Climatic predictors of temperature performance curve parameters in ectotherms imply complex responses to climate change

Determining organismal responses to climate change is one of biology's greatest challenges. Recent forecasts for future climates emphasize altered temperature variation and precipitation, but most studies of animals have largely focused on forecasting the outcome of changes in mean temperature. Theory suggests that extreme thermal variation and precipitation will influence species performance and hence affect their response to changes in climate. Using an information-theoretic approach, we show that in squamate ectotherms (mostly lizards and snakes), two fitness-influencing components of performance, the critical thermal maximum and the thermal optimum, are more closely related to temperature variation and to precipitation, respectively, than they are to mean thermal conditions. By contrast, critical thermal minimum is related to mean annual temperature. Our results suggest that temperature variation and precipitation regimes have had a strong influence on the evolution of ectotherm performance, so that forecasts for animal responses to climate change will have to incorporate these factors and not only changes in average temperature.     

Predicting Climate Change Impacts on the Amount and Duration of Autumn Colors in a New England Forest

Climate change affects the phenology of many species. As temperature and precipitation are thought to control autumn color change in temperate deciduous trees, it is possible that climate change might also affect the phenology of autumn colors. Using long-term data for eight tree species in a New England hardwood forest, we show that the timing and cumulative amount of autumn color are correlated with variation in temperature and precipitation at specific times of the year. A phenological model driven by accumulated cold degree-days and photoperiod reproduces most of the interspecific and interannual variability in the timing of autumn colors. We use this process-oriented model to predict changes in the phenology of autumn colors to 2099, showing that, while responses vary among species, climate change under standard IPCC projections will lead to an overall increase in the amount of autumn colors for most species.     

Similarities in butterfly emergence dates among populations suggest local adaptation to climate

Phenology shifts are the most widely cited examples of the biological impact of climate change, yet there are few assessments of potential effects on the fitness of individual organisms or the persistence of populations. Despite extensive evidence of climate‐driven advances in phenological events over recent decades, comparable patterns across species' geographic ranges have seldom been described. Even fewer studies have quantified concurrent spatial gradients and temporal trends between phenology and climate. Here we analyse a large data set (~129 000 phenology measures) over 37 years across the UK to provide the first phylogenetic comparative analysis of the relative roles of plasticity and local adaptation in generating spatial and temporal patterns in butterfly mean flight dates. Although populations of all species exhibit a plastic response to temperature, with adult emergence dates earlier in warmer years by an average of 6.4 days per °C, among‐population differences are significantly lower on average, at 4.3 days per °C. Emergence dates of most species are more synchronised over their geographic range than is predicted by their relationship between mean flight date and temperature over time, suggesting local adaptation. Biological traits of species only weakly explained the variation in differences between space‐temperature and time‐temperature phenological responses, suggesting that multiple mechanisms may operate to maintain local adaptation. As niche models assume constant relationships between occurrence and environmental conditions across a species' entire range, an important implication of the temperature‐mediated local adaptation detected here is that populations of insects are much more sensitive to future climate changes than current projections suggest.     

Links between plant species’ spatial and temporal responses to a warming climate

To generate realistic projections of species’ responses to climate change, we need to understand the factors that limit their ability to respond. Although climatic niche conservatism, the maintenance of a species’s climatic niche over time, is a critical assumption in niche-based species distribution models, little is known about how universal it is and how it operates. In particular, few studies have tested the role of climatic niche conservatism via phenological changes in explaining the reported wide variance in the extent of range shifts among species. Using historical records of the phenology and spatial distribution of British plants under a warming climate, we revealed that: (i) perennial species, as well as those with weaker or lagged phenological responses to temperature, experienced a greater increase in temperature during flowering (i.e. failed to maintain climatic niche via phenological changes); (ii) species that failed to maintain climatic niche via phenological changes showed greater northward range shifts; and (iii) there was a complementary relationship between the levels of climatic niche conservatism via phenological changes and range shifts. These results indicate that even species with high climatic niche conservatism might not show range shifts as instead they track warming temperatures during flowering by advancing their phenology.     

Trait means and reaction norms: the consequences of climate change/invasion interactions at the organism level

How the impacts of climate change on biological invasions will play out at the mechanistic level is not well understood. Two major hypotheses have been proposed: invasive species have a suite of traits that enhance their performance relative to indigenous ones over a reasonably wide set of circumstances; invasive species have greater phenotypic plasticity than their indigenous counterparts and will be better able to retain performance under altered conditions. Thus, two possibly independent, but complementary mechanistic perspectives can be adopted: based on trait means and on reaction norms. Here, to demonstrate how this approach might be applied to understand interactions between climate change and invasion, we investigate variation in the egg development times and their sensitivity to temperature amongst indigenous and introduced springtail species in a cool temperate ecosystem (Marion Island, 46°54′S 37°54′E) that is undergoing significant climate change. Generalized linear model analyses of the linear part of the development rate curves revealed significantly higher mean trait values in the invasive species compared to indigenous species, but no significant interactions were found when comparing the thermal reaction norms. In addition, the invasive species had a higher hatching success than the indigenous species at high temperatures. This work demonstrates the value of explicitly examining variation in trait means and reaction norms among indigenous and invasive species to understand the mechanistic basis of variable responses to climate change among these groups.     

An indicator highlights seasonal variation in the response of Lepidoptera communities to warming

The impacts of climate change on species and ecosystems are increasingly evident. While these tend to be clearest with respect to changes in phenology and distribution ranges, there are also important consequences for population sizes and community structure. There is an urgent need to develop ecological indicators that can be used to detect climate-driven changes in ecological communities, and identify how those impacts may vary spatially. Here we describe the development of a new community-based seasonal climate change indicator that uses national population and weather indices. We test this indicator using Lepidopteran and co-located weather data collected across a range of UK Environmental Change Network (ECN) sites. We compare our butterfly indicator with estimates derived from an alternative, previously published metric, the Community Temperature Index (CTI).First, we quantified the effect of temperature on population growth rates of moths and butterflies (Species Temperature Response, STR) by modelling annual variation in national population indices as a function of nationally averaged seasonal variation in temperature, using species and weather data independent of the ECN data. Then, we calculated average STRs for annually summarised species data from each ECN site, weighted by species’ abundance, to produce the Community Temperature Response (CTR). Finally, we tested the extent to which CTR correlated with spatial variation in temperature between sites and the extent to which temporal variation in CTR tracked both annual and seasonal warming trends.Mean site CTR was positively correlated with mean site temperature for moths but not butterflies. However, spatial variation in moth communities was well explained by mean site summer temperature and butterfly communities by winter temperature, respectively accounting for 74% and 63% of variation. Temporal variation in moth and butterfly CTR within sites did not vary with the mean annual temperature but responded to variation in the mean temperature of specific seasons. There were positive correlations between moth seasonal CTRs and seasonal temperatures in winter, spring and summer; and butterfly seasonal CTRs and seasonal temperatures in winter and summer. Butterfly CTR and CTI both correlated spatially and temporally with winter temperature.Our results highlight the need for seasonality to be considered when examining the impact of climate change on communities. Seasonal CTRs may be used to track the impact of changing temperatures on biodiversity and help identify potential mechanisms by which climate change is affecting communities. In the case of Lepidoptera, our results suggest that future warming may reassemble Lepidoptera communities.     

The effect of past changes in inter‐annual temperature variability on tree distribution limits

Aim The northern limits of temperate broadleaved species in Fennoscanndia are controlled by their requirements for summer warmth for successful regeneration and growth as well as by the detrimental effects of winter cold on plant tissue. However, occurrences of meteorological conditions with detrimental effects on individual species are rare events rather than a reflection of average conditions. We explore the effect of changes in inter‐annual temperature variability on the abundances of the tree species Tilia cordata, Quercus robur and Ulmus glabra near their distribution limits using a process‐based model of ecosystem dynamics. Location A site in central Sweden and a site in southern Finland were used as examples for the ecotone between boreal and temperate forests in Fennoscandia. The Finnish site was selected because of the availability of varve‐thickness data. Methods The dynamic vegetation model LPJ‐GUESS was run with four scenarios of inter‐annual temperature forcing for the last 10,000 years. In one scenario the variability in the thickness of summer and winter varves from the annually laminated lake in Finland was used as a proxy for past inter‐annual temperature variability. Two scenarios were devised to explore systematically the effect of stepwise changes in the variance and shape parameter of a probability distribution. All variability scenarios were run both with and without the long‐term trend in Holocene temperature change predicted by an atmospheric general circulation model. Results Directional changes in inter‐annual temperature variability have significant effects on simulated tree distribution limits through time. Variations in inter‐annual temperature variability alone are shown to alter vegetation composition by magnitudes similar to the magnitude of changes driven by variation in mean temperatures. Main conclusions The varve data indicate that inter‐annual climate variability has changed in the past. The model results show that past changes in species abundance can be explained by changes in the inter‐annual variability of climate parameters as well as by mean climate. Because inter‐annual climatic variability is predicted to change in the future, this component of climate change should be taken into account both when making projections of future plant distributions and when interpreting vegetation history.     

The Definition of British Water Beetle Species Pools (Coleoptera) and their Relationship to Altitude, Temperature, Precipitation and Land Cover Variables

Pooled water beetle species lists from 1826 British national grid 10-km squares were analysed using multivariate ordination and classification methods. The relationships of pool groups to the climate, altitude and land cover variables were assessed using constrained and partial ordinations. Ordination of the species pool data indicated a major trend between squares in the north-west of Scotland and those in southern England, illustrating differences in acid and basic water standing water. Secondary variation was from acid standing water to fast-flowing streams and rivers. Classification generated nine species pool groups. These showed a distinct north-west to south-east trend but there was no obvious coastal or brackish water effect on distribution. The climatic and land cover variables were all significantly related to each other, and to north-south variation in grid square location, but the constrained ordination results indicated that that the most important influence on water beetle species pool distribution was mean summer temperature. Although the amount of variation explained by the environmental variables was low, spatial variation in the environmental predictors was almost as important as the environmental variables themselves in determining species pool composition. Mean annual temperature was also strongly correlated with species pool distribution with two land cover variables slightly less important. Altitude and precipitation had the least influence. The water beetle national recording scheme database appears to be of sufficiently high quality for environmental investigations at the British scale. There is considerable potential for the synthesis of invertebrate species distribution, land cover and climate change predictions in the assessment of environmental change. The results, together with previous work on other invertebrate species, indicate that changing summer temperatures may have a considerable influence on the distribution British invertebrate species.     

Designing ecological climate change impact assessments to reflect key climatic drivers

Identifying the climatic drivers of an ecological system is a key step in assessing its vulnerability to climate change. The climatic dimensions to which a species or system is most sensitive – such as means or extremes – can guide methodological decisions for projections of ecological impacts and vulnerabilities. However, scientific workflows for combining climate projections with ecological models have received little explicit attention. We review Global Climate Model (GCM) performance along different dimensions of change and compare frameworks for integrating GCM output into ecological models. In systems sensitive to climatological means, it is straightforward to base ecological impact assessments on mean projected changes from several GCMs. Ecological systems sensitive to climatic extremes may benefit from what we term the ‘model space’ approach: a comparison of ecological projections based on simulated climate from historical and future time periods. This approach leverages the experimental framework used in climate modeling, in which historical climate simulations serve as controls for future projections. Moreover, it can capture projected changes in the intensity and frequency of climatic extremes, rather than assuming that future means will determine future extremes. Given the recent emphasis on the ecological impacts of climatic extremes, the strategies we describe will be applicable across species and systems. We also highlight practical considerations for the selection of climate models and data products, emphasizing that the spatial resolution of the climate change signal is generally coarser than the grid cell size of downscaled climate model output. Our review illustrates how an understanding of how climate model outputs are derived and downscaled can improve the selection and application of climatic data used in ecological modeling.     

Growth and development of an invasive forest insect under current and future projected temperature regimes

Temperature and its impact on fitness are fundamental for understanding range shifts and population dynamics under climate change. Geographic climate heterogeneity, behavioral and physiological plasticity, and thermal adaptation to local climates make predicting the responses of species to climate change complex. Using larvae from seven geographically distinct wild populations in the eastern United States of the non‐native forest pest Lymantria dispar dispar (L.), we conducted a simulated reciprocal transplant experiment in environmental chambers using six custom temperature regimes representing contemporary conditions near the southern and northern extremes of the US invasion front and projections under two climate change scenarios for the year 2050. Larval growth and development rates increased with climate warming compared with current thermal regimes and tended to be greater for individuals originally sourced from southern rather than northern populations. Although increases in growth and development rates with warming varied somewhat by region of the source population, there was not strong evidence of local adaptation, southern populations tended to outperform those from northern populations in all thermal regimes. Our study demonstrates the utility of simulating thermal regimes under climate change in environmental chambers and emphasizes how the impacts from future increases in temperature can vary based on geographic differences in climate‐related performance among populations.     

A framework integrating physiology,dispersal and land‐use to project species ranges under climate change

To study the potential effects of climate change on species, one of the most popular approaches are species distribution models (SDMs). However, they usually fail to consider important species‐specific biological traits, such as species’ physiological capacities or dispersal ability. Furthermore, there is consensus that climate change does not influence species distributions in isolation, but together with other anthropogenic impacts such as land‐use change, even though studies investigating the relative impacts of different threats on species and their geographic ranges are still rare. Here we propose a novel integrative approach which produces refined future range projections by combining SDMs based on distribution, climate, and physiological tolerance data with empirical data on dispersal ability as well as current and future land‐use. Range projections based on different combinations of these factors show strong variation in projected range size for our study species Emberiza hortulana. Using climate and physiological data alone, strong range gains are projected. However, when we account for land‐use change and dispersal ability, future range‐gain may even turn into a future range loss. Our study highlights the importance of accounting for biological traits and processes in species distribution models and of considering the additive effects of climate and land‐use change to achieve more reliable range projections. Furthermore, with our approach we present a new tool to assess species’ vulnerability to climate change which can be easily applied to multiple species.     

Modelling changes in species’ abundance in response to projected climate change

Aim Existing climate envelope models give an indication of broad scale shifts in distribution, but do not specifically provide information on likely future population changes useful for conservation prioritization and planning. We demonstrate how these techniques can be developed to model likely future changes in absolute density and population size as a result of climate change. Location Great Britain. Methods Generalized linear models were used to model breeding densities of two northerly‐ and two southerly‐distributed bird species as a function of climate and land use. Models were built using count data from extensive national bird monitoring data and incorporated detectability to estimate absolute abundance. Projections of likely future changes in the distribution and abundance of these species were made by applying these models to projections of future climate change under two emissions scenarios. Results Models described current spatial variation in abundance for three of the four species and produced modelled current estimates of national populations that were similar to previously published estimates for all species. Climate change was projected to result in national population declines in the two northerly‐distributed species, with declines for Eurasian curlew Numenius arquata projected to be particularly severe. Conversely, the abundances of the two southerly distributed species were projected to increase nationally. Projected maps of future abundance may be used to identify priority areas for the future conservation of each species. Main conclusions The analytical methods provide a framework to make projections of impacts of climate change on species abundance, rather than simply projected range changes. Outputs may be summarized at any spatial scale, providing information to inform future conservation planning at national, regional and local scales. Results suggest that as a consequence of climate change, northerly distributed bird species in Great Britain are likely to become an increasingly high conservation priority within the UK.     

Projecting insect voltinism under high and low greenhouse gas emission conditions

We develop individual-based Monte Carlo methods to explore how climate change can alter insect voltinism under varying greenhouse gas emissions scenarios by using input distributions of diapause termination or spring emergence, development rate, and diapause initiation, linked to daily temperature and photoperiod. We show concurrence of these projections with a field dataset, and then explore changes in grape berry moth, Paralobesia viteana (Clemens), voltinism that may occur with climate projections developed from the average of three climate models using two different future emissions scenarios from the International Panel of Climate Change (IPCC). Based on historical climate data from 1960 to 2008, and projected downscaled climate data until 2099 under both high (A1fi) and low (B1) greenhouse gas emission scenarios, we used concepts of P. viteana biology to estimate distributions of individuals entering successive generations per year. Under the low emissions scenario, we observed an earlier emergence from diapause and a shift in mean voltinism from 2.8 to 3.1 generations per year, with a fraction of the population achieving a fourth generation. Under the high emissions scenario, up to 3.6 mean generations per year were projected by the end of this century, with a very small fraction of the population achieving a fifth generation. Changes in voltinism in this and other species in response to climate change likely will cause significant economic and ecological impacts, and the methods presented here can be readily adapted to other species for which the input distributions are reasonably approximated.     

Models of climate associations and distributions of amphibians in Italy

Analyzing the relationships between the distribution of animal species and climatic variables is not only important for understanding which factors govern species distribution but also for improving our ability to predict future ecological responses to climate change. In the context of global climate change, amphibians are of particular interest because of their extreme sensitivity to the variation of temperature and precipitation regimes. We analyzed species–climate relationships for 17 amphibian species occurring in Italy using species distribution data at the 10 × 10 km resolution. A machine learning method, Random Forests, was used to model the distribution of amphibians in relation to a set of 18 climatic variables. The results showed that the variables which had the highest importance were those related to precipitation, indicating that precipitation is an important factor in determining amphibian distribution. Future projections showed a complex response of species distributions, emphasizing the potential severity of climate change on the distributions of amphibians in Italy. The species that will decrease the most are those occurring in mountainous and Mediterranean areas. Our results provide some preliminary information that could be useful for amphibian conservation, indicating if future conservation priorities for some species should be enhanced.     

Shifting global invasive potential of European plants with climate change

Global climate change and invasions by nonnative species rank among the top concerns for agents of biological loss in coming decades. Although each of these themes has seen considerable attention in the modeling and forecasting communities, their joint effects remain little explored and poorly understood. We developed ecological niche models for 1804 species from the European flora, which we projected globally to identify areas of potential distribution, both at present and across 4 scenarios of future (2055) climates. As expected from previous studies, projections based on the CGCM1 climate model were more extreme than those based on the HadCM3 model, and projections based on the a2 emissions scenario were more extreme than those based on the b2 emissions scenario. However, less expected were the highly nonlinear and contrasting projected changes in distributional areas among continents: increases in distributional potential in Europe often corresponded with decreases on other continents, and species seeing expanding potential on one continent often saw contracting potential on others. In conclusion, global climate change will have complex effects on invasive potential of plant species. The shifts and changes identified in this study suggest strongly that biological communities will see dramatic reorganizations in coming decades owing to shifting invasive potential by nonnative species.     

An empirical test of the relative and combined effects of land‐cover and climate change on local colonization and extinction

Land‐cover and climate change are two main drivers of changes in species ranges. Yet, the majority of studies investigating the impacts of global change on biodiversity focus on one global change driver and usually use simulations to project biodiversity responses to future conditions. We conduct an empirical test of the relative and combined effects of land‐cover and climate change on species occurrence changes. Specifically, we examine whether observed local colonization and extinctions of North American birds between 1981–1985 and 2001–2005 are correlated with land‐cover and climate change and whether bird life history and ecological traits explain interspecific variation in observed occurrence changes. We fit logistic regression models to test the impact of physical land‐cover change, changes in net primary productivity, winter precipitation, mean summer temperature, and mean winter temperature on the probability of Ontario breeding bird local colonization and extinction. Models with climate change, land‐cover change, and the combination of these two drivers were the top ranked models of local colonization for 30%, 27%, and 29% of species, respectively. Conversely, models with climate change, land‐cover change, and the combination of these two drivers were the top ranked models of local extinction for 61%, 7%, and 9% of species, respectively. The quantitative impacts of land‐cover and climate change variables also vary among bird species. We then fit linear regression models to test whether the variation in regional colonization and extinction rate could be explained by mean body mass, migratory strategy, and habitat preference of birds. Overall, species traits were weakly correlated with heterogeneity in species occurrence changes. We provide empirical evidence showing that land‐cover change, climate change, and the combination of multiple global change drivers can differentially explain observed species local colonization and extinction.     

Bird population trends are linearly affected by climate change along species thermal ranges

Beyond the effects of temperature increase on local population trends and on species distribution shifts, how populations of a given species are affected by climate change along a species range is still unclear. We tested whether and how species responses to climate change are related to the populations locations within the species thermal range. We compared the average 20 year growth rates of 62 terrestrial breeding birds in three European countries along the latitudinal gradient of the species ranges. After controlling for factors already reported to affect bird population trends (habitat specialization, migration distance and body mass), we found that populations breeding close to the species thermal maximum have lower growth rates than those in other parts of the thermal range, while those breeding close to the species thermal minimum have higher growth rates. These results were maintained even after having controlled for the effect of latitude per se. Therefore, the results cannot solely be explained by latitudinal clines linked to the geographical structure in local spring warming. Indeed, we found that populations are not just responding to changes in temperature at the hottest and coolest parts of the species range, but that they show a linear graded response across their European thermal range. We thus provide insights into how populations respond to climate changes. We suggest that projections of future species distributions, and also management options and conservation assessments, cannot be based on the assumption of a uniform response to climate change across a species range or at range edges only.     

Range position and climate sensitivity: The structure of among‐population demographic responses to climatic variation

Species’ distributions will respond to climate change based on the relationship between local demographic processes and climate and how this relationship varies based on range position. A rarely tested demographic prediction is that populations at the extremes of a species’ climate envelope (e.g., populations in areas with the highest mean annual temperature) will be most sensitive to local shifts in climate (i.e., warming). We tested this prediction using a dynamic species distribution model linking demographic rates to variation in temperature and precipitation for wood frogs (Lithobates sylvaticus) in North America. Using long‐term monitoring data from 746 populations in 27 study areas, we determined how climatic variation affected population growth rates and how these relationships varied with respect to long‐term climate. Some models supported the predicted pattern, with negative effects of extreme summer temperatures in hotter areas and positive effects on recruitment for summer water availability in drier areas. We also found evidence of interacting temperature and precipitation influencing population size, such as extreme heat having less of a negative effect in wetter areas. Other results were contrary to predictions, such as positive effects of summer water availability in wetter parts of the range and positive responses to winter warming especially in milder areas. In general, we found wood frogs were more sensitive to changes in temperature or temperature interacting with precipitation than to changes in precipitation alone. Our results suggest that sensitivity to changes in climate cannot be predicted simply by knowing locations within the species’ climate envelope. Many climate processes did not affect population growth rates in the predicted direction based on range position. Processes such as species‐interactions, local adaptation, and interactions with the physical landscape likely affect the responses we observed. Our work highlights the need to measure demographic responses to changing climate.