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 abiotic and biotic factors limiting establishment of predatory fishes at their expanding northern range boundaries in Ontario,Canada

There is a poor understanding of the importance of biotic interactions in determining species distributions with climate change. Theory from invasion biology suggests that the success of species introductions outside of their historical ranges may be either positively (biotic acceptance) or negatively (biotic resistance) related to native biodiversity. Using data on fish community composition from two survey periods separated by approximately 28 years during which climate was warming, we examined the factors influencing the establishment of three predatory centrarchids: Smallmouth Bass (Micropterus dolomieu), Largemouth Bass (M. salmoides), and Rock Bass (Ambloplites rupestris) in lakes at their expanding northern range boundaries in Ontario. Variance partitioning demonstrated that, at a regional scale, abiotic factors play a stronger role in determining the establishment of these species than biotic factors. Pairing lakes within watersheds where each species had established with lakes sharing similar abiotic conditions where the species had not established revealed both positive and negative relationships between the establishment of centrarchids and the historical presence of other predatory species. The establishment of these species near their northern range boundaries is primarily determined by abiotic factors at a regional scale; however, biotic factors become important at the lake‐to‐lake scale. Studies of exotic species invasions have previously highlighted how spatial scale mediates the importance of abiotic vs. biotic factors on species establishment. Our study demonstrates how concepts from invasion biology can inform our understanding of the factors controlling species distributions with changing climate.     

The role of biotic interactions in shaping distributions and realised assemblages of species: implications for species distribution modelling

Predicting which species will occur together in the future, and where, remains one of the greatest challenges in ecology, and requires a sound understanding of how the abiotic and biotic environments interact with dispersal processes and history across scales. Biotic interactions and their dynamics influence species' relationships to climate, and this also has important implications for predicting future distributions of species. It is already well accepted that biotic interactions shape species' spatial distributions at local spatial extents, but the role of these interactions beyond local extents (e.g. 10 km² to global extents) are usually dismissed as unimportant. In this review we consolidate evidence for how biotic interactions shape species distributions beyond local extents and review methods for integrating biotic interactions into species distribution modelling tools. Drawing upon evidence from contemporary and palaeoecological studies of individual species ranges, functional groups, and species richness patterns, we show that biotic interactions have clearly left their mark on species distributions and realised assemblages of species across all spatial extents. We demonstrate this with examples from within and across trophic groups. A range of species distribution modelling tools is available to quantify species environmental relationships and predict species occurrence, such as: (i) integrating pairwise dependencies, (ii) using integrative predictors, and (iii) hybridising species distribution models (SDMs) with dynamic models. These methods have typically only been applied to interacting pairs of species at a single time, require a priori ecological knowledge about which species interact, and due to data paucity must assume that biotic interactions are constant in space and time. To better inform the future development of these models across spatial scales, we call for accelerated collection of spatially and temporally explicit species data. Ideally, these data should be sampled to reflect variation in the underlying environment across large spatial extents, and at fine spatial resolution. Simplified ecosystems where there are relatively few interacting species and sometimes a wealth of existing ecosystem monitoring data (e.g. arctic, alpine or island habitats) offer settings where the development of modelling tools that account for biotic interactions may be less difficult than elsewhere.     

Spatial predictions at the community level: from current approaches to future frameworks

A fundamental goal of ecological research is to understand and model how processes generate patterns so that if conditions change, changes in the patterns can be predicted. Different approaches have been proposed for modelling species assemblage, but their use to predict spatial patterns of species richness and other community attributes over a range of spatial and temporal scales remains challenging. Different methods emphasize different processes of structuring communities and different goals. In this review, we focus on models that were developed for generating spatially explicit predictions of communities, with a particular focus on species richness, composition, relative abundance and related attributes. We first briefly describe the concepts and theories that span the different drivers of species assembly. A combination of abiotic processes and biotic mechanisms are thought to influence the community assembly process. In this review, we describe four categories of drivers: (i) historical and evolutionary, (ii) environmental, (iii) biotic, and (iv) stochastic. We discuss the different modelling approaches proposed or applied at the community level and examine them from different standpoints, i.e. the theoretical bases, the drivers included, the source data, and the expected outputs, with special emphasis on conservation needs under climate change. We also highlight the most promising novelties, possible shortcomings, and potential extensions of existing methods. Finally, we present new approaches to model and predict species assemblages by reviewing promising ‘integrative frameworks’ and views that seek to incorporate all drivers of community assembly into a unique modelling workflow. We discuss the strengths and weaknesses of these new solutions and how they may hasten progress in community‐level modelling.     

Biotic homogenisation and differentiation as directional change in beta diversity: synthesising driver–response relationships to develop conceptual models across ecosystems

Biotic homogenisation is defined as decreasing dissimilarity among ecological assemblages sampled within a given spatial area over time. Biotic differentiation, in turn, is defined as increasing dissimilarity over time. Overall, changes in the spatial dissimilarities among assemblages (termed ‘beta diversity’) is an increasingly recognised feature of broader biodiversity change in the Anthropocene. Empirical evidence of biotic homogenisation and biotic differentiation remains scattered across different ecosystems. Most meta-analyses quantify the prevalence and direction of change in beta diversity, rather than attempting to identify underlying ecological drivers of such changes. By conceptualising the mechanisms that contribute to decreasing or increasing dissimilarity in the composition of ecological assemblages across space, environmental managers and conservation practitioners can make informed decisions about what interventions may be required to sustain biodiversity and can predict potential biodiversity outcomes of future disturbances. We systematically reviewed and synthesised published empirical evidence for ecological drivers of biotic homogenisation and differentiation across terrestrial, marine, and freshwater realms to derive conceptual models that explain changes in spatial beta diversity. We pursued five key themes in our review: (i) temporal environmental change; (ii) disturbance regime; (iii) connectivity alteration and species redistribution; (iv) habitat change; and (v) biotic and trophic interactions. Our first conceptual model highlights how biotic homogenisation and differentiation can occur as a function of changes in local (alpha) diversity or regional (gamma) diversity, independently of species invasions and losses due to changes in species occurrence among assemblages. Second, the direction and magnitude of change in beta diversity depends on the interaction between spatial variation (patchiness) and temporal variation (synchronicity) of disturbance events. Third, in the context of connectivity and species redistribution, divergent beta diversity outcomes occur as different species have different dispersal characteristics, and the magnitude of beta diversity change associated with species invasions also depends strongly on alpha and gamma diversity prior to species invasion. Fourth, beta diversity is positively linked with spatial environmental variability, such that biotic homogenisation and differentiation occur when environmental heterogeneity decreases or increases, respectively. Fifth, species interactions can influence beta diversity via habitat modification, disease, consumption (trophic dynamics), competition, and by altering ecosystem productivity. Our synthesis highlights the multitude of mechanisms that cause assemblages to be more or less spatially similar in composition (taxonomically, functionally, phylogenetically) through time. We consider that future studies should aim to enhance our collective understanding of ecological systems by clarifying the underlying mechanisms driving homogenisation or differentiation, rather than focusing only on reporting the prevalence and direction of change in beta diversity, per se.     

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.     

Mutualism influences species distribution predictions for a bromeliad‐breeding anuran under climate change

Ecological niche models, or species distribution models, have been widely used to identify potentially suitable areas for species in future climate change scenarios. However, there are inherent errors to these models due to their inability to evaluate species occurrence influenced by non‐climatic factors. With the intuit to improve the modelling predictions for a bromeliad‐breeding treefrog (Phyllodytes melanomystax, Hylidae), we investigate how the climatic suitability of bromeliads influences the distribution model for the treefrog in the context of baseline and 2050 climate change scenarios. We used point occurrence data on the frog and the bromeliad (Vriesea procera, Bromeliaceae) to generate their predicted distributions based on baseline and 2050 climates. Using a consensus of five algorithms, we compared the accuracy of the models and the geographic predictions for the frog generated from two modelling procedures: (i) a climate‐only model for P. melanomystax and V. procera; and (ii) a climate‐biotic model for P. melanomystax, in which the climatic suitability of the bromeliad was jointly considered with the climatic variables. Both modelling approaches generated strong and similar predictive power for P. melanomystax, yet climate‐biotic modelling generated more concise predictions, particularly for the year 2050. Specifically, because the predicted area of the bromeliad overlaps with the predictions for the treefrog in the baseline climate, both modelling approaches produce reasonable similar predicted areas for the anuran. Alternatively, due to the predicted loss of northern climatically suitable areas for the bromeliad by 2050, only the climate‐biotic models provide evidence that northern populations of P. melanomystax will likely be negatively affected by 2050.     

Flow‐mediated predator–prey dynamics influence fish populations in a tropical river

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Higher spring temperatures increase food scarcity and limit the current and future distributions of crossbills

Aim

Understanding how climate affects species distributions remains a major challenge, with the relative importance of direct physiological effects versus biotic interactions still poorly understood. We focus on three species of resource specialists (crossbill Loxia finches) to assess the role of climate in determining the seasonal availability of their food, the importance of climate and the occurrence of their food plants for explaining their current distributions, and to predict changes in their distributions under future climate change scenarios.

Location

Europe.

Methods

We used datasets on the timing of seed fall in European Scots pine Pinus sylvestris forests (where different crossbill species occur) to estimate seed fall phenology and climate data to determine its influence on spatial and temporal variation in the timing of seed fall to provide a link between climate and seed scarcity for crossbills. We used large‐scale datasets on crossbill distribution, cover of the conifers relied on by the three crossbill species and climate variables associated with timing of seed fall, to assess their relative importance for predicting crossbill distributions. We used species distribution modelling to predict changes in their distributions under climate change projections for 2070.

Results

We found that seed fall occurred 1.5–2 months earlier in southern Europe than in Sweden and Scotland and was associated with variation in spring maximum temperatures and precipitation. These climate variables and area covered with conifers relied on by the crossbills explained much of their observed distributions. Projections under global change scenarios revealed reductions in potential crossbill distributions, especially for parrot crossbills.

Main conclusions

Ranges of resource specialists are directly influenced by the presence of their food plants, with climate conditions further affecting resource availability and the window of food scarcity indirectly. Future distributions will be determined by tree responses to changing climatic conditions and the impact of climate on seed fall phenology.

     

Biotic interactions influence the projected distribution of a specialist mammal under climate change

Aim

To measure the effects of including biotic interactions on climate‐based species distribution models (SDMs) used to predict distribution shifts under climate change. We evaluated the performance of distribution models for an endangered marsupial, the northern bettong (Bettongia tropica), comparing models that used only climate variables with models that also took into account biotic interactions.

Location

North‐east Queensland, Australia.

Methods

We developed separate climate‐based distribution models for the northern bettong, its two main resources and a competitor species. We then constructed models for the northern bettong by including climate suitability estimates for the resources and competitor as additional predictor variables to make climate + resource and climate + resource + competition models. We projected these models onto seven future climate scenarios and compared predictions of northern bettong distribution made by these differently structured models, using a ‘global’ metric, the I similarity statistic, to measure overlap in distribution and a ‘local’ metric to identify where predictions differed significantly.

Results

Inclusion of food resource biotic interactions improved model performance. Over moderate climate changes, up to 3.0 °C of warming, the climate‐only model for the northern bettong gave similar predictions of distribution to the more complex models including interactions, with differences only at the margins of predicted distributions. For climate changes beyond 3.0 °C, model predictions diverged significantly. The interactive model predicted less contraction of distribution than the simpler climate‐only model.

Main conclusions

Distribution models that account for interactions with other species, in particular direct resources, improve model predictions in the present‐day climate. For larger climate changes, shifts in distribution of interacting species cause predictions of interactive models to diverge from climate‐only models. Incorporating interactions with other species in SDMs may be needed for long‐term prediction of changes in distribution of species under climate change, particularly for specialized species strongly dependent on a small number of biotic interactions.     

Spatial analysis of taxonomic and genetic patterns and their potential for understanding evolutionary histories

Aim The aim of this research is to develop and investigate methods for the spatial analysis of diversity based on genetic and taxonomic units of difference. We use monophyletic groups of species to assess the potential for these diversity indices to elucidate the geographical components of macro‐scaled evolutionary processes. Location The range occupied by Pultenaea species in temperate and sub‐tropical eastern Australia, extending from western South Australia (133° E–32° S) to Tasmania (146° E–43° S) to coastal central Queensland (148° E–20° S). Methods We applied a series of both spatially explicit and spatially implicit analyses to explore the nature of diversity patterns in the genus Pultenaea, Fabaceae. We first analysed the eastern species as a whole and then the phylogenetic groups within them. We delineated patterns of endemism and biotic (taxon) regions that have been traditionally circumscribed in biogeographical studies of taxa. Centres of endemism were calculated using corrected weighted endemism at a range of spatial scales. Biotic regions were defined by comparing the similarity of species assemblages of grid cells using the Jaccard index and clustering similar cells using hierarchical clustering. On the basis that genetically coherent areas were likely to be more evolutionary informative than species patterns, genetic indices of similarity and difference were derived. A matrix of similarity distances between taxa was generated based on the number of shared informative characters of two sections of trnL‐F and ndhF chloroplast nuclear regions. To identify genetically similar areas, we clustered cells using the mean genetic similarities of the species contained within each pair of cells. Measures of the mean genetic similarity of species in areas were delineated using a geographically local multi‐scalar approach. Resultant patterns of genetic diversity are interpreted in relation to theories of the evolutionary relationships between species and species groups. Results Centres of Pultenaea endemism were defined, those of clades 1 congruent with the spatially separated centres of clades 2 and 3. The taxonomic classification analysis defined cells with shared groups of species, which in some cases clustered when plotted in geographic space, defining biotic regions. In some instances the distribution of biotic regions was congruent with centres of endemism, however larger scale groupings were also apparent. In clade 1 one set of species was replaced by another along the extent of the range, with some connectivity between some geographically disjunct regions due to the presence of widespread species. In the combined analysis of clade 2 and 3 species the major biotic (taxonomic) groups with geographic coherence were defined by species in the respective clades, representing the geographic separation of these clades. However distinctive biotic regions within these main groupings of clades 2 and 3 were also apparent. Clustering cells using the mean genetic similarities of the species contained within each pair of cells indicated that some of the taxonomically defined biotic boundaries were the result of changes in composition of closely related species. This was most apparent in clades 1 and 2 where most cells were highly genetically similar. In clade 3 genetically distinct groups remained and were in part defined by sister taxa with disjunct distributions. Gradients in mean genetic similarity became more apparent from small to larger scales of analysis. At larger scales of analysis, regions of different levels of genetic diversity were delineated. Regions with highest diversity levels (lowest level of similarity) often represented regions where the ranges of phylogenetically distinctive species intergraded. Main conclusions The combined analysis of diversity, phylogeny and geography has potential to reveal macro‐scaled evolutionary patterns from which evolutionary processes may be inferred. The spatial genetic diversity indices developed in this study contribute new methods for identifying coherent evolutionary units in the landscape, which overcome some of the limitations of using taxonomic data, and from which the role of geography in evolutionary processes can be tested. We also conclude that a multiple‐index approach to diversity pattern analysis is useful, especially where patterns may be the result of a long history of different environmental changes and related evolutionary events. The analysis contributes to the knowledge of large‐scale diversity patterns of Pultenaea which has relevance for the assessment of the conservation status of the genus.     

Sap‐feeding insects on forest trees along latitudinal gradients in northern Europe: a climate‐driven patterns

Knowledge of the latitudinal patterns in biotic interactions, and especially in herbivory, is crucial for understanding the mechanisms that govern ecosystem functioning and for predicting their responses to climate change. We used sap‐feeding insects as a model group to test the hypotheses that the strength of plant–herbivore interactions in boreal forests decreases with latitude and that this latitudinal pattern is driven primarily by midsummer temperatures. We used a replicated sampling design and quantitatively collected and identified all sap‐feeding insects from four species of forest trees along five latitudinal gradients (750–1300 km in length, ten sites in each gradient) in northern Europe (59 to 70°N and 10 to 60°E) during 2008–2011. Similar decreases in diversity of sap‐feeding insects with latitude were observed in all gradients during all study years. The sap‐feeder load (i.e. insect biomass per unit of foliar biomass) decreased with latitude in typical summers, but increased in an exceptionally hot summer and was independent of latitude during a warm summer. Analysis of combined data from all sites and years revealed dome‐shaped relationships between the loads of sap‐feeders and midsummer temperatures, peaking at 17 °C in Picea abies, at 19.5 °C in Pinus sylvestris and Betula pubescens and at 22 °C in B. pendula. From these relationships, we predict that the losses of forest trees to sap‐feeders will increase by 0–45% of the current level in southern boreal forests and by 65–210% in subarctic forests with a 1 °C increase in summer temperatures. The observed relationships between temperatures and the loads of sap‐feeders differ between the coniferous and deciduous tree species. We conclude that climate warming will not only increase plant losses to sap‐feeding insects, especially in subarctic forests, but can also alter plant‐plant interactions, thereby affecting both the productivity and the structure of future forest ecosystems.     

Determining the factors affecting the distribution of Muscari latifolium,an endemic plant of Turkey,and a mapping species distribution model

Species distribution modeling was used to determine factors among the large predictor candidate data set that affect the distribution of Muscari latifolium , an endemic bulbous plant species of Turkey, to quantify the relative importance of each factor and make a potential spatial distribution map of M. latifolium . Models were built using the Boosted Regression Trees method based on 35 presence and 70 absence records obtained through field sampling in the Gönen Dam watershed area of the Kazda?? Mountains in West Anatolia. Large candidate variables of monthly and seasonal climate, fine‐scale land surface, and geologic and biotic variables were simplified using a BRT simplifying procedure. Analyses performed on these resources, direct and indirect variables showed that there were 14 main factors that influence the species’ distribution. Five of the 14 most important variables influencing the distribution of the species are bedrock type, Quercus cerris density, precipitation during the wettest month, Pinus nigra density, and northness. These variables account for approximately 60% of the relative importance for determining the distribution of the species. Prediction performance was assessed by 10 random subsample data sets and gave a maximum the area under a receiver operating characteristic curve (AUC) value of 0.93 and an average AUC value of 0.8. This study provides a significant contribution to the knowledge of the habitat requirements and ecological characteristics of this species. The distribution of this species is explained by a combination of biotic and abiotic factors. Hence, using biotic interaction and fine‐scale land surface variables in species distribution models improved the accuracy and precision of the model. The knowledge of the relationships between distribution patterns and environmental factors and biotic interaction of M. latifolium can help develop a management and conservation strategy for this species.     

Parapatric species and the implications for climate change studies: a case study on hares in Europe

Parapatry is a biogeographical term used to refer to organisms whose ranges do not overlap, but are immediately adjacent to each other; they only co‐occur – if at all – in a narrow contact zone. Often there are no environmental barriers in the contact zones, hence competitive interaction is usually advocated as the factor that modulates species distribution ranges. Even though the effects of climate change on species distribution have been widely studied, few studies have explored these effects on the biogeographical relationships between closely related, parapatric, species. We modelled environmental favourability for three parapatric hare species in Europe – Lepus granatensis, L. europaeus and L. timidus – using ecogeographical variables and projected the models into the future according to the IPCC A2 emissions scenario. Favourabilities for present and future scenarios were combined using fuzzy logic with the following aims: (i) to determine the biogeographical relationships between hare species in parapatry, that is L. granatensis/L. europaeus and L. europaeus/L. timidus and (ii) to assess the effects of climate change on each species as well as on their interspecific interactions. In their contact area L. granatensis achieved higher favourability values than L. europaeus, suggesting that if both species have a similar population status, the former species may have some advantages over the latter if competitive relationships are established. Climate change had the most striking effect on the distribution of L. timidus, especially when interspecific interactions with L. europaeus were taken into account, which may compromise the co‐existence of L. timidus. The results of this study are relevant not only for understanding the distribution patterns of the hares studied and the effects of climate change on these patterns, but also for improving the general application of species distribution models to the prediction of the effects of climate change on biodiversity.     

Habitat shifts of endangered species under altered climate conditions: importance of biotic interactions

Predicting changes in potential habitat for endangered species as a result of global warming requires considering more than future climate conditions; it is also necessary to evaluate biotic associations. Most distribution models predicting species responses to climate change include climate variables and occasionally topographic and edaphic parameters, rarely are biotic interactions included. Here, we incorporate biotic interactions into niche models to predict suitable habitat for species under altered climates. We constructed and evaluated niche models for an endangered butterfly and a threatened bird species, both are habitat specialists restricted to semiarid shrublands of southern California. To incorporate their dependency on shrubs, we first developed climate‐based niche models for shrubland vegetation and individual shrub species. We also developed models for the butterfly's larval host plants. Outputs from these models were included in the environmental variable dataset used to create butterfly and bird niche models. For both animal species, abiotic–biotic models outperformed the climate‐only model, with climate‐only models over‐predicting suitable habitat under current climate conditions. We used the climate‐only and abiotic–biotic models to calculate amounts of suitable habitat under altered climates and to evaluate species' sensitivities to climate change. We varied temperature (+0.6, +1.7, and +2.8 °C) and precipitation (50%, 90%, 100%, 110%, and 150%) relative to current climate averages and within ranges predicted by global climate change models. Suitable habitat for each species was reduced at all levels of temperature increase. Both species were sensitive to precipitation changes, particularly increases. Under altered climates, including biotic variables reduced habitat by 68–100% relative to the climate‐only model. To design reserve systems conserving sensitive species under global warming, it is important to consider biotic interactions, particularly for habitat specialists and species with strong dependencies on other species.     

Host plant distributions and climate interact to affect the predicted geographic distribution of a Neotropical termite

Species distribution models (SDMs) have been widely used in the scientific literature. The majority of SDMs use climate data or other abiotic variables to forecast the potential distribution of a species in geographic space. Biotic interactions can affect the predicted spatial distribution of a species in many ways across multiple spatial scales, and incorporating these predictors in an SDM is a current topic in the scientific literature. Constrictotermes cyphergaster is a widely distributed termite in the Neotropics. This termite species nests in plants and more frequently nests in some arboreal species. Thus, this species is an excellent model to evaluate the influence of biotic interactions in SDMs. We evaluate the influences of climate and the geographic distribution of host plants on the potential distribution of C. cyphergaster. Three correlative models (MaxEnt) were built to predict the geographic distribution of the termite: (1) climate data, (2) biotic data (i.e., the geographic distribution of host plants), and (3) climate and biotic data. The models that were generated indicate that the potential geographic distribution of C. cyphergaster is concentrated in the Cerrado and Caatinga regions. In addition, path analysis and multiple regression revealed the importance of the direct effects of biological interactions in the geographic distribution of the termite, while climate affected the distribution of the termite mainly through indirect effects by influencing the geographic distributions of host plants. The current study endorses the importance of including biological interactions in SDMs. We recommend using biotic predictors in SDM studies of insect species, mainly because insects have important environmental services and biotic interaction data can improve the macroecological studies of this group.     

Species interactions and climate change: How the disruption of species co‐occurrence will impact on an avian forest guild

Interspecific interactions are crucial in determining species occurrence and community assembly. Understanding these interactions is thus essential for correctly predicting species' responses to climate change. We focussed on an avian forest guild of four hole‐nesting species with differing sensitivities to climate that show a range of well‐understood reciprocal interactions, including facilitation, competition and predation. We modelled the potential distributions of black woodpecker and boreal, tawny and Ural owl, and tested whether the spatial patterns of the more widespread species (excluding Ural owl) were shaped by interspecific interactions. We then modelled the potential future distributions of all four species, evaluating how the predicted changes will alter the overlap between the species' ranges, and hence the spatial outcomes of interactions. Forest cover/type and climate were important determinants of habitat suitability for all species. Field data analysed with N‐mixture models revealed effects of interspecific interactions on current species abundance, especially in boreal owl (positive effects of black woodpecker, negative effects of tawny owl). Climate change will impact the assemblage both at species and guild levels, as the potential area of range overlap, relevant for species interactions, will change in both proportion and extent in the future. Boreal owl, the most climate‐sensitive species in the guild, will retreat, and the range overlap with its main predator, tawny owl, will increase in the remaining suitable area: climate change will thus impact on boreal owl both directly and indirectly. Climate change will cause the geographical alteration or disruption of species interaction networks, with different consequences for the species belonging to the guild and a likely spatial increase of competition and/or intraguild predation. Our work shows significant interactions and important potential changes in the overlap of areas suitable for the interacting species, which reinforce the importance of including relevant biotic interactions in predictive climate change models for increasing forecast accuracy.     

Can we predict ectotherm responses to climate change using thermal performance curves and body temperatures?

Thermal performance curves (TPCs), which quantify how an ectotherm's body temperature (T_b) affects its performance or fitness, are often used in an attempt to predict organismal responses to climate change. Here, we examine the key – but often biologically unreasonable – assumptions underlying this approach; for example, that physiology and thermal regimes are invariant over ontogeny, space and time, and also that TPCs are independent of previously experienced T_b. We show how a critical consideration of these assumptions can lead to biologically useful hypotheses and experimental designs. For example, rather than assuming that TPCs are fixed during ontogeny, one can measure TPCs for each major life stage and incorporate these into stage‐specific ecological models to reveal the life stage most likely to be vulnerable to climate change. Our overall goal is to explicitly examine the assumptions underlying the integration of TPCs with T_b, to develop a framework within which empiricists can place their work within these limitations, and to facilitate the application of thermal physiology to understanding the biological implications of climate change.     

Continental‐scale tree‐ring‐based projection of Douglas‐fir growth: Testing the limits of space‐for‐time substitution

A central challenge in global change research is the projection of the future behavior of a system based upon past observations. Tree‐ring data have been used increasingly over the last decade to project tree growth and forest ecosystem vulnerability under future climate conditions. But how can the response of tree growth to past climate variation predict the future, when the future does not look like the past? Space‐for‐time substitution (SFTS) is one way to overcome the problem of extrapolation: the response at a given location in a warmer future is assumed to follow the response at a warmer location today. Here we evaluated an SFTS approach to projecting future growth of Douglas‐fir (Pseudotsuga menziesii), a species that occupies an exceptionally large environmental space in North America. We fit a hierarchical mixed‐effects model to capture ring‐width variability in response to spatial and temporal variation in climate. We found opposing gradients for productivity and climate sensitivity with highest growth rates and weakest response to interannual climate variation in the mesic coastal part of Douglas‐fir's range; narrower rings and stronger climate sensitivity occurred across the semi‐arid interior. Ring‐width response to spatial versus temporal temperature variation was opposite in sign, suggesting that spatial variation in productivity, caused by local adaptation and other slow processes, cannot be used to anticipate changes in productivity caused by rapid climate change. We thus substituted only climate sensitivities when projecting future tree growth. Growth declines were projected across much of Douglas‐fir's distribution, with largest relative decreases in the semiarid U.S. Interior West and smallest in the mesic Pacific Northwest. We further highlight the strengths of mixed‐effects modeling for reviving a conceptual cornerstone of dendroecology, Cook's 1987 aggregate growth model, and the great potential to use tree‐ring networks and results as a calibration target for next‐generation vegetation models.     

Is subarctic forest advance able to keep pace with climate change?

Recent climate warming and scenarios for further warming have led to expectations of rapid movement of ecological boundaries. Here we focus on the circumarctic forest–tundra ecotone (FTE), which represents an important bioclimatic zone with feedbacks from forest advance and corresponding tundra disappearance (up to 50% loss predicted this century) driving widespread ecological and climatic changes. We address FTE advance and climate history relations over the 20th century, using FTE response data from 151 sites across the circumarctic area and site‐specific climate data. Specifically, we investigate spatial uniformity of FTE advance, statistical associations with 20th century climate trends, and whether advance rates match climate change velocities (CCVs). Study sites diverged into four regions (Eastern Canada; Central and Western Canada and Alaska; Siberia; and Western Eurasia) based on their climate history, although all were characterized by similar qualitative patterns of behaviour (with about half of the sites showing advancing behaviour). The main associations between climate trend variables and behaviour indicate the importance of precipitation rather than temperature for both qualitative and quantitative behaviours, and the importance of non‐growing season as well as growing season months. Poleward latitudinal advance rates differed significantly among regions, being smallest in Eastern Canada (~10 m/year) and largest in Western Eurasia (~100 m/year). These rates were 1–2 orders of magnitude smaller than expected if vegetation distribution remained in equilibrium with climate. The many biotic and abiotic factors influencing FTE behaviour make poleward advance rates matching predicted 21st century CCVs (~10³–10⁴ m/year) unlikely. The lack of empirical evidence for swift forest relocation and the discrepancy between CCV and FTE response contradict equilibrium model‐based assumptions and warrant caution when assessing global‐change‐related biotic and abiotic implications, including land–atmosphere feedbacks and carbon sequestration.