Inclusion of local environmental conditions alters high-latitude vegetation change predictions based on bioclimatic models

Current predictions of how species will respond to climate change are typically based on coarse-grained climate surfaces utilizing bioclimate envelope modelling. However, the suitability of environmental conditions for a given species might result from a variety of factors including some unrelated to climate. To address this issue, we investigated whether the inclusion of topographical and soil information in bioclimatic envelope models would significantly alter predictions of climate change—induced fine-scale tree and shrub species range size changes at the tree-limit in subarctic Europe. Using generalized additive models and data on current climate and species distributions and three different climate scenarios for the period 2040–2069, we developed predictions of the currently suitable area and potential range size changes of seven tree and shrub species in an area of 1,100 km² at a resolution of 1-ha. The inclusion of topography and soil information increased the predictive accuracy of climate-only models for all studied species. The predicted changes in species distribution volumes were contradictory, and the predicted occurrences varied greatly depending on the model used. Our results therefore support the arguments that vegetation responses to climate change can be influenced by local environmental conditions and that attention should be paid to the combined effects of these factors. We conclude that disregarding local topography and soil conditions in bioclimatic models may result in biased projections of range expansions and the associated colonization, extinction and turnover assessments.     

Assessing the vulnerability of European butterflies to climate change using multiple criteria

Vulnerability of 100 European butterfly species to climate change was assessed using 13 different criteria and data on species distributions, climate, land cover and topography from 1,608 grid squares 30′ × 60′ in size, and species characteristics increasing the susceptibility to climate change. Four bioclimatic model-based criteria were developed for each species by comparing the present-day distribution and climatic suitability of the occupied grid cells with projected distribution and suitability in the future using the HadCM3-A2 climate scenario for 2051–2080. The proportions of disadvantageous land cover types (bare areas, water, snow and ice, artificial surfaces) and cultivated and managed land in the occupied grid squares and their surroundings were measured to indicate the amount of unfavourable land cover and dispersal barriers for butterflies, and topographical heterogeneity to indicate the availability of potential climatic refugia. Vulnerability was also assessed based on species dispersal ability, geographical localization and habitat specialization. Northern European species appeared to be amongst the most vulnerable European butterflies. However, there is much species-to-species variation, and species appear to be threatened due to different combinations of critical characteristics. Inclusion of additional criteria, such as life-history species characteristics, topography and land cover to complement the bioclimatic model-based species vulnerability measures can significantly deepen the assessments of species susceptibility to climate change.     

Inclusion of soil data improves the performance of bioclimatic envelope models for insect species distributions in temperate Europe

Aim To analyse the effect of the inclusion of soil and land‐cover data on the performance of bioclimatic envelope models for the regional‐scale prediction of butterfly (Rhopalocera) and grasshopper (Orthoptera) distributions. Location Temperate Europe (Belgium). Methods Distributional data were extracted from butterfly and grasshopper atlases at a resolution of 5 km for the period 1991–2006 in Belgium. For each group separately, the well‐surveyed squares (n = 366 for butterflies and n = 322 for grasshoppers) were identified using an environmental stratification design and were randomly divided into calibration (70%) and evaluation (30%) datasets. Generalized additive models were applied to the calibration dataset to estimate occurrence probabilities for 63 butterfly and 33 grasshopper species, as a function of: (1) climate, (2) climate and land‐cover, (3) climate and soil, and (4) climate, land‐cover and soil variables. Models were evaluated as: (1) the amount of explained deviance in the calibration dataset, (2) Akaike’s information criterion, and (3) the number of omission and commission errors in the evaluation dataset. Results Information on broad land‐cover classes or predominant soil types led to similar improvements in the performance relative to the climate‐only models for both taxonomic groups. In addition, the joint inclusion of land‐cover and soil variables in the models provided predictions that fitted more closely to the species distributions than the predictions obtained from bioclimatic models incorporating only land‐cover or only soil variables. The combined models exhibited higher discrimination ability between the presence and absence of species in the evaluation dataset. Main conclusions These results draw attention to the importance of soil data for species distribution models at regional scales of analysis. The combined inclusion of land‐cover and soil data in the models makes it possible to identify areas with suitable climatic conditions but unsuitable combinations of vegetation and soil types. While contingent on the species, the results indicate the need to consider soil information in regional‐scale species–climate impact models, particularly when predicting future range shifts of species under climate change.     

Predicting range expansion of the map butterfly in Northern Europe using bioclimatic models

The two main goals of this study are: (i) to examine the range shifts of a currently northwards expanding species, the map butterfly (Araschnia levana), in relation to annual variation in weather, and (ii) to test the capability of a bioclimatic envelope model, based on broad-scale European distribution data, to predict recent distributional changes (2000–2004) of the species in Finland. A significant relationship between annual maximum dispersal distance of the species and late summer temperature was detected. This suggests that the map butterfly has dispersed more actively in warmer rather than cooler summers, the most notable dispersal events being promoted by periods of exceptionally warm weather and southerly winds. The accuracy of the broad-scale bioclimatic model built for the species with European data using Generalized Additive Models (GAM) was good based on split-sample evaluation for a single period. However, the model’s performance was poor when applied to predict range shifts in Finland. Among the many potential explanations for the poor success of the transferred bioclimatic model, is the fact that bioclimatic envelope models do not generally account for species dispersal. This and other uncertainties support the view that bioclimatic models should be applied with caution when they are used to project future range shifts of species.     

Determinants of the biogeographical distribution of butterflies in boreal regions

Aim  We explored the relative contributions of climatic and land-cover factors in explaining the distribution patterns of butterflies in a boreal region.
Location  Finland, northern Europe.
Methods  Data from a national butterfly atlas survey carried out during 1991–2003, with a 10-km grain grid system, were used in these analyses. We used generalized additive models (GAM) and hierarchical partitioning (HP) to explore the main environmental correlates (climate and land-cover) of the realized niches of 98 butterfly species. The accuracy of the distribution models (GAMs) was validated by resubstitution and cross-validation approaches, using the area under the curve (AUC) derived from the receiver operating characteristic (ROC) plots.
Results  Predictive accuracies of the 98 individual environment–butterfly models varied from low to very high (cross-validated AUC values 0.48–0.99), with a mean of 0.79. The results of both the GAM and HP analyses were broadly concordant. Most of the variation in butterfly distributions is associated with growing degree-days, mean temperature of the coldest month and cover of built-up area in all six phylogenetic groups (butterfly families). There were no statistically significant differences in predictive accuracy among the different butterfly families.
Main conclusions  About three-quarters of the distributions of butterfly species in Finland appear to be governed principally by climatic, predominantly temperature-related, factors. This indicates that many butterfly species may respond rapidly to the projected climate change in boreal regions. By determining the ecological niches of multiple species, we can project their range shifts in response to changes in climate and land-cover, and identify species that are particularly sensitive to forecasted global changes.     

Equilibrium of global amphibian species distributions with climate

A common assumption in bioclimatic envelope modeling is that species distributions are in equilibrium with contemporary climate. A number of studies have measured departures from equilibrium in species distributions in particular regions, but such investigations were never carried out for a complete lineage across its entire distribution. We measure departures of equilibrium with contemporary climate for the distributions of the world amphibian species. Specifically, we fitted bioclimatic envelopes for 5544 species using three presence-only models. We then measured the proportion of the modeled envelope that is currently occupied by the species, as a metric of equilibrium of species distributions with climate. The assumption was that the greater the difference between modeled bioclimatic envelope and the occupied distribution, the greater the likelihood that species distribution would not be at equilibrium with contemporary climate. On average, amphibians occupied 30% to 57% of their potential distributions. Although patterns differed across regions, there were no significant differences among lineages. Species in the Neotropic, Afrotropics, Indo-Malay, and Palaearctic occupied a smaller proportion of their potential distributions than species in the Nearctic, Madagascar, and Australasia. We acknowledge that our models underestimate non equilibrium, and discuss potential reasons for the observed patterns. From a modeling perspective our results support the view that at global scale bioclimatic envelope models might perform similarly across lineages but differently across regions.     

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.     

Climate change, plant migration, and range collapse in a global biodiversity hotspot: the Banksia (Proteaceae) of Western Australia

Climate change has already altered global patterns of biodiversity by modifying the geographic distributions of species. Forecasts based on bioclimatic envelop modeling of distributions of species suggests greater impacts can be expected in the future, but such projections are contingent on assumptions regarding future climate and migration rates of species. Here, we present a first assessment of the potential impact of climate change on a global biodiversity hotspot in southwestern Western Australia. Across three representative scenarios of future climate change, we simulated migration of 100 Banksia (Proteaceae) species at a rate of 5 km decade^?1 and compared projected impacts with those under the commonly applied, but acknowledged as inadequate, assumptions of ‘full‐’ and ‘no‐migration.’ Across all climate × migration scenarios, 66% of species were projected to decline, whereas only 6% were projected to expand or remain stable. Between 5% and 25% of species were projected to suffer range losses of 100% by 2080, depending mainly on climate scenario. Species losses were driven primarily by changes in current precipitation regimes, with the greatest losses of species projected to occur in a transition zone between wet coastal areas and interior arid regions and which is projected to become more arid in the future. Because the ranges of most species tended to collapse in all climate scenarios, we found that climate change impacts to flora of southwestern Western Australia may be large, even under optimistic assumptions regarding migration abilities. Taken together, our results suggest that the future of biodiversity in southwestern Western Australia may lie largely in the degree to which this hotspot experiences increased drought and in the ability of species to tolerate such decreases in precipitation. More broadly, our study is among a growing number of theoretical studies suggesting the impacts of future climate change on global biodiversity may be considerable.     

Climate-driven,but dynamic and complex? A reconciliation of competing hypotheses for species’ distributions

Estimates of the percentage of species “committed to extinction” by climate change range from 15% to 37%. The question is whether factors other than climate need to be included in models predicting species’ range change. We created demographic range models that include climate vs. climate-plus-competition, evaluating their influence on the geographic distribution of Pinus edulis, a pine endemic to the semiarid southwestern U.S. Analyses of data on 23,426 trees in 1941 forest inventory plots support the inclusion of competition in range models. However, climate and competition together only partially explain this species’ distribution. Instead, the evidence suggests that climate affects other range-limiting processes, including landscape-scale, spatial processes such as disturbances and antagonistic biotic interactions. Complex effects of climate on species distributions—through indirect effects, interactions, and feedbacks—are likely to cause sudden changes in abundance and distribution that are not predictable from a climate-only perspective.     

Climatic niche of the Saker Falcon Falco cherrug: predicted new areas to direct population surveys in Central Asia

Accurate species distribution data across remote and extensive geographical areas are difficult to obtain. Here, we use bioclimatic envelope models to determine climatic constraints on the distribution of the migratory Saker Falcon Falco cherrug to identify areas in data-deficient regions that may contain unidentified populations. Sakers live at low densities across large ranges in remote regions, making distribution status difficult to assess. Using presence-background data and eight bioclimatic variables within a species distribution modelling framework, we applied MaxEnt to construct models for both breeding and wintering ranges. Occurrence data were spatially filtered and climatic variables tested for multicollinearity before selecting best fit models using the Akaike information criterion by tuning MaxEnt parameters. Model predictive performance tested using the continuous Boyce index (B) was high for both breeding (B_TEST = 0.921) and wintering models (B_TEST = 0.735), with low omission rates and minimal overfitting. The Saker climatic niche was defined by precipitation in the warmest quarter in the breeding range model, and mean temperature in the wettest quarter in the wintering range model. Our models accurately predicted areas of highest climate suitability and defined the climatic constraints on a wide-ranging rare species, suggesting that climate is a key determinant of Saker distribution across macro-scales. We recommend targeted population surveys for the Saker based on model predictions to areas of highest climatic suitability in key regions with distribution knowledge gaps, in particular the Qinghai-Tibet plateau in western China. Further applications of our models could identify protected areas and reintroduction sites, inform development conflicts, and assess the impact of climate change on distributions.     

The importance of biotic interactions for modelling species distributions under climate change

Aim There is a debate as to whether biotic interactions exert a dominant role in governing species distributions at macroecological scales. The prevailing idea is that climate is the key limiting factor; thus models that use present‐day climate–species range relationships are expected to provide reasonable means to quantify the impacts of climate change on species distributions. However, there is little empirical evidence that biotic interactions would not constrain species distributions at macroecological scales. We examine this idea, for the first time, and provide tests for two null hypotheses: (H₀ 1) – biotic interactions do not exert a significant role in explaining current distributions of a particular species of butterfly (clouded Apollo, Parnassius mnemosyne) in Europe; and (H₀ 2) – biotic interactions do not exert a significant role in predictions of altered species’ ranges under climate change. Location Europe. Methods Generalized additive modelling (GAM) was used to investigate relationships between species and climate; species and host plants; and species and climate + host plants. Because models are sensitive to the variable selection strategies utilised, four alternative approaches were used: AIC (Akaike's Information Criterion), BIC (Bayesian Information Criterion), BRUTO (Adaptive Backfitting), and CROSS (Cross Selection). Results In spite of the variation in the variables selected with different methods, both hypotheses (H₀ 1 and H₀ 2) were falsified, providing support for the proposition that biotic interactions significantly affect both the explanatory and predictive power of bioclimatic envelope models at macro scales. Main conclusions Our results contradict the widely held view that the effects of biotic interactions on individual species distributions are not discernible at macroecological scales. Results are contingent on the species, type of interaction and methods considered, but they call for more stringent evidence in support of the idea that purely climate‐based modelling would be sufficient to quantify the impacts of climate change on species distributions.     

The role of land cover in bioclimatic models depends on spatial resolution

Aim We explored the importance of climate and land cover in bird species distribution models on multiple spatial scales. In particular, we tested whether the integration of land cover data improves the performance of pure bioclimatic models. Location Finland, northern Europe. Methods The data of the bird atlas survey carried out in 1986–89 using a 10 × 10 km uniform grid system in Finland were employed in the analyses. Land cover and climatic variables were compiled using the same grid system. The dependent and explanatory variables were resampled to 20‐km, 40‐km and 80‐km resolutions. Generalized additive models (GAM) were constructed for each of the 88 land bird species studied in order to estimate the probability of occurrence as a function of (1) climate and (2) climate and land cover variables. Model accuracy was measured by a cross‐validation approach using the area under the curve (AUC) of a receiver operating characteristic (ROC) plot. Results In general, the accuracies of the 88 bird–climate models were good at all studied resolutions. However, the inclusion of land cover increased the performance of 79 and 78 of the 88 bioclimatic models at 10‐km and 20‐km resolutions, respectively. There was no significant improvement at the 40‐km resolution. In contrast to the finer resolutions, the inclusion of land cover variables decreased the modelling accuracy at 80km resolution. Main conclusions Our results suggest that the determinants of bird species distributions are hierarchically structured: climatic variables are large‐scale determinants, followed by land cover at finer resolutions. The majority of the land bird species in Finland are rather clearly correlated with climate, and bioclimate envelope models can provide useful tools for identifying the relationships between these species and the environment at resolutions ranging from 10 km to 80 km. However, the notable contribution of land cover to the accuracy of bioclimatic models at 10–20‐km resolutions indicates that the integration of climate and land cover information can improve our understanding and model predictions of biogeographical patterns under global change.     

Exploring consensus in 21st century projections of climatically suitable areas for African vertebrates

Africa is predicted to be highly vulnerable to 21st century climatic changes. Assessing the impacts of these changes on Africa's biodiversity is, however, plagued by uncertainties, and markedly different results can be obtained from alternative bioclimatic envelope models or future climate projections. Using an ensemble forecasting framework, we examine projections of future shifts in climatic suitability, and their methodological uncertainties, for over 2500 species of mammals, birds, amphibians and snakes in sub‐Saharan Africa. To summarize a priori the variability in the ensemble of 17 general circulation models, we introduce a consensus methodology that combines co‐varying models. Thus, we quantify and map the relative contribution to uncertainty of seven bioclimatic envelope models, three multi‐model climate projections and three emissions scenarios, and explore the resulting variability in species turnover estimates. We show that bioclimatic envelope models contribute most to variability, particularly in projected novel climatic conditions over Sahelian and southern Saharan Africa. To summarize agreements among projections from the bioclimatic envelope models we compare five consensus methodologies, which generally increase or retain projection accuracy and provide consistent estimates of species turnover. Variability from emissions scenarios increases towards late‐century and affects southern regions of high species turnover centred in arid Namibia. Twofold differences in median species turnover across the study area emerge among alternative climate projections and emissions scenarios. Our ensemble of projections underscores the potential bias when using a single algorithm or climate projection for Africa, and provides a cautious first approximation of the potential exposure of sub‐Saharan African vertebrates to climatic changes. The future use and further development of bioclimatic envelope modelling will hinge on the interpretation of results in the light of methodological as well as biological uncertainties. Here, we provide a framework to address methodological uncertainties and contextualize results.     

Predicting the impact of climate change on Australia's most endangered snake, Hoplocephalus bungaroides

Aim  To predict how the bioclimatic envelope of the broad-headed snake (BHS) ( Hoplocephalus bungaroides ) may be redistributed under future climate warming scenarios.
Location  South-eastern New South Wales, Australia.
Methods  We used 159 independent locations for the species and 35 climatic variables to model the bioclimatic envelope for the BHS using two modelling approaches – B ioclim and M axent . Predictions were made under current climatic conditions and we also predicted the species distribution under low and high climate change scenarios for 2030 and 2070.
Results  Broad-headed snakes currently encompass their entire bioclimatic envelope. Both modelling approaches predict that suitable climate space for BHS will be lost to varying degrees under both climate warming scenarios, and under the worst case, only 14% of known snake populations may persist.
Main conclusions  Areas of higher elevation within the current range will be most important for persistence of this species because they will remain relatively moist and cool even under climate change and will match the current climate envelope. Conservation efforts should focus on areas where suitable climate space may persist under climate warming scenarios. Long-term monitoring programs should be established both in these areas and where populations are predicted to become extirpated, so that we can accurately determine changes in the distribution of this species throughout its range.     

Do we need land‐cover data to model species distributions in Europe?

Aim To assess the influence of land cover and climate on species distributions across Europe. To quantify the importance of land cover to describe and predict species distributions after using climate as the main driver. Location The study area is Europe. Methods (1) A multivariate analysis was applied to describe land‐cover distribution across Europe and assess if the land cover is determined by climate at large spatial scales. (2) To evaluate the importance of land cover to predict species distributions, we implemented a spatially explicit iterative procedure to predict species distributions of plants (2603 species), mammals (186 species), breeding birds (440 species), amphibian and reptiles (143 species). First, we ran bioclimatic models using stepwise generalized additive models using bioclimatic variables. Secondly, we carried out a regression of land cover (LC) variables against residuals from the bioclimatic models to select the most relevant LC variables. Finally, we produced mixed models including climatic variables and those LC variables selected as decreasing the residual of bioclimatic models. Then we compared the explanatory and predictive power of the pure bioclimatic against the mixed model. Results (1) At the European coarse resolution, land cover is mainly driven by climate. Two bioclimatic axes representing a gradient of temperature and a gradient of precipitation explained most variation of land‐cover distribution. (2) The inclusion of land cover improved significantly the explanatory power of bioclimatic models and the most relevant variables across groups were those not explained or poorly explained by climate. However, the predictive power of bioclimatic model was not improved by the inclusion of LC variables in the iterative model selection process. Main conclusion Climate is the major driver of both species and land‐cover distributions over Europe. Yet, LC variables that are not explained or weakly associated with climate (inland water, sea or arable land) are interesting to describe particular mammal, bird and tree distributions. However, the addition of LC variables to pure bioclimatic models does not improve their predictive accuracy.     

Potential changes in the distributions of latitudinally restricted Australian butterfly species in response to climate change

This study assessed potential changes in the distributions of Australian butterfly species in response to global warming. The bioclimatic program, BIOCLIM, was used to determine the current climatic ranges of 77 butterfly species restricted to Australia. We found that the majority of these species had fairly wide climatic ranges in comparison to other taxa, with only 8% of butterfly species having a mean annual temperature range spanning less than 3 °C. The potential changes in the distributions of 24 butterfly species under four climate change scenarios for 2050 were also modelled using BIOCLIM. Results suggested that even species with currently wide climatic ranges may still be vulnerable to climate change; under a very conservative climate change scenario (with a temperature increase of 0.8–1.4 °C by 2050) 88% of species distributions decreased, and 54% of species distributions decreased by at least 20%. Under an extreme scenario (temperature increase of 2.1–3.9 °C by 2050) 92% of species distributions decreased, and 83% of species distributions decreased by at least 50%. Furthermore, the proportion of the current range that was contained within the predicted range decreased from an average of 63% under a very conservative scenario to less than 22% under the most extreme scenario. By assessing the climatic ranges that species are currently exposed to, the extent of potential changes in distributions in response to climate change and details of their life histories, we identified species whose characteristics may make them particularly vulnerable to climate change in the future.     

Patterns and Variability of Projected Bioclimatic Habitat for Pinus albicaulis in the Greater Yellowstone Area

Projected climate change at a regional level is expected to shift vegetation habitat distributions over the next century. For the sub-alpine species whitebark pine (Pinus albicaulis), warming temperatures may indirectly result in loss of suitable bioclimatic habitat, reducing its distribution within its historic range. This research focuses on understanding the patterns of spatiotemporal variability for future projected P.albicaulis suitable habitat in the Greater Yellowstone Area (GYA) through a bioclimatic envelope approach. Since intermodel variability from General Circulation Models (GCMs) lead to differing predictions regarding the magnitude and direction of modeled suitable habitat area, nine bias-corrected statistically down-scaled GCMs were utilized to understand the uncertainty associated with modeled projections. P.albicaulis was modeled using a Random Forests algorithm for the 1980–2010 climate period and showed strong presence/absence separations by summer maximum temperatures and springtime snowpack. Patterns of projected habitat change by the end of the century suggested a constant decrease in suitable climate area from the 2010 baseline for both Representative Concentration Pathways (RCPs) 8.5 and 4.5 climate forcing scenarios. Percent suitable climate area estimates ranged from 2–29% and 0.04–10% by 2099 for RCP 8.5 and 4.5 respectively. Habitat projections between GCMs displayed a decrease of variability over the 2010–2099 time period related to consistent warming above the 1910–2010 temperature normal after 2070 for all GCMs. A decreasing pattern of projected P.albicaulis suitable habitat area change was consistent across GCMs, despite strong differences in magnitude. Future ecological research in species distribution modeling should consider a full suite of GCM projections in the analysis to reduce extreme range contractions/expansions predictions. The results suggest that restoration strageties such as planting of seedlings and controlling competing vegetation may be necessary to maintain P.albicaulis in the GYA under the more extreme future climate scenarios.     

Inclusion of trophic interactions increases the vulnerability of an alpine butterfly species to climate change

Climate change is expected to have significant and complex impacts on ecological communities. In addition to direct effects of climate on species, there can also be indirect effects through an intermediary species, such as in host–plant interactions. Indirect effects are expected to be more pronounced in alpine environments because these ecosystems are sensitive to temperature changes and there are limited areas for migration of both species (i.e. closed systems), and because of simpler trophic interactions. We tested the hypothesis that climate change will reduce the range of an alpine butterfly (Parnassius smintheus) because of indirect effects through its host plant (Sedum sp.). To test for direct and indirect effects, we used the simulations of climate change to assess the distribution of P. smintheus with and without Sedum sp. We also compared the projected ranges of P. smintheus to four other butterfly species that are found in the alpine, but that are generalists feeding on many plant genera. We found that P. smintheus gained distributional area in climate‐only models, but these gains were significantly reduced with the inclusion of Sedum sp. and in dry‐climate scenarios which resulted in a reduction in net area. When compared to the more generalist butterfly species, P. smintheus exhibited the largest loss in suitable habitat. Our findings support the importance of including indirect effects in modelling species distributions in response to climate change. We highlight the potentially large and still neglected impacts climate change can have on the trophic structure of communities, which can lead to significant losses of biodiversity. In the future, communities will continue to favour species that are generalists as climate change induces asynchronies in the migration of species.     

Species traits are associated with the quality of bioclimatic models

Aim To analyse the effects of nine species trait variables on the accuracy of bioclimatic envelope models built for 98 butterfly species. Location Finland, northern Europe. Methods Data from a national butterfly atlas monitoring scheme (NAFI) collected from 1991–2003 with a resolution of 10 × 10 km were used in the analyses. Generalized additive models (GAMs) were constructed for 98 butterfly species to predict their occurrence as a function of climatic variables. Modelling accuracy was measured as the cross‐validation area under the curve (AUC) of the receiver–operating characteristic plot. Observed variation in modelling accuracy was related to species traits using multiple GAMs. The effects of phylogenetic relatedness among butterflies were accounted for by using generalized estimation equations. Results The values of the cross‐validation AUC for the 98 species varied between 0.56 and 1.00 with a mean of 0.79. Five species trait variables were included in the GAM that explained 71.4% of the observed variation in modelling accuracy. Four variables remained significant after accounting for phylogenetic relatedness. Species with high mobility and a long flight period were modelled less accurately than species with low mobility and a short flight period. Large species (>50 mm in wing span) were modelled more accurately than small ones. Species inhabiting mires had especially poor models, whereas the models for species inhabiting rocky outcrops, field verges and open fells were more accurate compared with other habitats. Main conclusions These results draw attention to the importance of species traits variables for species–climate impact models. Most importantly, species traits may have a strong impact on the performance of bioclimatic envelope models, and certain trait groups can be inherently difficult to model reliably. These uncertainties should be taken into account by downweighting or excluding species with such traits in studies applying bioclimatic modelling and making assessments of the impacts of climate change.