserves in a 50‐year period. Measured uncertainties varied in 6% being 1–3% attributed to dispersal assumptions and 2–5% to the choice of reserve‐selection method. Suitability approaches to reserve selection performed best, while reserve clustering performed poorly. We also found that 5% of species modelled would lose their entire climatic envelope in the studied area; 2% of the species modelled would have nonoverlapping distributions; 93% of the species modelled would maintain varying levels of overlapping distributions. We conclude there are opportunities to minimize species' extinctions within reserves but new approaches are needed to account for impacts of climate change on species; especially for those projected to have temporally nonoverlapping distributions.     

The influence of historical climate changes on Southern Ocean marine predator populations: a comparative analysis

The Southern Ocean ecosystem is undergoing rapid physical and biological changes that are likely to have profound implications for higher‐order predators. Here, we compare the long‐term, historical responses of Southern Ocean predators to climate change. We examine palaeoecological evidence for changes in the abundance and distribution of seabirds and marine mammals, and place these into context with palaeoclimate records in order to identify key environmental drivers associated with population changes. Our synthesis revealed two key factors underlying Southern Ocean predator population changes; (i) the availability of ice‐free ground for breeding and (ii) access to productive foraging grounds. The processes of glaciation and sea ice fluctuation were key; the distributions and abundances of elephant seals, snow petrels, gentoo, chinstrap and Adélie penguins all responded strongly to the emergence of new breeding habitat coincident with deglaciation and reductions in sea ice. Access to productive foraging grounds was another limiting factor, with snow petrels, king and emperor penguins all affected by reduced prey availability in the past. Several species were isolated in glacial refugia and there is evidence that refuge populations were supported by polynyas. While the underlying drivers of population change were similar across most Southern Ocean predators, the individual responses of species to environmental change varied because of species specific factors such as dispersal ability and environmental sensitivity. Such interspecific differences are likely to affect the future climate change responses of Southern Ocean marine predators and should be considered in conservation plans. Comparative palaeoecological studies are a valuable source of long‐term data on species’ responses to environmental change that can provide important insights into future climate change responses. This synthesis highlights the importance of protecting productive foraging grounds proximate to breeding locations, as well as the potential role of polynyas as future Southern Ocean refugia.     

Shared morphological consequences of global warming in North American migratory birds

Increasing temperatures associated with climate change are predicted to cause reductions in body size, a key determinant of animal physiology and ecology. Using a four‐decade specimen series of 70 716 individuals of 52 North American migratory bird species, we demonstrate that increasing annual summer temperature over the 40‐year period predicts consistent reductions in body size across these diverse taxa. Concurrently, wing length – an index of body shape that impacts numerous aspects of avian ecology and behaviour – has consistently increased across species. Our findings suggest that warming‐induced body size reduction is a general response to climate change, and reveal a similarly consistent and unexpected shift in body shape. We hypothesise that increasing wing length represents a compensatory adaptation to maintain migration as reductions in body size have increased the metabolic cost of flight. An improved understanding of warming‐induced morphological changes is important for predicting biotic responses to global change.