Disentangling scale dependencies in species environmental niches and distributions

Understanding species’ responses to environmental conditions, and how these ­species–environment associations shape spatial distributions, are longstanding goals in ecology and biogeography. However, an essential component of species–environment relationships – the spatial unit, or grain, at which they operate – remains unresolved. We identify three components of scale‐dependence in analyses of species–environment associations: 1) response grain, the grain at which species respond most strongly to their environment; 2) environment spatial structure, the pattern of spatial autocorrelation intrinsic to an environmental factor; and 3) analysis grain, the grain at which analyses are conducted and ecological inferences are made. We introduce a novel conceptual framework that defines these scale components in the context of analyzing species–environment relationships, and provide theoretical examples of their interactions for species with various ecological attributes. We then use a virtual species approach to investigate the impacts of each component on common methods of measuring and predicting species–environment relationships. We find that environment spatial structure has a substantial impact on the ability of even simple, univariate species distribution models (SDMs) to recover known species–­environment associations at coarse analysis grains. For simulated environments with ‘fine’ and ‘intermediate’ spatial structure, model explanatory power, and the frequency with which simple SDMs correctly estimated a virtual species’ response to the simulated environment, dramatically declined as analysis grain increased. Informed by these results, we use a scaling analysis to identify maximum analysis grains for individual environmental factors, and a scale optimization procedure to determine the grain of maximum predictive accuracy. Implementing these analysis grain thresholds and model performance standards in an example east African study system yields more accurate distribution predictions, compared to SDMs independently constructed at arbitrary analysis grains. Finally, we integrate our conceptual framework with virtual and empirical results to provide practical recommendations for researchers asking common questions about species–environment relationships.     

Trade-offs in covariate selection for species distribution models: a methodological comparison

Species distribution models (SDMs) are a common approach to describing species’ space-use and spatially-explicit abundance. With a myriad of model types, methods and parameterization options available, it is challenging to make informed decisions about how to build robust SDMs appropriate for a given purpose. One key component of SDM development is the appropriate parameterization of covariates, such as the inclusion of covariates that reflect underlying processes (e.g. abiotic and biotic covariates) and covariates that act as proxies for unobserved processes (e.g. space and time covariates). It is unclear how different SDMs apportion variance among a suite of covariates, and how parameterization decisions influence model accuracy and performance. To examine trade-offs in covariation parameterization in SDMs, we explore the attribution of spatiotemporal and environmental variation across a suite of SDMs. We first used simulated species distributions with known environmental preferences to compare three types of SDM: a machine learning model (boosted regression tree), a semi-parametric model (generalized additive model) and a spatiotemporal mixed-effects model (vector autoregressive spatiotemporal model, VAST). We then applied the same comparative framework to a case study with three fish species (arrowtooth flounder, pacific cod and walleye pollock) in the eastern Bering Sea, USA. Model type and covariate parameterization both had significant effects on model accuracy and performance. We found that including either spatiotemporal or environmental covariates typically reproduced patterns of species distribution and abundance across the three models tested, but model accuracy and performance was maximized when including both spatiotemporal and environmental covariates in the same model framework. Our results reveal trade-offs in the current generation of SDM tools between accurately estimating species abundance, accurately estimating spatial patterns, and accurately quantifying underlying species–environment relationships. These comparisons between model types and parameterization options can help SDM users better understand sources of model bias and estimate error.     

Incorporating spatial autocorrelation into species distribution models alters forecasts of climate‐mediated range shifts

Species distribution models (SDMs) are widely used to forecast changes in the spatial distributions of species and communities in response to climate change. However, spatial autocorrelation (SA) is rarely accounted for in these models, despite its ubiquity in broad‐scale ecological data. While spatial autocorrelation in model residuals is known to result in biased parameter estimates and the inflation of type I errors, the influence of unmodeled SA on species' range forecasts is poorly understood. Here we quantify how accounting for SA in SDMs influences the magnitude of range shift forecasts produced by SDMs for multiple climate change scenarios. SDMs were fitted to simulated data with a known autocorrelation structure, and to field observations of three mangrove communities from northern Australia displaying strong spatial autocorrelation. Three modeling approaches were implemented: environment‐only models (most frequently applied in species' range forecasts), and two approaches that incorporate SA; autologistic models and residuals autocovariate (RAC) models. Differences in forecasts among modeling approaches and climate scenarios were quantified. While all model predictions at the current time closely matched that of the actual current distribution of the mangrove communities, under the climate change scenarios environment‐only models forecast substantially greater range shifts than models incorporating SA. Furthermore, the magnitude of these differences intensified with increasing increments of climate change across the scenarios. When models do not account for SA, forecasts of species' range shifts indicate more extreme impacts of climate change, compared to models that explicitly account for SA. Therefore, where biological or population processes induce substantial autocorrelation in the distribution of organisms, and this is not modeled, model predictions will be inaccurate. These results have global importance for conservation efforts as inaccurate forecasts lead to ineffective prioritization of conservation activities and potentially to avoidable species extinctions.     

Forecasting species range dynamics with process‐explicit models: matching methods to applications

Knowing where species occur is fundamental to many ecological and environmental applications. Species distribution models (SDMs) are typically based on correlations between species occurrence data and environmental predictors, with ecological processes captured only implicitly. However, there is a growing interest in approaches that explicitly model processes such as physiology, dispersal, demography and biotic interactions. These models are believed to offer more robust predictions, particularly when extrapolating to novel conditions. Many process–explicit approaches are now available, but it is not clear how we can best draw on this expanded modelling toolbox to address ecological problems and inform management decisions. Here, we review a range of process–explicit models to determine their strengths and limitations, as well as their current use. Focusing on four common applications of SDMs – regulatory planning, extinction risk, climate refugia and invasive species – we then explore which models best meet management needs. We identify barriers to more widespread and effective use of process‐explicit models and outline how these might be overcome. As well as technical and data challenges, there is a pressing need for more thorough evaluation of model predictions to guide investment in method development and ensure the promise of these new approaches is fully realised.     

Spatial autocorrelation in predictors reduces the impact of positional uncertainty in occurrence data on species distribution modelling

Aim To investigate the impact of positional uncertainty in species occurrences on the predictions of seven commonly used species distribution models (SDMs), and explore its interaction with spatial autocorrelation in predictors. Methods A series of artificial datasets covering 155 scenarios including different combinations of five positional uncertainty scenarios and 31 spatial autocorrelation scenarios were simulated. The level of positional uncertainty was defined by the standard deviation of a normally distributed zero‐mean random variable. Each dataset included two environmental gradients (predictor variables) and one set of species occurrence sample points (response variable). Seven commonly used models were selected to develop SDMs: generalized linear models, generalized additive models, boosted regression trees, multivariate adaptive regression spline, random forests, genetic algorithm for rule‐set production and maximum entropy. A probabilistic approach was employed to model and simulate five levels of error in the species locations. To analyse the propagation of positional uncertainty, Monte Carlo simulation was applied to each scenario for each SDM. The models were evaluated for performance using simulated independent test data with Cohen’s Kappa and the area under the receiver operating characteristic curve. Results Positional uncertainty in species location led to a reduction in prediction accuracy for all SDMs, although the magnitude of the reduction varied between SDMs. In all cases the magnitude of this impact varied according to the degree of spatial autocorrelation in predictors and the levels of positional uncertainty. It was shown that when the range of spatial autocorrelation in the predictors was less than or equal to three times the standard deviation of the positional error, the models were less affected by error and, consequently, had smaller decreases in prediction accuracy. When the range of spatial autocorrelation in predictors was larger than three times the standard deviation of positional error, the prediction accuracy was low for all scenarios. Main conclusions The potential impact of positional uncertainty in species occurrences on the predictions of SDMs can be understood by comparing it with the spatial autocorrelation range in predictor variables.     

Do species distribution models explain spatial structure within tree species ranges?

Aim To evaluate the ability of species distribution models (SDMs) to predict the spatial structure of tree species within their geographical ranges (how trees are distributed within their ranges). Location Continental Spain. Methods We used an extensive dataset consisting of c. 90,000 plots (1 plot km^?2) where presence/absence data for 23 common Mediterranean and Atlantic tree species had been surveyed. We first generated SDMs relating the presence or absence of each species to a set of 16 environmental predictors, following a stepwise modelling process based on maximum likelihood methods. Superimposing spatial correlograms generated from the predictions of the SDMs over those generated from the raw data allowed a model–observation comparison of the nature, scale and intensity (level of aggregation) of spatial structure with the species ranges. Results SDMs predicted accurately the nature and scale of the spatial structure of trees. However, for most species, the observed intensity of spatial structure (level of aggregation of species in space) was substantially greater than that predicted by the SDMs. On average, the intensity of spatial aggregation was twice that predicted by SDMs. In addition, we also found a negative correlation between intensity of aggregation and species range size. Main conclusions Standard SDM predictions of spatial structure patterns differ among species. SDMs are apparently able to reproduce both the scale and intensity of species spatial structure within their ranges. However, one or more missing processes not included in SDMs results in species being substantially more aggregated in space than can be captured by the SDMs. This result adds to recent calls for a new generation of more biologically realistic SDMs. In particular, future SDMs should incorporate ecological processes that are likely to increase the intensity of spatial aggregation, such as source–sink dynamics, fine‐scale environmental heterogeneity and disequilibrium.     

Temporal resolutions in species distribution models of highly mobile marine animals: Recommendations for ecologists and managers

While ecologists have long recognized the influence of spatial resolution on species distribution models (SDMs), they have given relatively little attention to the influence of temporal resolution. Considering temporal resolutions is critical in distribution modelling of highly mobile marine animals, as they interact with dynamic oceanographic processes that vary at time‐scales from seconds to decades. We guide ecologists in selecting temporal resolutions that best match ecological questions and ecosystems, and managers in applying these models. We group the temporal resolutions of environmental variables used in SDMs into three classes: instantaneous, contemporaneous and climatological. We posit that animal associations with fine‐scale and ephemeral features are best modelled with instantaneous covariates. Associations with large scale and persistent oceanographic features are best modelled with climatological covariates. Associations with mesoscale features are best modelled with instantaneous or contemporaneous covariates if ephemeral processes are present or interannual variability occurs, and climatological covariates if seasonal processes dominate and interannual variability is weak.     

Limited transferability of stream‐fish distribution models among river catchments: reasons and implications

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Towards novel approaches to modelling biotic interactions in multispecies assemblages at large spatial extents

Aim Biotic interactions – within guilds or across trophic levels – have widely been ignored in species distribution models (SDMs). This synthesis outlines the development of ‘species interaction distribution models’ (SIDMs), which aim to incorporate multispecies interactions at large spatial extents using interaction matrices. Location Local to global. Methods We review recent approaches for extending classical SDMs to incorporate biotic interactions, and identify some methodological and conceptual limitations. To illustrate possible directions for conceptual advancement we explore three principal ways of modelling multispecies interactions using interaction matrices: simple qualitative linkages between species, quantitative interaction coefficients reflecting interaction strengths, and interactions mediated by interaction currencies. We explain methodological advancements for static interaction data and multispecies time series, and outline methods to reduce complexity when modelling multispecies interactions. Results Classical SDMs ignore biotic interactions and recent SDM extensions only include the unidirectional influence of one or a few species. However, novel methods using error matrices in multivariate regression models allow interactions between multiple species to be modelled explicitly with spatial co‐occurrence data. If time series are available, multivariate versions of population dynamic models can be applied that account for the effects and relative importance of species interactions and environmental drivers. These methods need to be extended by incorporating the non‐stationarity in interaction coefficients across space and time, and are challenged by the limited empirical knowledge on spatio‐temporal variation in the existence and strength of species interactions. Model complexity may be reduced by: (1) using prior ecological knowledge to set a subset of interaction coefficients to zero, (2) modelling guilds and functional groups rather than individual species, and (3) modelling interaction currencies and species’ effect and response traits. Main conclusions There is great potential for developing novel approaches that incorporate multispecies interactions into the projection of species distributions and community structure at large spatial extents. Progress can be made by: (1) developing statistical models with interaction matrices for multispecies co‐occurrence datasets across large‐scale environmental gradients, (2) testing the potential and limitations of methods for complexity reduction, and (3) sampling and monitoring comprehensive spatio‐temporal data on biotic interactions in multispecies communities.     

Hierarchical models facilitate spatial analysis of large data sets: a case study on invasive plant species in the northeastern United States

Many critical ecological issues require the analysis of large spatial point data sets – for example, modelling species distributions, abundance and spread from survey data. But modelling spatial relationships, especially in large point data sets, presents major computational challenges. We use a novel Bayesian hierarchical statistical approach, 'spatial predictive process' modelling, to predict the distribution of a major invasive plant species, Celastrus orbiculatus , in the northeastern USA. The model runs orders of magnitude faster than traditional geostatistical models on a large data set of c . 4000 points, and performs better than generalized linear models, generalized additive models and geographically weighted regression in cross-validation. We also use this approach to model simultaneously the distributions of a set of four major invasive species in a spatially explicit multivariate model. This multispecies analysis demonstrates that some pairs of species exhibit negative residual spatial covariation, suggesting potential competitive interaction or divergent responses to unmeasured factors.     

Accounting for multi‐scale spatial autocorrelation improves performance of invasive species distribution modelling (iSDM)

Aim Analyses of species distributions are complicated by various origins of spatial autocorrelation (SAC) in biogeographical data. SAC may be particularly important for invasive species distribution models (iSDMs) because biological invasions are strongly influenced by dispersal and colonization processes that typically create highly structured distribution patterns. We examined the efficacy of using a multi‐scale framework to account for different origins of SAC, and compared non‐spatial models with models that accounted for SAC at multiple levels. Location We modelled the spatial distribution of an invasive forest pathogen, Phytophthora ramorum, in western USA. Methods We applied one conventional statistical method (generalized linear model, GLM) and one nonparametric technique (maximum entropy, Maxent) to a large dataset on P. ramorum occurrence (n = 3787) to develop four types of model that included environmental variables and that either ignored spatial context or incorporated it at a broad scale using trend surface analysis, a local scale using autocovariates, or multiple scales using spatial eigenvector mapping. We evaluated model accuracies and amounts of explained spatial structure, and examined the changes in predictive power of the environmental and spatial variables. Results Accounting for different scales of SAC significantly enhanced the predictive capability of iSDMs. Dramatic improvements were observed when fine‐scale SAC was included, suggesting that local range‐confining processes are important in P. ramorum spread. The importance of environmental variables was relatively consistent across all models, but the explanatory power decreased in spatial models for factors with strong spatial structure. While accounting for SAC reduced the amount of residual autocorrelation for GLM but not for Maxent, it still improved the performance of both approaches, supporting our hypothesis that dispersal and colonization processes are important factors to consider in distribution models of biological invasions. Main conclusions Spatial autocorrelation has become a paradigm in biogeography and ecological modelling. In addition to avoiding the violation of statistical assumptions, accounting for spatial patterns at multiple scales can enhance our understanding of dynamic processes that explain ecological mechanisms of invasion and improve the predictive performance of static iSDMs.     

Accounting for spatial autocorrelation in null models of tree species association

A commonly used null model for species association among forest trees is a well‐mixed community (WMC). A WMC represents a non‐spatial, or spatially implicit, model, in which species form nearest‐neighbor pairs at a rate equal to the product of their community proportions. WMC models assume that the outcome of random dispersal and demographic processes is complete spatial randomness (CSR) in the species’ spatial distributions. Yet, stochastic dispersal processes often lead to spatial autocorrelation (SAC) in tree species densities, giving rise to clustering, segregation, and other nonrandom patterns. Although methods exist to account for SAC in spatially‐explicit models, its impact on non‐spatial models often remains unaccounted for. To investigate the potential for SAC to bias tests based upon non‐spatial models, we developed a spatially‐heterogeneous (SH) modelling approach that incorporates measured levels of SAC. Using the mapped locations of individuals in a tropical tree community, we tested the hypothesis that the identity of nearest‐neighbors represents a random draw from neighborhood species pools. Correlograms of Moran's I confirmed that, for 50 of 51 dominant species, stem density was significantly autocorrelated over distances ranging from 50 to 200 m. The observed patterns of SAC were consistent with dispersal limitation, with most species occurring in distinct patches. For nearly all of the 106 species in the community, the frequency of pairwise association was statistically indistinguishable from that projected by the null models. However, model comparisons revealed that non‐spatial models more strongly underestimated observed species‐pair frequencies, particularly for conspecific pairs. Overall, the CSR models projected more significant facilitative interactions than did SH models, yielding a more liberal test of niche differences. Our results underscore the importance of accounting for stochastic spatial processes in tests of association, regardless of whether spatial or non‐spatial models are employed.     

Disentangling drivers of spatial autocorrelation in species distribution models

Species distribution models (SDMs) are frequently used to understand the influence of site properties on species occurrence. For robust model inference, SDMs need to account for the spatial autocorrelation of virtually all species occurrence data. Current methods do not routinely distinguish between extrinsic and intrinsic drivers of spatial autocorrelation, although these may have different implications for conservation. Here, we present and test a method that disentangles extrinsic and intrinsic drivers of spatial autocorrelation using repeated observations of a species. We focus on unknown habitat characteristics and conspecific interactions as extrinsic and intrinsic drivers, respectively. We model the former with spatially correlated random effects and the latter with an autocovariate, such that the spatially correlated random effects are constant across the repeated observations whereas the autocovariate may change. We tested the performance of our model on virtual species data and applied it to observations of the corncrake Crex crex in the Netherlands. Applying our model to virtual species data revealed that it was well able to distinguish between the two different drivers of spatial autocorrelation, outperforming models with no or a single component for spatial autocorrelation. This finding was independent of the direction of the conspecific interactions (i.e. conspecific attraction versus competitive exclusion). The simulations confirmed that the ability of our model to disentangle both drivers of autocorrelation depends on repeated observations. In the case study, we discovered that the corncrake has a stronger response to habitat characteristics compared to a model that did not include spatially correlated random effects, whereas conspecific interactions appeared to be less important. This implies that future conservation efforts should primarily focus on maximizing habitat availability. Our study shows how to systematically disentangle extrinsic and intrinsic drivers of spatial autocorrelation. The method we propose can help to correctly identify the main drivers of species distributions.     

Predicting species distribution: offering more than simple habitat models

In the last two decades, interest in species distribution models (SDMs) of plants and animals has grown dramatically. Recent advances in SDMs allow us to potentially forecast anthropogenic effects on patterns of biodiversity at different spatial scales. However, some limitations still preclude the use of SDMs in many theoretical and practical applications. Here, we provide an overview of recent advances in this field, discuss the ecological principles and assumptions underpinning SDMs, and highlight critical limitations and decisions inherent in the construction and evaluation of SDMs. Particular emphasis is given to the use of SDMs for the assessment of climate change impacts and conservation management issues. We suggest new avenues for incorporating species migration, population dynamics, biotic interactions and community ecology into SDMs at multiple spatial scales. Addressing all these issues requires a better integration of SDMs with ecological theory.     

Equilibrium or not? Modelling potential distribution of invasive species in different stages of invasion

Aim The assumption of equilibrium between organisms and their environment is a standard working postulate in species distribution models (SDMs). However, this assumption is typically violated in models of biological invasions where range expansions are highly constrained by dispersal and colonization processes. Here, we examined how stage of invasion affects the extent to which occurrence data represent the ecological niche of organisms and, in turn, influences spatial prediction of species’ potential distributions. Location Six ecoregions in western Oregon, USA. Methods We compiled occurrence data from 697 field plots collected over a 9‐year period (2001–09) of monitoring the spread of invasive forest pathogen Phytophthora ramorum. Using these data, we applied ecological‐niche factor analysis to calibrate models of potential distribution across different years of colonization. We accounted for natural variation and uncertainties in model evaluation by further investigating three hypothetical scenarios of varying equilibrium in a simulated virtual species, for which the ‘true’ potential distribution was known. Results We confirm our hypothesis that SDMs calibrated in early stages of invasion are less accurate than models calibrated under scenarios closer to equilibrium. SDMs that are developed in early stages of invasion tend to underpredict the potential range compared to models that are built in later stages of invasion. Main conclusions A full environmental niche of invasive species cannot be effectively captured with data from a realized distribution that is restricted by processes preventing full occupancy of suitable habitats. If SDMs are to be used effectively in conservation and management, stage of invasion needs to be considered to avoid underestimation of habitats at risk of invasion.     

Community versus single‐species distribution models for British plants

Aim Species distribution models are increasingly used to predict the impacts of global change on whole ecological communities by modelling the individualistic niche responses of large numbers of species. However, it is not clear whether this single‐species ensemble approach is preferable to community‐wide strategies that represent interspecific associations or shared responses to environmental gradients. Here, we test the performance of two multi‐species modelling approaches against equivalent single‐species models. Location Great Britain. Methods Single‐ and multi‐species distribution models were fitted for 701 native British plant species at a 10‐km grid scale. Two machine learning methods were used – classification and regression trees (CARTs) and artificial neural networks (ANNs). The single‐species versions are widely used in ecology but their multivariate extensions are less well known and have not previously been evaluated against one another. We compared their abilities to predict species distributions, community compositions and species richness in an independent geographical region reserved from model‐fitting. Results The single‐ and multi‐species models performed similarly, although the community models gave slightly poorer predictive accuracy by all measures. However, from the point of view of the whole community they were much simpler than the array of single‐species models, involving orders of magnitude fewer parameters. Multi‐species approaches also left greater residual spatial autocorrelation than the individualistic models and, contrary to expectation, were relatively less accurate for rarer species. However, the fitted multi‐species response curves had lower tendency for pronounced discontinuities that are unlikely to be a feature of realized niche responses. Main conclusions Although community distribution models were slightly less accurate than single‐species models, they offered a highly simplified way of modelling spatial patterns in British plant diversity. Moreover, an advantage of the multi‐species approach was that the modelling of shared environmental responses resolved more realistic response curves. However, there was a slight tendency for community models to predict rare species less accurately, which is potentially disadvantageous for conservation applications. We conclude that multi‐species distribution models may have potential for understanding and predicting the structure of ecological communities, but were slightly inferior to single‐species ensembles for our data.     

A new method for dealing with residual spatial autocorrelation in species distribution models

Species distribution modelling (SDM) is a widely used tool and has many applications in ecology and conservation biology. Spatial autocorrelation (SAC), a pattern in which observations are related to one another by their geographic distance, is common in georeferenced ecological data. SAC in the residuals of SDMs violates the ‘independent errors’ assumption required to justify the use of statistical models in modelling species’ distributions. The autologistic modelling approach accounts for SAC by including an additional term (the autocovariate) representing the similarity between the value of the response variable at a location and neighbouring locations. However, autologistic models have been found to introduce bias in the estimation of parameters describing the influence of explanatory variables on habitat occupancy. To address this problem we developed an extension to the autologistic approach by calculating the autocovariate on SAC in residuals (the RAC approach). Performance of the new approach was tested on simulated data with a known spatial structure and on strongly autocorrelated mangrove species’ distribution data collected in northern Australia. The RAC approach was implemented as generalized linear models (GLMs) and boosted regression tree (BRT) models. We found that the BRT models with only environmental explanatory variables can account for some SAC, but applying the standard autologistic or RAC approaches further reduced SAC in model residuals and substantially improved model predictive performance. The RAC approach showed stronger inferential performance than the standard autologistic approach, as parameter estimates were more accurate and statistically significant variables were accurately identified. The new RAC approach presented here has the potential to account for spatial autocorrelation while maintaining strong predictive and inferential performance, and can be implemented across a range of modelling approaches.     

Avian SDMs: current state,challenges, and opportunities

Quantifying species distributions using species distribution models (SDMs) has emerged as a central method in modern biogeography. These empirical models link species occurrence data with spatial environmental information. Since their emergence in the 1990s, thousands of scientific papers have used SDMs to study organisms across the entire tree of life, with birds commanding considerable attention. Here, we review the current state of avian SDMs and point to challenges and future opportunities for specific applications, ranging from conservation biology, invasive species and predicting seabird distributions, to more general topics such as modeling avian diversity, niche evolution and seasonal distributions at a biogeographic scale. While SDMs have been criticized for being phenomenological in nature, and for their inability to explicitly account for a variety of processes affecting populations, we conclude that they remain a powerful tool to learn about past, current, and future species distributions – at least when their limitations and assumptions are recognized and addressed. We close our review by providing an outlook on prospects and synergies with other disciplines in which avian SDMs can play an important role.     

Species interactions weakly modify climate‐induced tree co‐occurrence patterns

Aims

Species distributions are hypothesized to be underlain by a complex association of processes that span multiple spatial scales including biotic interactions, dispersal limitation, fine‐scale resource gradients and climate. Species disequilibrium with climate may reflect the effects of non‐climatic processes on species distributions, yet distribution models have rarely directly considered non‐climatic processes. Here, we use a Joint Species Distribution Model (JSDM) to investigate the influence of non‐climatic factors on species co‐occurrence patterns and to directly quantify the relative influences of climate and alternative processes that may generate correlated responses in species distributions, such as species interactions, on tree co‐occurrence patterns.

Location

US Rocky Mountains.

Methods

We apply a Bayesian JSDM to simultaneously model the co‐occurrence patterns of ten dominant tree species across the Rocky Mountains, and evaluate climatic and residual correlations from the fitted model to determine the relative contribution of each component to observed co‐occurrence patterns. We also evaluate predictions generated from the fitted model relative to a single‐species modelling approach.

Results

For most species, correlation due to climate covariates exceeded residual correlation, indicating an overriding influence of broad‐scale climate on co‐occurrence patterns. Accounting for covariance among species did not significantly improve predictions relative to a single‐species approach, providing limited evidence for a strong independent influence of species interactions on distribution patterns.

Conclusions

Overall, our findings indicate that climate is an important driver of regional biodiversity patterns and that interactions between dominant tree species contribute little to explain species co‐occurrence patterns among Rocky Mountain trees.     

Benchmarking novel approaches for modelling species range dynamics

Increasing biodiversity loss due to climate change is one of the most vital challenges of the 21st century. To anticipate and mitigate biodiversity loss, models are needed that reliably project species’ range dynamics and extinction risks. Recently, several new approaches to model range dynamics have been developed to supplement correlative species distribution models (SDMs), but applications clearly lag behind model development. Indeed, no comparative analysis has been performed to evaluate their performance. Here, we build on process‐based, simulated data for benchmarking five range (dynamic) models of varying complexity including classical SDMs, SDMs coupled with simple dispersal or more complex population dynamic models (SDM hybrids), and a hierarchical Bayesian process‐based dynamic range model (DRM). We specifically test the effects of demographic and community processes on model predictive performance. Under current climate, DRMs performed best, although only marginally. Under climate change, predictive performance varied considerably, with no clear winners. Yet, all range dynamic models improved predictions under climate change substantially compared to purely correlative SDMs, and the population dynamic models also predicted reasonable extinction risks for most scenarios. When benchmarking data were simulated with more complex demographic and community processes, simple SDM hybrids including only dispersal often proved most reliable. Finally, we found that structural decisions during model building can have great impact on model accuracy, but prior system knowledge on important processes can reduce these uncertainties considerably. Our results reassure the clear merit in using dynamic approaches for modelling species’ response to climate change but also emphasize several needs for further model and data improvement. We propose and discuss perspectives for improving range projections through combination of multiple models and for making these approaches operational for large numbers of species.