Snowshoe hares display limited phenotypic plasticity to mismatch in seasonal camouflage

As duration of snow cover decreases owing to climate change, species undergoing seasonal colour moults can become colour mismatched with their background. The immediate adaptive solution to this mismatch is phenotypic plasticity, either in phenology of seasonal colour moults or in behaviours that reduce mismatch or its consequences. We observed nearly 200 snowshoe hares across a wide range of snow conditions and two study sites in Montana, USA, and found minimal plasticity in response to mismatch between coat colour and background. We found that moult phenology varied between study sites, likely due to differences in photoperiod and climate, but was largely fixed within study sites with only minimal plasticity to snow conditions during the spring white-to-brown moult. We also found no evidence that hares modify their behaviour in response to colour mismatch. Hiding and fleeing behaviours and resting spot preference of hares were more affected by variables related to season, site and concealment by vegetation, than by colour mismatch. We conclude that plasticity in moult phenology and behaviours in snowshoe hares is insufficient for adaptation to camouflage mismatch, suggesting that any future adaptation to climate change will require natural selection on moult phenology or behaviour.     

Testing the match–mismatch hypothesis in bighorn sheep in the context of climate change

In species with long gestation, females commit to reproduction several months before parturition. If cues driving conception date are uncoupled from spring conditions, parturition could be mistimed. Mismatch may increase with global change if the rate of temporal changes in autumn cues differs from the rate of change in spring conditions. Using 17 years of data on climate and vegetation phenology, we show that autumn temperature and precipitation, but not vegetation phenology, explain parturition date in bighorn sheep. Although autumn cues drive the timing of conception, they do not predict conditions at parturition in spring. We calculated the mismatch between individual parturition date and spring green-up, assessed whether mismatch increased over time and investigated the consequences of mismatch on lamb neonatal survival, weaning mass and overwinter survival. Mismatch fluctuated over time but showed no temporal trend. Temporal changes in green-up date did not lead to major fitness consequence of mismatch. Detailed data on individually marked animals revealed no effect of mismatch on neonatal or overwinter survival, but lamb weaning mass was negatively affected by mismatch. Capital breeders might be less sensitive to mismatch than income breeders because they are less dependent on daily food acquisition. Herbivores in seasonal environments may access sufficient forage to sustain lactation before or after the spring ‘peak’ green-up, and partly mitigate the consequences of a mismatch. Thus, the effect of phenological mismatch on fitness may be affected by species life history, highlighting the complexity in quantifying trophic mismatches in the context of climate change.     

The importance of pre-and postcopulatory sexual selection promoting adaptation to increasing temperatures

Global temperatures are increasing rapidly affecting species globally. Understanding if and how different species can adapt fast enough to keep up with increasing temperatures is of vital importance. One mechanism that can accelerate adaptation and promote evolutionary rescue is sexual selection. Two different mechanisms by which sexual selection can facilitate adaptation are pre- and postcopulatory sexual selection. However, the relative effects of these different forms of sexual selection in promoting adaptation are unknown. Here, we present the results from an experimental study in which we exposed fruit flies Drosophila melanogaster to either no mate choice or 1 of 2 different sexual selection regimes (pre- and postcopulatory sexual selection) for 6 generations, under different thermal regimes. Populations showed evidence of thermal adaptation under precopulatory sexual selection, but this effect was not detected in the postcopulatory sexual selection and the no choice mating regime. We further demonstrate that sexual dimorphism decreased when flies evolved under increasing temperatures, consistent with recent theory predicting more sexually concordant selection under environmental stress. Our results suggest an important role for precopulatory sexual selection in promoting thermal adaptation and evolutionary rescue.     

Eco‐evolution on the edge during climate change

We urgently need to predict species responses to climate change to minimize future biodiversity loss and ensure we do not waste limited resources on ineffective conservation strategies. Currently, most predictions of species responses to climate change ignore the potential for evolution. However, evolution can alter species ecological responses, and different aspects of evolution and ecology can interact to produce complex eco‐evolutionary dynamics under climate change. Here we review how evolution could alter ecological responses to climate change on species warm and cool range margins, where evolution could be especially important. We discuss different aspects of evolution in isolation, and then synthesize results to consider how multiple evolutionary processes might interact and affect conservation strategies. On species cool range margins, the evolution of dispersal could increase range expansion rates and allow species to adapt to novel conditions in their new range. However, low genetic variation and genetic drift in small range‐front populations could also slow or halt range expansions. Together, these eco‐evolutionary effects could cause a three‐step, stop‐and‐go expansion pattern for many species. On warm range margins, isolation among populations could maintain high genetic variation that facilitates evolution to novel climates and allows species to persist longer than expected without evolution. This ‘evolutionary extinction debt’ could then prevent other species from shifting their ranges. However, as climate change increases isolation among populations, increasing dispersal mortality could select for decreased dispersal and cause rapid range contractions. Some of these eco‐evolutionary dynamics could explain why many species are not responding to climate change as predicted. We conclude by suggesting that resurveying historical studies that measured trait frequencies, the strength of selection, or heritabilities could be an efficient way to increase our eco‐evolutionary knowledge in climate change biology.     

Spatio‐temporal variation in fitness responses to contrasting environments in Arabidopsis thaliana

The evolutionary response of organisms to global climate change is expected to be strongly conditioned by preexisting standing genetic variation. In addition, natural selection imposed by global climate change on fitness‐related traits can be heterogeneous over time. We estimated selection of life‐history traits of an entire genetic lineage of the plant Arabidopsis thaliana occurring in north‐western Iberian Peninsula that were transplanted over multiple years into two environmentally contrasting field sites in southern Spain, as southern environments are expected to move progressively northwards with climate change in the Iberian Peninsula. The results indicated that natural selection on flowering time prevailed over that on recruitment. Selection favored early flowering in six of eight experiments and late flowering in the other two. Such heterogeneity of selection for flowering time might be a powerful mechanism for maintaining genetic diversity in the long run. We also found that north‐western A. thaliana accessions from warmer environments exhibited higher fitness and higher phenotypic plasticity for flowering time in southern experimental facilities. Overall, our transplant experiments suggested that north‐western Iberian A. thaliana has the means to cope with increasingly warmer environments in the region as predicted by trends in global climate change models.     

Compensating for climate change–induced cue‐environment mismatches: evidence for contemporary evolution of a photoperiodic reaction norm in Colias butterflies

Anthropogenic climate change alters seasonal conditions without altering photoperiod and can thus create a cue‐environment mismatch for organisms that use photoperiod as a cue for seasonal plasticity. We investigated whether evolution of the photoperiodic reaction norm has compensated for this mismatch in Colias eurytheme. This butterfly’s wing melanization has a thermoregulatory function and changes seasonally. In 1971, Hoffmann quantified how larval photoperiod determines adult wing melanization. We recreated his experiment 47 years later using a contemporary population. Comparing our results to his, we found decreased melanization at short photoperiods but no change in melanization at long photoperiods, which is consistent with the greater increase in spring than summer temperatures recorded for this region. Our study shows that evolution can help correct cue‐environment mismatches but not in the same way under all conditions. Studies of contemporary evolution may miss important changes if they focus on only a limited range of conditions.     

Genetic structure of a recent climate change‐driven range extension

The life‐history strategies of some species make them strong candidates for rapid exploitation of novel habitat under new climate regimes. Some early‐responding species may be considered invasive, and negatively impact on ‘naïve’ ecosystems. The barrens‐forming sea urchin Centrostephanus rodgersii is one such species, having a high dispersal capability and a high‐latitude range margin limited only by a developmental temperature threshold. Within this species’ range in eastern Australian waters, sea temperatures have increased at greater than double the global average rate. The coinciding poleward range extension of C. rodgersii has caused major ecological changes, threatening reef biodiversity and fisheries productivity. We investigated microsatellite diversity and population structure associated with range expansion by this species. Generalized linear model analyses revealed no reduction in genetic diversity in the newly colonized region. A ‘seascape genetics’ analysis of genetic distances found no spatial genetic structure associated with the range extension. The distinctive genetic characteristic of the extension zone populations was reduced population‐specific F_ST, consistent with very rapid population expansion. Demographic and genetic simulations support our inference of high connectivity between pre‐ and post‐extension zones. Thus, the range shift appears to be a poleward extension of the highly‐connected rangewide population of C. rodgersii. This is consistent with advection of larvae by the intensified warm water East Australian current, which has also increased Tasmanian Sea temperatures above the species’ lower developmental threshold. Thus, ocean circulation changes have improved the climatic suitability of novel habitat for C. rodgersii and provided the supply of recruits necessary for colonization.     

Long‐term monitoring of an amphibian community after a climate change‐ and infectious disease‐driven species extirpation

Infectious disease and climate change are considered major threats to biodiversity and act as drivers behind the global amphibian decline. This is, to a large extent, based on short‐term studies that are designed to detect the immediate and strongest biodiversity responses to a threatening process. What few long‐term studies are available, although typically focused on single species, report outcomes that often diverge significantly from the short‐term species responses. Here, we report the results of an 18‐year survey of an amphibian community exposed to both climate warming and the emergence of lethal chytridiomycosis. Our study shows that the impacts of infectious disease are ongoing but restricted to two out of nine species that form the community, despite the fact all species can become infected with the fungus. Climate warming appears to be affecting four out of the nine species, but the response of three of these is an increase in abundance. Our study supports a decreasing role of infectious disease on the community, and an increasing and currently positive effect of climate warming. We caution that if the warming trends continue, the net positive effect will turn negative as amphibian breeding habitat becomes unavailable as water bodies dry, a pattern that already may be underway.     

Herbarium specimens reveal the footprint of climate change on flowering trends across north‐central North America

Shifting flowering phenology with rising temperatures is occurring worldwide, but the rarity of co‐occurring long‐term observational and temperature records has hindered the evaluation of phenological responsiveness in many species and across large spatial scales. We used herbarium specimens combined with historic temperature data to examine the impact of climate change on flowering trends in 141 species collected across 116,000 km² in north‐central North America. On average, date of maximum flowering advanced 2.4 days °C⁻¹, although species‐specific responses varied from − 13.5 to + 7.3 days °C⁻¹. Plant functional types exhibited distinct patterns of phenological responsiveness with significant differences between native and introduced species, among flowering seasons, and between wind‐ and biotically pollinated species. This study is the first to assess large‐scale patterns of phenological responsiveness with broad species representation and is an important step towards understanding current and future impacts of climate change on species performance and biodiversity.     

Climate change‐induced migration patterns and extinction risks of Theaceae species in China

Theaceae, an economically important angiosperm family, is widely distributed in tropical and subtropical forests in Asia. In China, Theaceae has particularly high abundances and endemism, comprising ~75% of the total genera and ~46% of the total species worldwide. Therefore, predicting the response of Theaceae species to climate change is vital. In this study, we collected distribution data for 200 wild Theaceae species in China, and predicted their distribution patterns under current and future climactic conditions by species distribution modeling (SDM). We revealed that Theaceae species richness is highest in southeastern China and on Hainan Island, reaching its highest value (137 species) in Fujian Province. According to the IUCN Red List criteria for assessing species threat levels under two dispersal assumptions (no dispersal and full dispersal), we evaluated the conservation status of all Theaceae species by calculating loss of suitable habitat under future climate scenarios. We predicted that nine additional species will become threatened due to climate change in the future; one species will be classified as critically endangered (CR), two as endangered (EN), and six as vulnerable (VU). Given their extinction risks associated with climate change, we recommended that these species be added to the Red List. Our investigation of migration patterns revealed regional differences in the number of emigrant, immigrant, and persistent species, indicating the need for targeted conservation strategies. Regions containing numerous emigrants are concentrated in Northern Taiwan and coastal regions of Zhejiang and Fujian provinces, while regions containing numerous immigrants include central Sichuan Province, the southeastern Tibet Autonomous Region, southwest Yunnan Province, northwest Sichuan Province, and the junction of Guangxi and Hunan provinces. Lastly, regions containing persistent species are widely distributed in southern China. Importantly, regions with high species turnover are located on the northern border of the entire Theaceae species distribution ranges owing to upwards migration; these regions are considered most sensitive to climate change and conservation planning should therefore be prioritized here. This study will contribute valuable information for reducing the negative impacts of climate change on Theaceae species, which will ultimately improve biodiversity conservation efficiency.     

Fitness declines towards range limits and local adaptation to climate affect dispersal evolution during climate‐induced range shifts

Dispersal ability will largely determine whether species track their climatic niches during climate change, a process especially important for populations at contracting (low‐latitude/low‐elevation) range limits that otherwise risk extinction. We investigate whether dispersal evolution at contracting range limits is facilitated by two processes that potentially enable edge populations to experience and adjust to the effects of climate deterioration before they cause extinction: (i) climate‐induced fitness declines towards range limits and (ii) local adaptation to a shifting climate gradient. We simulate a species distributed continuously along a temperature gradient using a spatially explicit, individual‐based model. We compare range‐wide dispersal evolution during climate stability vs. directional climate change, with uniform fitness vs. fitness that declines towards range limits (RLs), and for a single climate genotype vs. multiple genotypes locally adapted to temperature. During climate stability, dispersal decreased towards RLs when fitness was uniform, but increased when fitness declined towards RLs, due to highly dispersive genotypes maintaining sink populations at RLs, increased kin selection in smaller populations, and an emergent fitness asymmetry that favoured dispersal in low‐quality habitat. However, this initial dispersal advantage at low‐fitness RLs did not facilitate climate tracking, as it was outweighed by an increased probability of extinction. Locally adapted genotypes benefited from staying close to their climate optima; this selected against dispersal under stable climates but for increased dispersal throughout shifting ranges, compared to cases without local adaptation. Dispersal increased at expanding RLs in most scenarios, but only increased at the range centre and contracting RLs given local adaptation to climate.     

Signatures of diversifying selection at EST‐SSR loci and association with climate in natural Eucalyptus populations

Understanding the environmental parameters that drive adaptation among populations is important in predicting how species may respond to global climatic changes and how gene pools might be managed to conserve adaptive genetic diversity. Here, we used Bayesian F_ST outlier tests and allele–climate association analyses to reveal two Eucalyptus EST‐SSR loci as strong candidates for diversifying selection in natural populations of a southwestern Australian forest tree, Eucalyptus gomphocephala (Myrtaceae). The Eucalyptus homolog of a CONSTANS‐like gene was an F_ST outlier, and allelic variation showed significant latitudinal clinal associations with annual and winter solar radiation, potential evaporation, summer precipitation and aridity. A second F_ST outlier locus, homologous to quinone oxidoreductase, was significantly associated with measures of temperature range, high summer temperature and summer solar radiation, with important implications for predicting the effect of temperature on natural populations in the context of climate change. We complemented these data with investigations into neutral population genetic structure and diversity throughout the species range. This study provides an investigation into selection signatures at gene‐homologous EST‐SSRs in natural Eucalyptus populations, and contributes to our understanding of the relationship between climate and adaptive genetic variation, informing the conservation of both putatively neutral and adaptive components of genetic diversity.     

Climate change‐induced vegetation change as a driver of increased subarctic biogenic volatile organic compound emissions

Emissions of biogenic volatile organic compounds (BVOCs) have been earlier shown to be highly temperature sensitive in subarctic ecosystems. As these ecosystems experience rapidly advancing pronounced climate warming, we aimed to investigate how warming affects the BVOC emissions in the long term (up to 13 treatment years). We also aimed to assess whether the increased litterfall resulting from the vegetation changes in the warming subarctic would affect the emissions. The study was conducted in a field experiment with factorial open‐top chamber warming and annual litter addition treatments on subarctic heath in Abisko, northern Sweden. After 11 and 13 treatment years, BVOCs were sampled from plant communities in the experimental plots using a push–pull enclosure technique and collection into adsorbent cartridges during the growing season and analyzed with gas chromatography–mass spectrometry. Plant species coverage in the plots was analyzed by the point intercept method. Warming by 2 °C caused a 2‐fold increase in monoterpene and 5‐fold increase in sesquiterpene emissions, averaged over all measurements. When the momentary effect of temperature was diminished by standardization of emissions to a fixed temperature, warming still had a significant effect suggesting that emissions were also indirectly increased. This indirect increase appeared to result from increased plant coverage and changes in vegetation composition. The litter addition treatment also caused significant increases in the emission rates of some BVOC groups, especially when combined with warming. The combined treatment had both the largest vegetation changes and the highest BVOC emissions. The increased emissions under litter addition were probably a result of a changed vegetation composition due to alleviated nutrient limitation and stimulated microbial production of BVOCs. We suggest that the changes in the subarctic vegetation composition induced by climate warming will be the major factor indirectly affecting the BVOC emission potentials and composition.     

Behavioural responses to human‐induced environmental change

The initial response of individuals to human‐induced environmental change is often behavioural. This can improve the performance of individuals under sudden, large‐scale perturbations and maintain viable populations. The response can also give additional time for genetic changes to arise and, hence, facilitate adaptation to new conditions. On the other hand, maladaptive responses, which reduce individual fitness, may occur when individuals encounter conditions that the population has not experienced during its evolutionary history, which can decrease population viability. A growing number of studies find human disturbances to induce behavioural responses, both directly and by altering factors that influence fitness. Common causes of behavioural responses are changes in the transmission of information, the concentration of endocrine disrupters, the availability of resources, the possibility of dispersal, and the abundance of interacting species. Frequent responses are alterations in habitat choice, movements, foraging, social behaviour and reproductive behaviour. Behavioural responses depend on the genetically determined reaction norm of the individuals, which evolves over generations. Populations first respond with individual behavioural plasticity, whereafter changes may arise through innovations and the social transmission of behavioural patterns within and across generations, and, finally, by evolution of the behavioural response over generations. Only a restricted number of species show behavioural adaptations that make them thrive in severely disturbed environments. Hence, rapid human‐induced disturbances often decrease the diversity of native species, while facilitating the spread of invasive species with highly plastic behaviours. Consequently, behavioural responses to human‐induced environmental change can have profound effects on the distribution, adaptation, speciation and extinction of populations and, hence, on biodiversity. A better understanding of the mechanisms of behavioural responses and their causes and consequences could improve our ability to predict the effects of human‐induced environmental change on individual species and on biodiversity.     

Long‐term changes in autumn migration dates at the Strait of Gibraltar reflect population trends of soaring birds

A growing body of work shows that climate change is the cause of a number of directional shifts in the spring phenology of migratory birds. However, changes in autumn phenology are well studied and their consistency across species, as well as their link with population trends, remains uncertain. We investigate changes in the autumn migration dates of 11 species of soaring birds over the Strait of Gibraltar over a 16‐year period. Using models corrected for phylogeny, we assessed whether ecological and morphological characteristics, as well as population trends, account for interspecific shifts in migration times. We recorded different phenological changes in different periods of the migration season and suggest that these differences are due to age‐dependent responses. The variable best predicting advances in migration dates was population trend: species that did not advance their autumn migration dates were those showing a decline in their European breeding populations. We repeated our tests on a dataset representing the migration date of soaring birds across the Pyrenees Mountains and found that population trends at this site also predicted phenological shifts. Our results suggest that flexibility in migratory strategy and population trends may be related, such that different adaptive capacity in migration timing may be more relevant than other ecological traits in determining the conservation status of migratory birds in Europe and perhaps other regions.     

Differences in photoperiod‐induced diapause plasticity among different populations of the bark beetle Ips typographus and its predator Thanasimus formicarius

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Estimation of fitness from energetics and life‐history data: An example using mussels

Changing environments have the potential to alter the fitness of organisms through effects on components of fitness such as energy acquisition, metabolic cost, growth rate, survivorship, and reproductive output. Organisms, on the other hand, can alter aspects of their physiology and life histories through phenotypic plasticity as well as through genetic change in populations (selection). Researchers examining the effects of environmental variables frequently concentrate on individual components of fitness, although methods exist to combine these into a population level estimate of average fitness, as the per capita rate of population growth for a set of identical individuals with a particular set of traits. Recent advances in energetic modeling have provided excellent data on energy intake and costs leading to growth, reproduction, and other life‐history parameters; these in turn have consequences for survivorship at all life‐history stages, and thus for fitness. Components of fitness alone (performance measures) are useful in determining organism response to changing conditions, but are often not good predictors of fitness; they can differ in both form and magnitude, as demonstrated in our model. Here, we combine an energetics model for growth and allocation with a matrix model that calculates population growth rate for a group of individuals with a particular set of traits. We use intertidal mussels as an example, because data exist for some of the important energetic and life‐history parameters, and because there is a hypothesized energetic trade‐off between byssus production (affecting survivorship), and energy used for growth and reproduction. The model shows exactly how strong this trade‐off is in terms of overall fitness, and it illustrates conditions where fitness components are good predictors of actual fitness, and cases where they are not. In addition, the model is used to examine the effects of environmental change on this trade‐off and on both fitness and on individual fitness components.     

Direct benefits of choosing a high‐fitness mate can offset the indirect costs associated with intralocus sexual conflict

Intralocus sexual conflict generates a cost to mate choice: high‐fitness partners transmit genetic variation that confers lower fitness to offspring of the opposite sex. Our earlier work in the fruit fly, Drosophila melanogaster, revealed that these indirect genetic costs were sufficient to reverse potential “good genes” benefits of sexual selection. However, mate choice can also confer direct fitness benefits by inducing larger numbers of progeny. Here, we consider whether direct benefits through enhanced fertility could offset the costs associated with intralocus sexual conflict in D. melanogaster. Using hemiclonal analysis, we found that females mated to high‐fitness males produced 11% more offspring compared to those mated to low‐fitness males, and high‐fitness females produced 34% more offspring than low‐fitness females. These direct benefits more than offset the reduction in offspring fitness caused by intralocus sexual conflict, creating a net fitness benefit for each sex to pairing with a high‐fitness partner. Our findings highlight the need to consider both direct and indirect effects when investigating the fitness impacts of mate choice. Direct fitness benefits may shelter sexually antagonistic alleles from selection, suggesting a novel mechanism for the maintenance of fitness variation.     

Structural and functional responses of plant communities to climate change‐mediated alterations in the hydrology of riparian areas in temperate Europe

The hydrology of riparian areas changes rapidly these years because of climate change‐mediated alterations in precipitation patterns. In this study, we used a large‐scale in situ experimental approach to explore effects of drought and flooding on plant taxonomic diversity and functional trait composition in riparian areas in temperate Europe. We found significant effects of flooding and drought in all study areas, the effects being most pronounced under flooded conditions. In near‐stream areas, taxonomic diversity initially declined in response to both drought and flooding (although not significantly so in all years) and remained stable under drought conditions, whereas the decline continued under flooded conditions. For most traits, we found clear indications that the functional diversity also declined under flooded conditions, particularly in near‐stream areas, indicating that fewer strategies succeeded under flooded conditions. Consistent changes in community mean trait values were also identified, but fewer than expected. This can have several, not mutually exclusive, explanations. First, different adaptive strategies may coexist in a community. Second, intraspecific variability was not considered for any of the traits. For example, many species can elongate shoots and petioles that enable them to survive shallow, prolonged flooding but such abilities will not be captured when applying mean trait values. Third, we only followed the communities for 3 years. Flooding excludes species intolerant of the altered hydrology, whereas the establishment of new species relies on time‐dependent processes, for instance the dispersal and establishment of species within the areas. We expect that altered precipitation patterns will have profound consequences for riparian vegetation in temperate Europe. Riparian areas will experience loss of taxonomic and functional diversity and, over time, possibly also alterations in community trait responses that may have cascading effects on ecosystem functioning.