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.

Many studies in recent years have investigated the effects of climate change on the future of biodiversity. In this review, we first examine the different possible effects of climate change that can operate at individual, population, species, community, ecosystem and biome scales, notably showing that species can respond to climate change challenges by shifting their climatic niche along three non-exclusive axes: time (e.g. phenology), space (e.g. range) and self (e.g. physiology). Then, we present the principal specificities and caveats of the most common approaches used to estimate future biodiversity at global and subcontinental scales and we synthesise their results. Finally, we highlight several challenges for future research both in theoretical and applied realms. Overall, our review shows that current estimates are very variable, depending on the method, taxonomic group, biodiversity loss metrics, spatial scales and time periods considered. Yet, the majority of models indicate alarming consequences for biodiversity, with the worstcase scenarios leading to extinction rates that would qualify as the sixth mass extinction in the history of the earth.

Aim  Concerns over how global change will influence species distributions, in conjunction with increased emphasis on understanding niche dynamics in evolutionary and community contexts, highlight the growing need for robust methods to quantify niche differences between or within taxa. We propose a statistical framework to describe and compare environmental niches from occurrence and spatial environmental data. Location  Europe, North America and South America. Methods  The framework applies kernel smoothers to densities of species occurrence in gridded environmental space to calculate metrics of niche overlap and test hypotheses regarding niche conservatism. We use this framework and simulated species with pre‐defined distributions and amounts of niche overlap to evaluate several ordination and species distribution modelling techniques for quantifying niche overlap. We illustrate the approach with data on two well‐studied invasive species. Results  We show that niche overlap can be accurately detected with the framework when variables driving the distributions are known. The method is robust to known and previously undocumented biases related to the dependence of species occurrences on the frequency of environmental conditions that occur across geographical space. The use of a kernel smoother makes the process of moving from geographical space to multivariate environmental space independent of both sampling effort and arbitrary choice of resolution in environmental space. However, the use of ordination and species distribution model techniques for selecting, combining and weighting variables on which niche overlap is calculated provide contrasting results. Main conclusions  The framework meets the increasing need for robust methods to quantify niche differences. It is appropriate for studying niche differences between species, subspecies or intra‐specific lineages that differ in their geographical distributions. Alternatively, it can be used to measure the degree to which the environmental niche of a species or intra‐specific lineage has changed over time.

BIOMOD is a computer platform for ensemble forecasting of species distributions, enabling the treatment of a range of methodological uncertainties in models and the examination of species-environment relationships. BIOMOD includes the ability to model species distributions with several techniques, test models with a wide range of approaches, project species distributions into different environmental conditions (e.g. climate or land use change scenarios) and dispersal functions. It allows assessing species temporal turnover, plot species response curves, and test the strength of species interactions with predictor variables. BIOMOD is implemented in R and is a freeware, open source, package.Species distribution models (SDM, Guisan and Thuiller 2005) are being used in nearly all branches of life and environmental sciences. A quick search in ISI Web of Science (18/02/08) using ''species distribution models'' OR ''niche models'' OR ''habitat models'' OR ''bioclimatic models'' highlights 21 973 papers, 74% of which published in the past 10 yr, in fields as varied as environmental sciences (53% of the records), zoology (15%), marine and freshwater biology (15%), life sciences and biomedicine (9%), biodiversity and conservation (8%), evolutionary biology (8%), fisheries (6%), forestry (6%), oceanography (5%), genetics and heredity (5%), amongst others. Advancement of knowledge in these fields is now intertwined with technical innovation in species distribution modelling and dependent on the existence of suitable software for fitting models and examining results. One difficulty with the use of species distribution models is that the number of techniques available is large and is increasing steadily, making it difficult for ''non-aficionados'' to select the most appropriate methodology for their needs (Elith et al. 2006, Heikkinen et al. 2006. Recent analyses have also demonstrated that discrepancies between different techniques can be very large, making the choice of the appropriate model even more difficult. This is particularly true when models are used to project distributions of species into independent situations, which is the example of projections of species distributions under future climate change scenarios (Thuiller 2004, Pearson et al. 2006. A possible solution to account for this inter-model variability is to fit ensembles of forecasts by simulating across more than one set of initial conditions, model classes, model parameters, and boundary conditions (for a review see Araújo and New 2007) and analyse the resulting range of uncertainties with bounding box, consensus and probabilistic methodologies rather than lining up with a single modelling outcome New 2007, Thuiller 2007). BIOMOD offers such a platform for ensemble forecasting (Fig. 1) using freeware and open-source R software (R Development Core Team 2008). It overcomes some of the limitations of existing software (e.g. being able to fit and compare different models) and incorporates several features for testing models (e.g. k-fold cross validation) ...

Climate change has already triggered species distribution shifts in many parts of the world. Increasing impacts are expected for the future, yet few studies have aimed for a general understanding of the regional basis for species vulnerability. We projected late 21st century distributions for 1,350 European plants species under seven climate change scenarios. Application of the International Union for Conservation of Nature and Natural Resources Red List criteria to our projections shows that many European plant species could become severely threatened. More than half of the species we studied could be vulnerable or threatened by 2080. Expected species loss and turnover per pixel proved to be highly variable across scenarios (27-42% and 45-63% respectively, averaged over Europe) and across regions (2.5-86% and 17-86%, averaged over scenarios). Modeled species loss and turnover were found to depend strongly on the degree of change in just two climate variables describing temperature and moisture conditions. Despite the coarse scale of the analysis, species from mountains could be seen to be disproportionably sensitive to climate change (Ϸ60% species loss). The boreal region was projected to lose few species, although gaining many others from immigration. The greatest changes are expected in the transition between the Mediterranean and Euro-Siberian regions. We found that risks of extinction for European plants may be large, even in moderate scenarios of climate change and despite inter-model variability.Intergovernmental Panel on Climate Change storylines ͉ species extinction ͉ species turnover ͉ niche-based model

Summary1. Species distribution models are increasingly used to address questions in conservation biology, ecology and evolution. The most effective species distribution models require data on both species presence and the available environmental conditions (known as background or pseudo-absence data) in the area. However, there is still no consensus on how and where to sample these pseudoabsences and how many. 2. In this study, we conducted a comprehensive comparative analysis based on simple simulated species distributions to propose guidelines on how, where and how many pseudo-absences should be generated to build reliable species distribution models. Depending on the quantity and quality of the initial presence data (unbiased vs. climatically or spatially biased), we assessed the relative effect of the method for selecting pseudo-absences (random vs. environmentally or spatially stratified) and their number on the predictive accuracy of seven common modelling techniques (regression, classification and machine-learning techniques). 3. When using regression techniques, the method used to select pseudo-absences had the greatest impact on the model's predictive accuracy. Randomly selected pseudo-absences yielded the most reliable distribution models. Models fitted with a large number of pseudo-absences but equally weighted to the presences (i.e. the weighted sum of presence equals the weighted sum of pseudoabsence) produced the most accurate predicted distributions. For classification and machine-learning techniques, the number of pseudo-absences had the greatest impact on model accuracy, and averaging several runs with fewer pseudo-absences than for regression techniques yielded the most predictive models. 4. Overall, we recommend the use of a large number (e.g. 10 000) of pseudo-absences with equal weighting for presences and absences when using regression techniques (e.g. generalised linear model and generalised additive model); averaging several runs (e.g. 10) with fewer pseudo-absences (e.g. 100) with equal weighting for presences and absences with multiple adaptive regression splines and discriminant analyses; and using the same number of pseudo-absences as available presences (averaging several runs if few pseudo-absences) for classification techniques such as boosted regression trees, classification trees and random forest. In addition, we recommend the random selection of pseudo-absences when using regression techniques and the random selection of geographically and environmentally stratified pseudo-absences when using classification and machine-learning techniques.

913 may also lead to dependence between species (phylogenetic structure) or populations of species (genetic structure) with more recent divergence will tend to be more similar than those which diverged longer ago (Harvey and Pagel 1991). While such underlying structures in the data are not fundamentally problematic for statistical analyses, they tend to create two undesirable outcomes. First, model error, as well as neglected processes and variables connected to these structures, often leads to dependence structures in the model residuals, which violates the critical assumption of independence present in many models and methods (Legendre and Fortin 1989, Miller et al. 2007). Second, because predictor variables are often correlated with underlying dependence structures (e.g. climate with space), models may use predic-tors to overfit the residual dependence structure and thereby remove it, partially or completely.

Increasing concern over the implications of climate change for biodiversity has led to the use of species-climate envelope models to project species extinction risk under climatechange scenarios. However, recent studies have demonstrated significant variability in model predictions and there remains a pressing need to validate models and to reduce uncertainties. Model validation is problematic as predictions are made for events that have not yet occurred. Resubstituition and data partitioning of present-day data sets are, therefore, commonly used to test the predictive performance of models. However, these approaches suffer from the problems of spatial and temporal autocorrelation in the calibration and validation sets. Using observed distribution shifts among 116 British breeding-bird species over the past $ 20 years, we are able to provide a first independent validation of four envelope modelling techniques under climate change. Results showed good to fair predictive performance on independent validation, although rules used to assess model performance are difficult to interpret in a decision-planning context. We also showed that measures of performance on nonindependent data provided optimistic estimates of models' predictive ability on independent data. Artificial neural networks and generalized additive models provided generally more accurate predictions of species range shifts than generalized linear models or classification tree analysis. Data for independent model validation and replication of this study are rare and we argue that perfect validation may not in fact be conceptually possible. We also note that usefulness of models is contingent on both the questions being asked and the techniques used. Implementations of species-climate envelope models for testing hypotheses and predicting future events may prove wrong, while being potentially useful if put into appropriate context.