Marine assemblages respond rapidly to winter climate variability

Even species within the same assemblage have varied responses to climate change, and there is a poor understanding for why some taxa are more sensitive to climate than others. In addition, multiple mechanisms can drive species' responses, and responses may be specific to certain life stages or times of year. To test how marine species respond to climate variability, we analyzed 73 diverse taxa off the southeast US coast in 26 years of scientific trawl survey data and determined how changes in distribution and biomass relate to temperature. We found that winter temperatures were particularly useful for explaining interannual variation in species' distribution and biomass, although the direction and magnitude of the response varied among species from strongly negative, to little response, to strongly positive. Across species, the response to winter temperature varied greatly, with much of this variation being explained by thermal preference. A separate analysis of annual commercial fishery landings revealed that winter temperatures may also impact several important fisheries in the southeast United States. Based on the life stages of the species surveyed, winter temperature appears to act through overwinter mortality of juveniles or as a cue for migration timing. We predict that this assemblage will be responsive to projected increases in temperature and that winter temperature may be broadly important for species relationships with climate on a global scale.     

Differential response to abiotic stress controls species distributions at biogeographic transition zones

Understanding range limits is critical to predicting species responses to climate change. Subtropical environments, where many species overlap at their range margins, are cooler, more light‐limited and variable than tropical environments. It is thus likely that species respond variably to these multi‐stressor regimes and that factors other than mean climatic conditions drive biodiversity patterns. Here, we tested these hypotheses for scleractinian corals at their high‐latitude range limits in eastern Australia and investigated the role of mean climatic conditions and of parameters linked to abiotic stress in explaining the distribution and abundance of different groups of species. We found that environmental drivers varied among taxa and were predominantly linked to abiotic stress. The distribution and abundance of tropical species and gradients in species richness (alpha diversity) and turnover (beta diversity) were best explained by light limitation, whereas minimum temperatures and temperature fluctuations best explained gradients in subtropical species, species nestedness and functional diversity. Variation in community structure (considering species composition and abundance) was most closely linked to the combined thermal and light regime. Our study demonstrates the role of abiotic stress in controlling the distribution of species towards their high‐latitude range limits and suggests that, at biogeographic transition zones, robust predictions of the impacts of climate change require approaches that account for various aspects of physiological stress and for species abundances and characteristics. These findings support the hypothesis that abiotic stress controls high‐latitude range limits and caution that projections solely based on mean temperature could underestimate species’ vulnerabilities to climate change.     

The impact of climate change measured at relevant spatial scales: new hope for tropical lizards

Much attention has been given to recent predictions that widespread extinctions of tropical ectotherms, and tropical forest lizards in particular, will result from anthropogenic climate change. Most of these predictions, however, are based on environmental temperature data measured at a maximum resolution of 1 km², whereas individuals of most species experience thermal variation on a much finer scale. To address this disconnect, we combined thermal performance curves for five populations of Anolis lizard from the Bay Islands of Honduras with high‐resolution temperature distributions generated from physical models. Previous research has suggested that open‐habitat species are likely to invade forest habitat and drive forest species to extinction. We test this hypothesis, and compare the vulnerabilities of closely related, but allopatric, forest species. Our data suggest that the open‐habitat populations we studied will not invade forest habitat and may actually benefit from predicted warming for many decades. Conversely, one of the forest species we studied should experience reduced activity time as a result of warming, while two others are unlikely to experience a significant decline in performance. Our results suggest that global‐scale predictions generated using low‐resolution temperature data may overestimate the vulnerability of many tropical ectotherms to climate change.     

Caribbean mesophotic coral ecosystems are unlikely climate change refugia

Deeper coral reefs experience reduced temperatures and light and are often shielded from localized anthropogenic stressors such as pollution and fishing. The deep reef refugia hypothesis posits that light‐dependent stony coral species at deeper depths are buffered from thermal stress and will avoid bleaching‐related mass mortalities caused by increasing sea surface temperatures under climate change. This hypothesis has not been tested because data collection on deeper coral reefs is difficult. Here we show that deeper (mesophotic) reefs, 30–75 m depth, in the Caribbean are not refugia because they have lower bleaching threshold temperatures than shallow reefs. Over two thermal stress events, mesophotic reef bleaching was driven by a bleaching threshold that declines 0.26 °C every +10 m depth. Thus, the main premise of the deep reef refugia hypothesis that cooler environments are protective is incorrect; any increase in temperatures above the local mean warmest conditions can lead to thermal stress and bleaching. Thus, relatively cooler temperatures can no longer be considered a de facto refugium for corals and it is likely that many deeper coral reefs are as vulnerable to climate change as shallow water reefs.     

Calling behaviour under climate change: geographical and seasonal variation of calling temperatures in ectotherms

Calling behaviour is strongly temperature‐dependent and critical for sexual selection and reproduction in a variety of ectothermic taxa, including anuran amphibians, which are the most globally threatened vertebrates. However, few studies have explored how species respond to distinct thermal environments at time of displaying calling behaviour, and thus it is still unknown whether ongoing climate change might compromise the performance of calling activity in ectotherms. Here, we used new audio‐trapping techniques (automated sound recording and detection systems) between 2006 and 2009 to examine annual calling temperatures of five temperate anurans and their patterns of geographical and seasonal variation at the thermal extremes of species ranges, providing insights into the thermal breadths of calling activity of species, and the mechanisms that enable ectotherms to adjust to changing thermal environments. All species showed wide thermal breadths during calling behaviour (above 15 °C) and increases in calling temperatures in extremely warm populations and seasons. Thereby, calling temperatures differed both geographically and seasonally, both in terrestrial and aquatic species, and were 8–22 °C below the specific upper critical thermal limits (CT_max) and strongly associated with the potential temperatures of each thermal environment (operative temperatures during the potential period of breeding). This suggests that calling behaviour in ectotherms may take place at population‐specific thermal ranges, diverging when species are subjected to distinct thermal environments, and might imply plasticity of thermal adjustment mechanisms (seasonal and developmental acclimation) that supply species with means of coping with climate change. Furthermore, the thermal thresholds of calling at the onset of the breeding season were dissimilar between conspecific populations, suggesting that other factors besides temperature are needed to trigger the onset of reproduction. Our findings imply that global warming would not directly inhibit calling behaviour in the study species, although might affect other temperature‐dependent features of their acoustic communication system.     

Modeling plant species distributions under future climates: how fine scale do climate projections need to be?

Recent studies suggest that species distribution models (SDMs) based on fine‐scale climate data may provide markedly different estimates of climate‐change impacts than coarse‐scale models. However, these studies disagree in their conclusions of how scale influences projected species distributions. In rugged terrain, coarse‐scale climate grids may not capture topographically controlled climate variation at the scale that constitutes microhabitat or refugia for some species. Although finer scale data are therefore considered to better reflect climatic conditions experienced by species, there have been few formal analyses of how modeled distributions differ with scale. We modeled distributions for 52 plant species endemic to the California Floristic Province of different life forms and range sizes under recent and future climate across a 2000‐fold range of spatial scales (0.008–16 km²). We produced unique current and future climate datasets by separately downscaling 4 km climate models to three finer resolutions based on 800, 270, and 90 m digital elevation models and deriving bioclimatic predictors from them. As climate‐data resolution became coarser, SDMs predicted larger habitat area with diminishing spatial congruence between fine‐ and coarse‐scale predictions. These trends were most pronounced at the coarsest resolutions and depended on climate scenario and species' range size. On average, SDMs projected onto 4 km climate data predicted 42% more stable habitat (the amount of spatial overlap between predicted current and future climatically suitable habitat) compared with 800 m data. We found only modest agreement between areas predicted to be stable by 90 m models generalized to 4 km grids compared with areas classified as stable based on 4 km models, suggesting that some climate refugia captured at finer scales may be missed using coarser scale data. These differences in projected locations of habitat change may have more serious implications than net habitat area when predictive maps form the basis of conservation decision making.     

Experimental whole‐stream warming alters community size structure

How ecological communities respond to predicted increases in temperature will determine the extent to which Earth's biodiversity and ecosystem functioning can be maintained into a warmer future. Warming is predicted to alter the structure of natural communities, but robust tests of such predictions require appropriate large‐scale manipulations of intact, natural habitat that is open to dispersal processes via exchange with regional species pools. Here, we report results of a two‐year whole‐stream warming experiment that shifted invertebrate assemblage structure via unanticipated mechanisms, while still conforming to community‐level metabolic theory. While warming by 3.8 °C decreased invertebrate abundance in the experimental stream by 60% relative to a reference stream, total invertebrate biomass was unchanged. Associated shifts in invertebrate assemblage structure were driven by the arrival of new taxa and a higher proportion of large, warm‐adapted species (i.e., snails and predatory dipterans) relative to small‐bodied, cold‐adapted taxa (e.g., chironomids and oligochaetes). Experimental warming consequently shifted assemblage size spectra in ways that were unexpected, but consistent with thermal optima of taxa in the regional species pool. Higher temperatures increased community‐level energy demand, which was presumably satisfied by higher primary production after warming. Our experiment demonstrates how warming reassembles communities within the constraints of energy supply via regional exchange of species that differ in thermal physiological traits. Similar responses will likely mediate impacts of anthropogenic warming on biodiversity and ecosystem function across all ecological communities.     

The shape of abundance distributions across temperature gradients in reef fishes

Improving predictions of ecological responses to climate change requires understanding how local abundance relates to temperature gradients, yet many factors influence local abundance in wild populations. We evaluated the shape of thermal‐abundance distributions using 98 422 abundance estimates of 702 reef fish species worldwide. We found that curved ceilings in local abundance related to sea temperatures for most species, where local abundance declined from realised thermal ‘optima’ towards warmer and cooler environments. Although generally supporting the abundant‐centre hypothesis, many species also displayed asymmetrical thermal‐abundance distributions. For many tropical species, abundances did not decline at warm distribution edges due to an unavailability of warmer environments at the equator. Habitat transitions from coral to macroalgal dominance in subtropical zones also influenced abundance distribution shapes. By quantifying the factors constraining species’ abundance, we provide an important empirical basis for improving predictions of community re‐structuring in a warmer world.     

Responses of arthropod populations to warming depend on latitude: evidence from urban heat islands

Biological effects of climate change are expected to vary geographically, with a strong signature of latitude. For ectothermic animals, there is systematic latitudinal variation in the relationship between climate and thermal performance curves, which describe the relationship between temperature and an organism's fitness. Here, we ask whether these documented latitudinal patterns can be generalized to predict arthropod responses to warming across mid‐ and high temperate latitudes, for taxa whose thermal physiology has not been measured. To address this question, we used a novel natural experiment consisting of a series of urban warming gradients at different latitudes. Specifically, we sampled arthropods from a single common street tree species across temperature gradients in four US cities, located from 35.8 to 42.4° latitude. We captured 6746 arthropods in 34 families from 111 sites that varied in summer average temperature by 1.7–3.4 °C within each city. Arthropod responses to warming within each city were characterized as Poisson regression coefficients describing change in abundance per °C for each family. Family responses in the two midlatitude cities were heterogeneous, including significantly negative and positive effects, while those in high‐latitude cities varied no more than expected by chance within each city. We expected high‐latitude taxa to increase in abundance with warming, and they did so in one of the two high‐latitude cities; in the other, Queens (New York City), most taxa declined with warming, perhaps due to habitat loss that was correlated with warming in this city. With the exception of Queens, patterns of family responses to warming were consistent with predictions based on known latitudinal patterns in arthropod physiology relative to regional climate. Heterogeneous responses in midlatitudes may be ecologically disruptive if interacting taxa respond oppositely to warming.     

Range expansion through fragmented landscapes under a variable climate

Ecological responses to climate change may depend on complex patterns of variability in weather and local microclimate that overlay global increases in mean temperature. Here, we show that high‐resolution temporal and spatial variability in temperature drives the dynamics of range expansion for an exemplar species, the butterfly Hesperia comma. Using fine‐resolution (5 m) models of vegetation surface microclimate, we estimate the thermal suitability of 906 habitat patches at the species' range margin for 27 years. Population and metapopulation models that incorporate this dynamic microclimate surface improve predictions of observed annual changes to population density and patch occupancy dynamics during the species' range expansion from 1982 to 2009. Our findings reveal how fine‐scale, short‐term environmental variability drives rates and patterns of range expansion through spatially localised, intermittent episodes of expansion and contraction. Incorporating dynamic microclimates can thus improve models of species range shifts at spatial and temporal scales relevant to conservation interventions.     

Global variation in thermal tolerances and vulnerability of endotherms to climate change

The relationships among species'' physiological capacities and the geographical variation of ambient climate are of key importance to understanding the distribution of life on the Earth. Furthermore, predictions of how species will respond to climate change will profit from the explicit consideration of their physiological tolerances. The climatic variability hypothesis, which predicts that climatic tolerances are broader in more variable climates, provides an analytical framework for studying these relationships between physiology and biogeography. However, direct empirical support for the hypothesis is mostly lacking for endotherms, and few studies have tried to integrate physiological data into assessments of species'' climatic vulnerability at the global scale. Here, we test the climatic variability hypothesis for endotherms, with a comprehensive dataset on thermal tolerances derived from physiological experiments, and use these data to assess the vulnerability of species to projected climate change. We find the expected relationship between thermal tolerance and ambient climatic variability in birds, but not in mammals—a contrast possibly resulting from different adaptation strategies to ambient climate via behaviour, morphology or physiology. We show that currently most of the species are experiencing ambient temperatures well within their tolerance limits and that in the future many species may be able to tolerate projected temperature increases across significant proportions of their distributions. However, our findings also underline the high vulnerability of tropical regions to changes in temperature and other threats of anthropogenic global changes. Our study demonstrates that a better understanding of the interplay among species'' physiology and the geography of climate change will advance assessments of species'' vulnerability to climate change.     

The complex drivers of thermal acclimation and breadth in ectotherms

Thermal acclimation capacity, the degree to which organisms can alter their optimal performance temperature and critical thermal limits with changing temperatures, reflects their ability to respond to temperature variability and thus might be important for coping with global climate change. Here, we combine simulation modelling with analysis of published data on thermal acclimation and breadth (range of temperatures over which organisms perform well) to develop a framework for predicting thermal plasticity across taxa, latitudes, body sizes, traits, habitats and methodological factors. Our synthesis includes > 2000 measures of acclimation capacities from > 500 species of ectotherms spanning fungi, invertebrates, and vertebrates from freshwater, marine and terrestrial habitats. We find that body size, latitude, and methodological factors often interact to shape acclimation responses and that acclimation rate scales negatively with body size, contributing to a general negative association between body size and thermal breadth across species. Additionally, we reveal that acclimation capacity increases with body size, increases with latitude (to mid‐latitudinal zones) and seasonality for smaller but not larger organisms, decreases with thermal safety margin (upper lethal temperature minus maximum environmental temperatures), and is regularly underestimated because of experimental artefacts. We then demonstrate that our framework can predict the contribution of acclimation plasticity to the IUCN threat status of amphibians globally, suggesting that phenotypic plasticity is already buffering some species from climate change.     

Spatial scale affects bioclimate model projections of climate change impacts on mountain plants

Plant species have responded to recent increases in global temperatures by shifting their geographical ranges poleward and to higher altitudes. Bioclimate models project future range contractions of montane species as suitable climate space shifts uphill. The species–climate relationships underlying such models are calibrated using data at either ‘macro’ scales (coarse resolution, e.g. 50 km × 50 km, and large spatial extent) or ‘local’ scales (fine resolution, e.g. 50 m × 50 m, and small spatial extent), but the two approaches have not been compared. This study projected macro (European) and local models for vascular plants at a mountain range in Scotland, UK, under low (+1.7 °C) and high (+3.3 °C) climate change scenarios for the 2080s. Depending on scenario, the local models projected that seven or eight out of 10 focal montane species would lose all suitable climate space at the site. However, the European models projected such a loss for only one species. The cause of this divergence was investigated by cross‐scale comparisons of estimated temperatures at montane species' warm range edges. The results indicate that European models overestimated species' thermal tolerances because the input coarse resolution climate data were biased against the cold, high‐altitude habitats of montane plants. Although tests at other mountain ranges are required, these results indicate that recent large‐scale modelling studies may have overestimated montane species' ability to cope with increasing temperatures, thereby underestimating the potential impacts of climate change. Furthermore, the results suggest that montane species persistence in microclimatic refugia might not be as widespread as previously speculated.     

Spatial scale,topography and thermoregulatory behaviour interact when modelling species’ thermal niches

The spatial scale at which climate and species’ occupancy data are gathered, and the resolution at which ecological models are run, can strongly influence predictions of species performance and distributions. Running model simulations at coarse rather than fine spatial resolutions, for example, can determine if a model accurately predicts the distribution of a species. The impacts of spatial scale on a model's accuracy are particularly pronounced across mountainous terrain. Understanding how these discrepancies arise requires a modelling approach in which the underlying processes that determine a species’ distribution are explicitly described. Here we use a process‐based model to explore how spatial resolution, topography and behaviour alter predictions of a species thermal niche, which in turn constrains its survival and geographic distribution. The model incorporates biophysical equations to predict the operative temperature (T_e), thermal‐dependent performance and survival of a typical insect, with a complex life‐cycle, in its microclimate. We run this model with geographic data from a mountainous terrain in South Africa using climate data at three spatial resolutions. We also explore how behavioural thermoregulation affects predictions of a species performance and survival by allowing the animal to select the optimum thermal location within each coarse‐grid cell. At the regional level, coarse‐resolution models predicted lower T_e at low elevations and higher T_e at high elevations than models run at fine‐resolutions. These differences were more prominent on steep, north‐facing slopes. The discrepancies in T_e in turn affected estimates of the species thermal niche. The modelling framework revealed how spatial resolution and topography influence predictions of species distribution models, including the potential impacts of climate change. These systematic biases must be accounted for when interpreting the outputs of future modelling studies, particularly when species distributions are predicted to shift from uniform to topographically heterogeneous landscapes.     

Using a historic drought and high‐heat event to validate thermal exposure predictions for ground‐dwelling birds

Deviations from typical environmental conditions can provide insight into how organisms may respond to future weather extremes predicted by climate modeling. During an episodic and multimonth heat wave event (i.e., ambient temperature up to 43.4°C), we studied the thermal ecology of a ground‐dwelling bird species in Western Oklahoma, USA. Specifically, we measured black bulb temperature (T_bb) and vegetation parameters at northern bobwhite (Colinus virginianus; hereafter bobwhite) adult and brood locations as well as at stratified random points in the study area. On the hottest days (i.e., ≥39°C), adults and broods obtained thermal refuge using tall woody cover that remained on average up to 16.51°C cooler than random sites on the landscape which reached >57°C. We also found that refuge sites used by bobwhites moderated thermal conditions by more than twofold compared to stratified random sites on the landscape but that T_bb commonly exceeded thermal stress thresholds for bobwhites (39°C) for several hours of the day within thermal refuges. The serendipitous high heat conditions captured in our study represent extreme heat for our study region as well as thermal stress for our study species, and subsequently allowed us to assess ground‐dwelling bird responses to temperatures that are predicted to become more common in the future. Our findings confirm the critical importance of tall woody cover for moderating temperatures and functioning as important islands of thermal refuge for ground‐dwelling birds, especially during extreme heat. However, the potential for extreme heat loads within thermal refuges that we observed (albeit much less extreme than the landscape) indicates that the functionality of tall woody cover to mitigate heat extremes may be increasingly limited in the future, thereby reinforcing predictions that climate change represents a clear and present danger for these species.     

Does climate variability influence the demography of wild primates? Evidence from long‐term life‐history data in seven species

Earth's rapidly changing climate creates a growing need to understand how demographic processes in natural populations are affected by climate variability, particularly among organisms threatened by extinction. Long‐term, large‐scale, and cross‐taxon studies of vital rate variation in relation to climate variability can be particularly valuable because they can reveal environmental drivers that affect multiple species over extensive regions. Few such data exist for animals with slow life histories, particularly in the tropics, where climate variation over large‐scale space is asynchronous. As our closest relatives, nonhuman primates are especially valuable as a resource to understand the roles of climate variability and climate change in human evolutionary history. Here, we provide the first comprehensive investigation of vital rate variation in relation to climate variability among wild primates. We ask whether primates are sensitive to global changes that are universal (e.g., higher temperature, large‐scale climate oscillations) or whether they are more sensitive to global change effects that are local (e.g., more rain in some places), which would complicate predictions of how primates in general will respond to climate change. To address these questions, we use a database of long‐term life‐history data for natural populations of seven primate species that have been studied for 29–52 years to investigate associations between vital rate variation, local climate variability, and global climate oscillations. Associations between vital rates and climate variability varied among species and depended on the time windows considered, highlighting the importance of temporal scale in detection of such effects. We found strong climate signals in the fertility rates of three species. However, survival, which has a greater impact on population growth, was little affected by climate variability. Thus, we found evidence for demographic buffering of life histories, but also evidence of mechanisms by which climate change could affect the fates of wild primates.     

Tropical forests are thermally buffered despite intensive selective logging

Tropical rainforests are subject to extensive degradation by commercial selective logging. Despite pervasive changes to forest structure, selectively logged forests represent vital refugia for global biodiversity. The ability of these forests to buffer temperature‐sensitive species from climate warming will be an important determinant of their future conservation value, although this topic remains largely unexplored. Thermal buffering potential is broadly determined by: (i) the difference between the “macroclimate” (climate at a local scale, m to ha) and the “microclimate” (climate at a fine‐scale, mm to m, that is distinct from the macroclimate); (ii) thermal stability of microclimates (e.g. variation in daily temperatures); and (iii) the availability of microclimates to organisms. We compared these metrics in undisturbed primary forest and intensively logged forest on Borneo, using thermal images to capture cool microclimates on the surface of the forest floor, and information from dataloggers placed inside deadwood, tree holes and leaf litter. Although major differences in forest structure remained 9–12 years after repeated selective logging, we found that logging activity had very little effect on thermal buffering, in terms of macroclimate and microclimate temperatures, and the overall availability of microclimates. For 1°C warming in the macroclimate, temperature inside deadwood, tree holes and leaf litter warmed slightly more in primary forest than in logged forest, but the effect amounted to <0.1°C difference between forest types. We therefore conclude that selectively logged forests are similar to primary forests in their potential for thermal buffering, and subsequent ability to retain temperature‐sensitive species under climate change. Selectively logged forests can play a crucial role in the long‐term maintenance of global biodiversity.     

Heat resistance throughout ontogeny: body size constrains thermal tolerance

Heat tolerance is a trait of paramount ecological importance and may determine a species' ability to cope with ongoing climate change. Although critical thermal limits have consequently received substantial attention in recent years, their potential variation throughout ontogeny remained largely neglected. We investigate whether such neglect may bias conclusions regarding a species' sensitivity to climate change. Using a tropical butterfly, we found that developmental stages clearly differed in heat tolerance. It was highest in pupae followed by larvae, adults and finally eggs and hatchlings. Strikingly, most of the variation found in thermal tolerance was explained by differences in body mass, which may thus impose a severe constraint on adaptive variation in stress tolerance. Furthermore, temperature acclimation was beneficial by increasing heat knock‐down time and therefore immediate survival under heat stress, but it affected reproduction negatively. Extreme temperatures strongly reduced survival and subsequent reproductive success even in our highly plastic model organism, exemplifying the potentially dramatic impact of extreme weather events on biodiversity. We argue that predictions regarding a species' fate under changing environmental conditions should consider variation in thermal tolerance throughout ontogeny, variation in body mass and acclimation responses as important predictors of stress tolerance.