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

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

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

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

Water–energy,land‐cover and heterogeneity drivers of the distribution of plant species richness in a mountain region of the European Alps

Aim To evaluate the relative importance of water–energy, land‐cover, environmental heterogeneity and spatial variables on the regional distribution of Red‐Listed and common vascular plant species richness. Location Trento Province (c. 6200 km²) on the southern border of the European Alps (Italy), subdivided regularly into 228 3′ × 5′ quadrants. Methods Data from a floristic inventory were separated into two subsets, representing Red‐Listed and common (i.e. all except Red‐Listed) plant species richness. Both subsets were separately related to water–energy, land‐cover and environmental heterogeneity variables. We simultaneously applied ordinary least squares regression with variation partitioning and hierarchical partitioning, attempting to identify the most important factors controlling species richness. We combined the analysis of environmental variables with a trend surface analysis and a spatial autocorrelation analysis. Results At the regional scale, plant species richness of both Red‐Listed and common species was primarily related to energy availability and land cover, whereas environmental heterogeneity had a lesser effect. The greatest number of species of both subsets was found in quadrants with the largest energy availability and the greatest degree of urbanization. These findings suggest that the elevation range within our study region imposes an energy‐driven control on the distribution of species richness, which resembles that of the broader latitude gradient. Overall, the two species subsets had similar trends concerning the relative importance of water–energy, land cover and environmental heterogeneity, showing a few differences regarding the selection of some predictors of secondary importance. The incorporation of spatial variables did not improve the explanatory power of the environmental models and the high original spatial autocorrelation in the response variables was reduced drastically by including the selected environmental variables. Main conclusions Water–energy and land cover showed significant pure effects in explaining plant species richness, indicating that climate and land cover should both be included as explanatory variables in modelling species richness in human‐affected landscapes. However, the high degree of shared variation between the two groups made the relative effects difficult to separate. The relatively low range of variation in the environmental heterogeneity variables within our sampling domain might have caused the low importance of this complex factor.     

Land use improves spatial predictions of mountain plant abundance but not presence‐absence

Question: Does a land‐use variable improve spatial predictions of plant species presence‐absence and abundance models at the regional scale in a mountain landscape? Location: Western Swiss Alps. Methods: Presence‐absence generalized linear models (GLM) and abundance ordinal logistic regression models (LRM) were fitted to data on 78 mountain plant species, with topo‐climatic and/or land‐use variables available at a 25‐m resolution. The additional contribution of land use when added to topo‐climatic models was evaluated by: (1) assessing the changes in model fit and (2) predictive power, (3) partitioning the deviance respectively explained by the topo‐climatic variables and the land‐use variable through variation partitioning, and (5) comparing spatial projections. Results: Land use significantly improved the fit of presence‐absence models but not their predictive power. In contrast, land use significantly improved both the fit and predictive power of abundance models. Variation partitioning also showed that the individual contribution of land use to the deviance explained by presence‐absence models was, on average, weak for both GLM and LRM (3.7% and 4.5%, respectively), but changes in spatial projections could nevertheless be important for some species. Conclusions: In this mountain area and at our regional scale, land use is important for predicting abundance, but not presence‐absence. The importance of adding land‐use information depends on the species considered. Even without a marked effect on model fit and predictive performance, adding land use can affect spatial projections of both presence‐absence and abundance models.     

Anticipated climate and land‐cover changes reveal refuge areas for Borneo's orang‐utans

Habitat loss and climate change pose a double jeopardy for many threatened taxa, making the identification of optimal habitat for the future a conservation priority. Using a case study of the endangered Bornean orang‐utan, we identify environmental refuges by integrating bioclimatic models with projected deforestation and oil‐palm agriculture suitability from the 1950s to 2080s. We coupled a maximum entropy algorithm with information on habitat needs to predict suitable habitat for the present day and 1950s. We then projected to the 2020s, 2050s and 2080s in models incorporating only land‐cover change, climate change or both processes combined. For future climate, we incorporated projections from four model and emission scenario combinations. For future land cover, we developed spatial deforestation predictions from 10 years of satellite data. Refuges were delineated as suitable forested habitats identified by all models that were also unsuitable for oil palm – a major threat to tropical biodiversity. Our analyses indicate that in 2010 up to 260 000 km² of Borneo was suitable habitat within the core orang‐utan range; an 18–24% reduction since the 1950s. Land‐cover models predicted further decline of 15–30% by the 2080s. Although habitat extent under future climate conditions varied among projections, there was majority consensus, particularly in north‐eastern and western regions. Across projections habitat loss due to climate change alone averaged 63% by 2080, but 74% when also considering land‐cover change. Refuge areas amounted to 2000–42 000 km² depending on thresholds used, with 900–17 000 km² outside the current species range. We demonstrate that efforts to halt deforestation could mediate some orang‐utan habitat loss, but further decline of the most suitable areas is to be expected given projected changes to climate. Protected refuge areas could therefore become increasingly important for ongoing translocation efforts. We present an approach to help identify such areas for highly threatened species given environmental changes expected this century.     

Climate and satellite-derived land cover for predicting breeding bird distribution in the Great Lakes Basin

Aim We examined relationships between breeding bird distribution of 10 forest songbirds in the Great Lakes Basin, large‐scale climate and the distribution of land cover types as estimated by advanced very high resolution radiometer (AVHRR) and multi‐spectral scanner (MSS) land cover classifications. Our objective was to examine the ability of regional climate, AVHRR (1 km resolution) land cover and MSS (200 m resolution) land cover to predict the distribution of breeding forest birds at the scale of the Great Lakes Basin and at the resolution of Breeding Bird Atlas data (5–10 km²). Specifically we addressed the following questions. (1) How well do AVHRR or MSS classifications capture the variation in distribution of bird species? (2) Is one land cover classification more useful than the other for predicting distribution? (3) How do models based on climate compare with models based on land cover? (4) Can the combination of both climate and land cover improve the predictive ability of these models. Location Modelling was conducted over the area of the Great Lakes Basin including parts of Ontario, Canada and parts of Illinois, Indiana, Michigan, New York, Ohio, Pennsylvania Wisconsin, and Minnesota, USA. Methods We conducted single variable logistic regression with the forest classes of AVHRR and MSS land cover using evidence of breeding as the response variable. We conducted multiple logistic regression with stepwise selection to select models from five sets of explanatory variables (AVHRR, MSS, climate, AVHRR + climate, MSS + climate). Results Generally, species were related to both AVHRR and MSS land cover types in the direction expected based on the known local habitat use of the species. Neither land cover classification appeared to produce consistently more intuitive results. Good models were generated using each of the explanatory data sets examined here. And at least one but usually all five variable sets produced acceptable or excellent models for each species. Main conclusions Both climate and large scale land cover were effective predictors of the distribution of the 10 forest bird species examined here. Models generated from these data had good classification accuracy of independent validation data. Good models were produced from all explanatory data sets or combinations suggesting that the distribution of climate, AVHRR land cover, and MSS land cover all captured similar variance in the distribution of the birds. It is difficult to separate the effects of climate and vegetation on the species’ distributions at this scale.     

Does local habitat fragmentation affect large‐scale distributions? The case of a specialist grassland bird

Aim

Although the negative effects of habitat fragmentation have been widely documented at the landscape scale, much less is known about its impacts on species distributions at the biogeographical scale. We hypothesize that fragmentation influences the large‐scale distribution of area‐ and edge‐sensitive species by limiting their occurrence in regions with fragmented habitats , despite otherwise favourable environmental conditions. We test this hypothesis by assessing the interplay of climate and landscape factors influencing the distribution of the calandra lark, a grassland specialist that is highly sensitive to habitat fragmentation.

Location

Iberia Peninsula, Europe.

Methods

Ecological niche modelling was used to investigate the relative influence of climate/topography, landscape fragmentation and spatial structure on calandra lark distribution. Modelling assumed explicitly a hierarchically structured effect among explanatory variables, with climate/topography operating at broader spatial scales than landscape variables. An eigenvector‐based spatial filtering approach was used to cancel bias introduced by spatial autocorrelation. The information theoretic approach was used in model selection, and variation partitioning was used to isolate the unique and shared effects of sets of explanatory variables.

Results

Climate and topography were the most influential variables shaping the distribution of calandra lark, but incorporating landscape metrics contributed significantly to model improvement. The probability of calandra lark occurrence increased with total habitat area and declined with the number of patches and edge density. Variation partitioning showed a strong overlap between variation explained by climate/topography and landscape variables. After accounting for spatial structure in species distribution, the explanatory power of environmental variables remained largely unchanged.

Main conclusions

We have shown here that landscape fragmentation can influence species distributions at the biogeographical scale. Incorporating fragmentation metrics into large‐scale ecological niche models may contribute for a better understanding of mechanism driving species distributions and for improving predictive modelling of range shifts associated with land use and climate changes.

     

Biotic interactions improve prediction of boreal bird distributions at macro-scales

Aim The role of biotic interactions in influencing species distributions at macro‐scales remains poorly understood. Here we test whether predictions of distributions for four boreal owl species at two macro‐scales (10 × 10 km and 40 × 40 km grid resolutions) are improved by incorporating interactions with woodpeckers into climate envelope models. Location Finland, northern Europe. Methods Distribution data for four owl and six woodpecker species, along with data for six land cover and three climatic variables, were collated from 2861 10 × 10 km grid cells. Generalized additive models were calibrated using a 50% random sample of the species data from western Finland, and by repeating this procedure 20 times for each of the four owl species. Models were fitted using three sets of explanatory variables: (1) climate only; (2) climate and land cover; and (3) climate, land cover and two woodpecker interaction variables. Models were evaluated using three approaches: (1) examination of explained deviance; (2) four‐fold cross‐validation using the model calibration data; and (3) comparison of predicted and observed values for independent grid cells in eastern Finland. The model accuracy for approaches (2) and (3) was measured using the area under the curve of a receiver operating characteristic plot. Results At 10‐km resolution, inclusion of the distribution of woodpeckers as a predictor variable significantly improved the explanatory power, cross‐validation statistics and the predictive accuracy of the models. Inclusion of land cover led to similar improvements at 10‐km resolution, although these improvements were less apparent at 40‐km resolution for both land cover and biotic interactions. Main conclusions Predictions of species distributions at macro‐scales may be significantly improved by incorporating biotic interactions and land cover variables into models. Our results are important for models used to predict the impacts of climate change, and emphasize the need for comprehensive evaluation of the reliability of species–climate impact models.     

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

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

Assessing uncertainties in land cover projections

Understanding uncertainties in land cover projections is critical to investigating land‐based climate mitigation policies, assessing the potential of climate adaptation strategies and quantifying the impacts of land cover change on the climate system. Here, we identify and quantify uncertainties in global and European land cover projections over a diverse range of model types and scenarios, extending the analysis beyond the agro‐economic models included in previous comparisons. The results from 75 simulations over 18 models are analysed and show a large range in land cover area projections, with the highest variability occurring in future cropland areas. We demonstrate systematic differences in land cover areas associated with the characteristics of the modelling approach, which is at least as great as the differences attributed to the scenario variations. The results lead us to conclude that a higher degree of uncertainty exists in land use projections than currently included in climate or earth system projections. To account for land use uncertainty, it is recommended to use a diverse set of models and approaches when assessing the potential impacts of land cover change on future climate. Additionally, further work is needed to better understand the assumptions driving land use model results and reveal the causes of uncertainty in more depth, to help reduce model uncertainty and improve the projections of land cover.     

宏生态尺度上景观破碎化对物种丰富度的影响

生物多样性的地理格局及其形成机制是宏生态学与生物地理学的研究热点。大量研究表明,景观尺度上的生境破碎化对物种多样性的分布格局具有重要作用,但目前尚不清楚这种作用是否足以在宏生态尺度上对生物多样性地理格局产生显著影响。利用中国大陆鸟类和哺乳动物的物种分布数据,在100 km×100 km网格的基础上生成了这两个类群生物的物种丰富度地理格局,进一步利用普通最小二乘法模型和空间自回归模型研究了物种丰富度与气候、生境异质性、景观破碎化的相关关系。结果表明,景观破碎化因子与鸟类和哺乳动物的物种丰富度都具有显著的关联关系,其方差贡献率可达约30%—50%(非空间模型)和60%—80%(空间模型),略低于或接近于气候和生境异质性因子。方差分解结果显示,景观破碎化因子与气候和生境异质性因子的方差贡献率的重叠部分达20%—40%。相对鸟类而言,景观破碎化对哺乳动物物种丰富度的地理格局具有更高的解释率。     

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

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

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

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

The contributions of topoclimate and land cover to species distributions and abundance: fine‐resolution tests for a mountain butterfly fauna

Aim Models relating species distributions to climate or habitat are widely used to predict the effects of global change on biodiversity. Most such approaches assume that climate governs coarse‐scale species ranges, whereas habitat limits fine‐scale distributions. We tested the influence of topoclimate and land cover on butterfly distributions and abundance in a mountain range, where climate may vary as markedly at a fine scale as land cover. Location Sierra de Guadarrama (Spain, southern Europe) Methods We sampled the butterfly fauna of 180 locations (89 in 2004, 91 in 2005) in a 10,800 km² region, and derived generalized linear models (GLMs) for species occurrence and abundance based on topoclimatic (elevation and insolation) or habitat (land cover, geology and hydrology) variables sampled at 100‐m resolution using GIS. Models for each year were tested against independent data from the alternate year, using the area under the receiver operating characteristic curve (AUC) (distribution) or Spearman's rank correlation coefficient (r_s) (abundance). Results In independent model tests, 74% of occurrence models achieved AUCs of > 0.7, and 85% of abundance models were significantly related to observed abundance. Topoclimatic models outperformed models based purely on land cover in 72% of occurrence models and 66% of abundance models. Including both types of variables often explained most variation in model calibration, but did not significantly improve model cross‐validation relative to topoclimatic models. Hierarchical partitioning analysis confirmed the overriding effect of topoclimatic factors on species distributions, with the exception of several species for which the importance of land cover was confirmed. Main conclusions Topoclimatic factors may dominate fine‐resolution species distributions in mountain ranges where climate conditions vary markedly over short distances and large areas of natural habitat remain. Climate change is likely to be a key driver of species distributions in such systems and could have important effects on biodiversity. However, continued habitat protection may be vital to facilitate range shifts in response to climate change.     

Patterns of beta diversity in Europe: the role of climate,land cover and distance across scales

Aim We test the prediction that beta diversity (species turnover) and the decay of community similarity with distance depend on spatial resolution (grain). We also study whether patterns of beta diversity are related to variability in climate, land cover or geographic distance and how the independent effects of these variables depend on the spatial grain of the data. Location Europe, Great Britain, Finland and Catalonia. Methods We used data on European birds, plants, butterflies, amphibians and reptiles, and data on British plants, Catalonian birds and Finnish butterflies. We fitted two or three nested grids of varying resolutions to each of these datasets. For each grid we calculated differences in climate, differences in land‐cover composition (CORINE) and beta diversity (β_sim, β_Jaccard) between all pairs of grid cells. In a separate analysis we looked specifically at pairs of adjacent grid cells (the first distance class). We then used variation partitioning to identify the magnitude of independent statistical associations (i.e. independent effects in the statistical sense) of climate, land cover and geographic distance with spatial patterns of beta diversity. Results Beta diversity between grid cells at any given distance decreased with increasing grain. Geographic distance was always the most important predictor of beta diversity for all pairwise comparisons at the extent of Europe. Climate and land cover had weaker but distinct and grain‐dependent effects. Climate was more important at relatively coarse grains, whereas land‐cover effects were stronger at finer grains. In the country‐wide analyses, climate and land cover were more important than geographic distance. Climatic and land‐cover models performed poorly and showed no systematic grain dependence for beta diversity between adjacent grid cells. Main conclusions We found that relationships between geographic distance and beta diversity, as well as the environmental correlates of beta diversity, are systematically grain dependent. The strong independent effect of distance indicates that, contrary to the current belief, a substantial fraction of species are missing from areas with a suitable environment. Moreover, the effects of geographic distance (at continental extents) and land cover (at fine grains) indicate that any species distribution modelling should take both environment and dispersal limitation into account.     

Can niche‐based distribution models outperform spatial interpolation?

Aim Distribution modelling relates sparse data on species occurrence or abundance to environmental information to predict the population of a species at any point in space. Recently, the importance of spatial autocorrelation in distributions has been recognized. Spatial autocorrelation can be categorized as exogenous (stemming from autocorrelation in the underlying variables) or endogenous (stemming from activities of the organism itself, such as dispersal). Typically, one asks whether spatial models explain additional variability (endogenous) in comparison to a fully specified habitat model. We turned this question around and asked: can habitat models explain additional variation when spatial structure is accounted for in a fully specified spatially explicit model? The aim was to find out to what degree habitat models may be inadvertently capturing spatial structure rather than true explanatory mechanisms. Location We used data from 190 species of the North American Breeding Bird Survey covering the conterminous United States and southern Canada. Methods We built 13 different models on 190 bird species using regression trees. Our habitat‐based models used climate and landcover variables as independent variables. We also used random variables and simulated ranges to validate our results. The two spatially explicit models included only geographical coordinates or a contagion term as independent variables. As another angle on the question of mechanism vs. spatial structure we pitted a model using related bird species as predictors against a model using randomly selected bird species. Results The spatially explicit models outperformed the traditional habitat models and the random predictor species outperformed the related predictor species. In addition, environmental variables produced a substantial R² in predicting artificial ranges. Main conclusions We conclude that many explanatory variables with suitable spatial structure can work well in species distribution models. The predictive power of environmental variables is not necessarily mechanistic, and spatial interpolation can outperform environmental explanatory variables.     

Modelling biome shifts and tree cover change for 2050 in West Africa

Aim Africa is expected to face severe changes in climatic conditions. Our objectives are: (1) to model trends and the extent of future biome shifts that may occur by 2050, (2) to model a trend in tree cover change, while accounting for human impact, and (3) to evaluate uncertainty in future climate projections. Location West Africa. Methods We modelled the potential future spatial distribution of desert, grassland, savanna, deciduous and evergreen forest in West Africa using six bioclimatic models. Future tree cover change was analysed with generalized additive models (GAMs). We used climate data from 17 general circulation models (GCMs) and included human population density and fire intensity to model tree cover. Consensus projections were derived via weighted averages to: (1) reduce inter‐model variability, and (2) describe trends extracted from different GCM projections. Results The strongest predicted effect of climate change was on desert and grasslands, where the bioclimatic envelope of grassland is projected to expand into the desert by an area of 2 million km². While savannas are predicted to contract in the south (by 54 ± 22 × 10⁴ km²), deciduous and evergreen forest biomes are expected to expand (64 ± 13 × 10⁴ km² and 77 ± 26 × 10⁴ km²). However, uncertainty due to different GCMs was particularly high for the grassland and the evergreen biome shift. Increasing tree cover (1–10%) was projected for large parts of Benin, Burkina Faso, Côte d’Ivoire, Ghana and Togo, but a decrease was projected for coastal areas (1–20%). Furthermore, human impact negatively affected tree cover and partly changed the direction of the projected change from increase to decrease. Main conclusions Considering climate change alone, the model results of potential vegetation (biomes) show a ‘greening’ trend by 2050. However, the modelled effects of human impact suggest future forest degradation. Thus, it is essential to consider both climate change and human impact in order to generate realistic future tree cover projections.     

Assessing ecosystem threats from global and regional change: hierarchical modeling of risk to sagebrush ecosystems from climate change, land use and invasive species in Nevada, USA

Global change poses significant challenges for ecosystem conservation. At regional scales, climate change may lead to extensive shifts in species distributions and widespread extirpations or extinctions. At landscape scales, land use and invasive species disrupt ecosystem function and reduce species richness. However, a lack of spatially explicit models of risk to ecosystems makes it difficult for science to inform conservation planning and land management. Here, I model risk to sagebrush ( Artemisia spp.) ecosystems in the state of Nevada, USA from climate change, land use/land cover change, and species invasion. Risk from climate change is based on an ensemble of 10 atmosphere-ocean general circulation model (AOGCM) projections applied to two bioclimatic envelope models (Mahalanobis distance and Maxent). Risk from land use is based on the distribution of roads, agriculture, and powerlines, and on the spatial relationships between land use and probability of cheatgrass Bromus tectorum invasion in Nevada. Risk from land cover change is based on probability and extent of pinyon-juniper ( Pinus monophylla; Juniperus spp.) woodland expansion. Climate change is most likely to negatively impact sagebrush ecosystems at the edges of its current range, particularly in southern Nevada, southern Utah, and eastern Washington. Risk from land use and woodland expansion is pervasive throughout Nevada, while cheatgrass invasion is most problematic in the northern part of the state. Cumulatively, these changes pose major challenges for conservation of sagebrush and sagebrush obligate species. This type of comprehensive assessment of ecosystem risk provides managers with spatially explicit tools important for conservation planning.     

Spatial heterogeneity of climate and land-cover constraints on distributions of tick-borne pathogens

Aim To compare the geographical distributions of two tick‐borne pathogens vectored by different tick species, to examine the relative importance of climate, land cover and host density in structuring these distributions, and to assess the spatial variability of these environmental constraints across the species ranges. Location South‐central and south‐eastern North America. Methods Presence/absence data for two tick‐borne pathogens, Ehrlichia chaffeensis and Anaplasma phagocytophilum, were obtained for 567 counties from a regional data base based on white‐tailed deer (Odocoileus virginianus) serology. Environmental variables describing climate, land cover and deer density were calculated for these counties. Global logistic regression analysis was used to screen the environmental variables and select a parsimonious subset of predictors. Local analysis was carried out using geographically weighted regression (GWR) to explore spatial variability in the parameters of the regression models. Cluster analysis was applied to the GWR output to identify zones with distinctive species–habitat relationships. Results Global habitat models for E. chaffeensis and A. phagocytophilum included temperature, humidity, precipitation and forest cover as explanatory variables. The E. chaffeensis model also included forest fragmentation, whereas the A. phagocytophilum model included deer density. Local analyses revealed that climate was the primary correlate of pathogen presence in the eastern portion of the study area, whereas forest cover and fragmentation constrained the western range boundaries. Habitat relationships for all variables were weak in and around the Mississippi Delta. Main conclusions Efforts to model pathogen and disease ranges, and to predict shifts in response to global change should consider future scenarios of land‐cover change as well as climate change, and should address the possibility of spatial heterogeneity in species–habitat relationships. The methods presented here outline an approach for objectively delineating geographical zones with similar species–environment relationships, which can then be used to stratify landscapes for the purposes of further explanatory and predictive modelling.     

Factors controlling the spatial species richness pattern of four groups of terrestrial vertebrates in an area between two different biogeographic regions in northern Spain

Aim To examine the influence of environmental variables on species richness patterns of amphibians, reptiles, mammals and birds and to assess the general usefulness of regional atlases of fauna. Location Navarra (10,421 km²) is located in the north of the Iberian Peninsula, in a territory shared by Mediterranean and Eurosiberian biogeographic regions. Important ecological patterns, climate, topography and land‐cover vary significantly from north to south. Methods Maps of vertebrate distribution and climatological and environmental data bases were used in a geographic information systems framework. Generalized additive models and partial regression analysis were used as statistical tools to differentiate (A) the purely spatial fraction, (B) the spatially structured environmental fraction and (C) the purely environmental fraction. In this way, we can evaluate the explanatory capacity of each variable, avoiding false correlations and assessing true causality. Final models were obtained through a stepwise procedure. Results Energy‐related features of climate, aridity and land‐cover variables show significant correlation with the species richness of reptiles, mammals and birds. Mammals and birds exhibit a spatial pattern correlated with variables such as aridity index and vegetation land‐cover. However, the high values of the spatially structured environmental fraction B and the low values of the purely environmental fraction A suggest that these predictor variables have a limited causal relationship with species richness for these vertebrate groups. An increment in land‐cover diversity is correlated with an increment of specific richness in reptiles, mammals and birds. No variables were found to be statistically correlated with amphibian species richness. Main conclusions Although aridity and land‐cover are the best predictor variables, their causal relationship with species richness must be considered with caution. Historical factors exhibiting a similar spatial pattern may be considered equally important in explaining the patterns of species richness. Also, land‐cover diversity appears as an important factor for maintaining biological diversity. Partial regression analysis has proved a useful technique in dealing with spatial autocorrelation. These results highlight the usefulness of coarsely sampled data and cartography at regional scales to predict and explain species richness patterns for mammals and birds. The accuracy of models appears to be related to the range perception of each group and the scale of the information.