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.     

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.     

Land-cover data improve bioclimatic models for anurans and turtles at a regional scale

Aim We investigated whether accounting for land cover could improve bioclimatic models for eight species of anurans and three species of turtles at a regional scale. We then tested whether accounting for spatial autocorrelation could significantly improve bioclimatic models after statistically controlling for the effects of land cover. Location Nova Scotia, eastern Canada. Methods Species distribution data were taken from a recent (1999–2003) herpetofaunal atlas. Generalized linear models were used to relate the presence or absence of each species to climate and land‐cover variables at a 10‐km resolution. We then accounted for spatial autocorrelation using an autocovariate or third‐order trend surface of the geographical coordinates of each grid square. Finally, variance partitioning was used to explore the independent and joint contributions of climate, land cover and spatial autocorrelation. Results The inclusion of land cover significantly increased the explanatory power of bioclimatic models for 10 of the 11 species. Furthermore, including land cover significantly increased predictive performance for eight of the 11 species. Accounting for spatial autocorrelation improved model fit for rare species but generally did not improve prediction success. Variance partitioning demonstrated that this lack of improvement was a result of the high correlation between climate and trend‐surface variables. Main conclusions The results of this study suggest that accounting for the effects of land cover can significantly improve the explanatory and predictive power of bioclimatic models for anurans and turtles at a regional scale. We argue that the integration of climate and land‐cover data is likely to produce more accurate spatial predictions of contemporary herpetofaunal diversity. However, the use of land‐cover simulations in climate‐induced range‐shift projections introduces additional uncertainty into the predictions of bioclimatic models. Further research is therefore needed to determine whether accounting for the effects of land cover in range‐shift projections is merited.     

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.     

Prediction of butterfly diversity hotspots in Belgium: a comparison of statistically focused and land use-focused models

Aim We evaluate differences between and the applicability of three linear predictive models to determine butterfly hotspots in Belgium for nature conservation purposes. Location The study is carried out in Belgium for records located to Universal Transverse Mercator (UTM) grid cells of 5 × 5 km. Methods We first determine the relationship between factors correlated to butterfly diversity by means of modified t‐tests and principal components analysis; subsequently, we predict hotspots using linear models based on land use, climate and topographical variables of well‐surveyed UTM grid cells (n = 197). The well‐surveyed squares are divided into a training set and an evaluation set to test the model predictions. We apply three different models: (1) a ‘statistically focused’ model where variables are entered in descending order of statistical significance, (2) a ‘land use‐focused’ model where land use variables known to be related to butterfly diversity are forced into the model and (3) a ‘hybrid’ model where the variables of the ‘land use‐focused model’ are entered first and subsequently complemented by the remaining variables entered in descending order of statistical significance. Results A principal components analyses reveals that climate, and to a large extent, land use are locked into topography, and that topography and climate are the variables most strongly correlated with butterfly diversity in Belgium. In the statistically focused model, biogeographical region alone explains 65% of the variability; other variables entering the statistically focused model are the area of coniferous and deciduous woodland, elevation and the number of frost days; the statistically focused model explains 77% of the variability in the training set and 66% in the evaluation set. In the land use‐focused model, biogeographical region, deciduous and mixed woodland, natural grassland, heathland and bog, woodland edge, urban and agricultural area and biotope diversity are forced into the model; the land use‐focused model explains 68% of the variability in the training set and 57% in the evaluation set. In the hybrid model, all variables from the land use‐focused model are entered first and the covariates elevation, number of frost days and natural grassland area are added on statistical grounds; the hybrid model explains 78% of the variability in the training set and 67% in the evaluation set. Applying the different models to determine butterfly diversity hotspots resulted in the delimitation of spatially different areas. Main conclusions The best predictions of butterfly diversity in Belgium are obtained by the hybrid model in which land use variables relevant to butterfly richness are entered first after which climatic and topographic variables were added on strictly statistical grounds. The land use‐focused model does not predict butterfly diversity in a satisfactory manner. When using predictive models to determine butterfly diversity, conservation biologists need to be aware of the consequences of applying such models. Although, in conservation biology, land use‐focused models are preferable to statistically focused models, one should always check whether the applied model makes sense on the ground. Predictive models can target mapping efforts towards potentially species‐rich sites and permits the incorporation of un‐surveyed sites into nature conservancy policies. Species richness distribution maps produced by predictive modelling should therefore be used as pro‐active conservation tools.     

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.     

Precipitation of the warmest quarter and temperature of the warmest month are key to understanding the effect of climate change on plant species diversity in Southern African savannah

In this study, we test for the key bioclimatic variables that significantly explain the current distribution of plant species richness in a southern African ecosystem as a preamble to predicting plant species richness under a changed climate. We used 54,000 records of georeferenced plant species data to calculate species richness and spatially interpolated climate data to derive nineteen bioclimatic variables. Next, we determined the key bioclimatic variables explaining variation in species richness across Zimbabwe using regression analysis. Our results show that two bioclimatic variables, that is, precipitation of the warmest quarter (R² = 0.92, P < 0.001) and temperature of the warmest month (R² = 0.67, P < 0.001) significantly explain variation in plant species richness. In addition, results of bioclimatic modelling using future climate change projections show a reduction in the current bio‐climatically suitable area that supports high plant species richness. However, in high‐altitude areas, plant richness is less sensitive to climate change while low‐altitude areas show high sensitivity. Our results have important implications to biodiversity conservation in areas sensitive to climate change; for example, high‐altitude areas are likely to continue being biodiversity hotspots, as such future conservation efforts should be concentrated in these areas.     

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.     

Modelling the distribution of a threatened habitat: the California sage scrub

Aim Using predictive species distribution and ecological niche modelling our objectives are: (1) to identify important climatic drivers of distribution at regional scales of a locally complex and dynamic system – California sage scrub; (2) to map suitable sage scrub habitat in California; and (3) to distinguish between bioclimatic niches of floristic groups within sage scrub to assess the conservation significance of analysing such species groups. Location Coastal mediterranean‐type shrublands of southern and central California. Methods Using point localities from georeferenced herbarium records, we modelled the potential distribution and bioclimatic envelopes of 14 characteristic sage scrub species and three floristic groups (south‐coastal, coastal–interior disjunct and broadly distributed species) based upon current climate conditions. Maxent was used to map climatically suitable habitat, while principal components analysis followed by canonical discriminant analysis were used to distinguish between floristic groups and visualize species and group distributions in multivariate ecological space. Results Geographical distribution patterns of individual species were mirrored in the habitat suitability maps of floristic groups, notably the disjunct distribution of the coastal–interior species. Overlap in the distributions of floristic groups was evident in both geographical and multivariate niche space; however, discriminant analysis confirmed the separability of floristic groups based on bioclimatic variables. Higher performance of floristic group models compared with sage scrub as a whole suggests that groups have differing climate requirements for habitat suitability at regional scales and that breaking sage scrub into floristic groups improves the discrimination between climatically suitable and unsuitable habitat. Main conclusions The finding that presence‐only data and climatic variables can produce useful information on habitat suitability of California sage scrub species and floristic groups at a regional scale has important implications for ongoing efforts of habitat restoration for sage scrub. In addition, modelling at a group level provides important information about the differences in climatic niches within California sage scrub. Finally, the high performance of our floristic group models highlights the potential a community‐level modelling approach holds for investigating plant distribution patterns.     

Potential spatial interaction of the invasive species Harmonia axyridis (Pallas) with native and endemic coccinellids

Biological invasions represent a serious menace to local species assemblages, mainly due to interspecific relationships such as competition and predation. One important invasive species worldwide is Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), which has invaded many regions of the world, threatening the native and endemic coccinellid assemblages due to negative interspecific interactions. These interactions have been widely studied at a local scale, but have been less studied at regional scales. Our aim was to estimate and analyse the potential spatial interaction associated with the co‐occurrence of H. axyridis with native and endemic species in Chile, considering bioclimatic and land cover variables. First, we created species distribution models (SDM) for H. axyridis, native and endemic coccinellids and six representative coccinellid species using maximum entropy technique. Then, we overlapped each SDM with land cover types to estimate the bioclimatic suitability within each land cover type. Finally, we identified the co‐occurrences of organisms according to the SDM and the land cover types, estimating in what land covers H. axyridis and the other coccinellids are more likely to co‐occur. Our results show that the suitable area for H. axyridis occurs from 30° to 42°S in Chile, while for native and endemic species this area is greater. The six selected species are mainly concentrated in central Chile, but differ in their potential suitable areas; Adalia angulifera Mulsant and Scymnus bicolor (Germain) have the largest range, and Mimoscymnus macula (Germain) has the most restricted one. The highest level of potential spatial interactions with H. axyridis occurs in central Chile, specifically in croplands and scrublands, and the lowest in primary native forest for all the species. Our results provide a spatially explicit baseline for coccinellid conservation and management of this invasive species.     

Fine‐scale determinants of butterfly species richness and composition in a mountain region

Aim Global patterns of species richness are often considered to depend primarily on climate. We aimed to determine how topography and land cover affect species richness and composition at finer scales. Location Sierra de Guadarrama (central Iberian Peninsula). Methods We sampled the butterfly fauna of 180 locations (89 in 2004, 91 in 2005) at 600–2300 m elevation in a region of 10800 km². We recorded environmental variables at 100‐m resolution using GIS, and derived generalized linear models for species density (number of species per unit area) and expected richness (number of species standardized to number of individuals) based on variables of topoclimate (elevation and insolation) or land cover (vegetation type, geology and hydrology), or both (combined). We evaluated the models against independent data from the alternative study year. We also tested for differences in species composition among sites and years using constrained ordination (canonical correspondence analysis), and used variation partitioning analyses to quantify the independent and combined roles of topoclimate and land cover. Results Topoclimatic, land cover and combined models were significantly related to observed species density and expected richness. Topoclimatic and combined models outperformed models based on land cover variables, showing a humped elevational diversity gradient. Both topoclimate and land cover made significant contributions to models of species composition. Main conclusions Topoclimatic factors may dominate species richness patterns in regions with pronounced elevational gradients, as long as large areas of natural habitat remain. In contrast, both topoclimate and land cover may have important effects on species composition. Biodiversity conservation in mountainous regions therefore requires protection and management of natural habitats over a wide range of topoclimatic conditions, which may assist in facilitating range shifts and alleviating declines in species richness related to climate change.     

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.     

Species traits are associated with the quality of bioclimatic models

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

Geographical distribution modelling of the bronze bug: a worldwide invasion

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Disregarding topographical heterogeneity biases species turnover assessments based on bioclimatic models

We investigated whether the inclusion of topographical heterogeneity in bioclimatic envelope models would significantly alter predictions of climate change – induced broad-scale butterfly species range size changes in Europe. Using generalized additive models, and data on current climate and species distributions and two different climate scenarios (HadCM3A2 and HadCM3B2) for the period 2051–2080, we developed predictions of the currently suitable area and potential range size changes of 100 European butterfly species. The inclusion of elevation range increased the predictive accuracy of climate-only models for 86 of the 100 species. The differences in projected future distributions were most notable in mountainous areas, where the climate–topography models projected only ca. half of the species losses than the climate-only models. By contrast, climate–topography models estimated double the losses of species than climate-only models in the flatlands regions. Our findings suggest that disregarding topographical heterogeneity may cause a significant source of error in broad-scale bioclimatic modelling. Mountainous regions are likely to be even more important for future conservation of species than had until now been predicted, based on bioclimatic envelope models that did not take an explicit account of elevational range of grid squares.     

Local vs regional factors as determinants of the invasibility of indigenous forest fragments by alien plant species

Both local and regional filters can determine the invasion of alien species into native plant communities. However, their relative importance is essentially unknown. We used plot data from fragments of indigenous forests in southeastern New Zealand to infer which factors are important in explaining invasibility, measured as alien species richness. Twenty-eight predictor variables comprising both local factors (stand structure and soil) and regional ones (climate and land cover) were assessed. Reduction or increase in deviance in linear models was assessed, both individually and with a forward and backward stepwise variable selection procedure using the Akaike information criterion (AIC).
We found that higher alien species richness was mainly associated with forest fragments of small area in warm and dry climates and where there were only small areas of surrounding indigenous forest. Local soil and stand structure variables had considerably smaller effects on alien species richness than the regional land cover and climate variables. Alien species richness showed no relationship with native species richness. We conclude that in the forest fragments investigated here, of the variables included in the analyses, regional land cover and climate variables are potentially important drivers for alien species richness at plot level. This has implications for projections of alien species spread in the future under different climate change and land use scenarios.     

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.     

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

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

Partitioning global change: Assessing the relative importance of changes in climate and land cover for changes in avian distribution

Understanding the relative impact of climate change and land cover change on changes in avian distribution has implications for the future course of avian distributions and appropriate management strategies. Due to the dynamic nature of climate change, our goal was to investigate the processes that shape species distributions, rather than the current distributional patterns. To this end, we analyzed changes in the distribution of Eastern Wood Pewees (Contopus virens) and Red‐eyed Vireos (Vireo olivaceus) from 1997 to 2012 using Breeding Bird Survey data and dynamic correlated‐detection occupancy models. We estimated the local colonization and extinction rates of these species in relation to changes in climate (hours of extreme temperature) and changes in land cover (amount of nesting habitat). We fit six nested models to partition the deviance explained by spatial and temporal components of land cover and climate. We isolated the temporal components of environmental variables because this is the essence of global change. For both species, model fit was significantly improved when we modeled vital rates as a function of spatial variation in climate and land cover. Model fit improved only marginally when we added temporal variation in climate and land cover to the model. Temporal variation in climate explained more deviance than temporal variation in land cover, although both combined only explained 20% (Eastern Wood Pewee) and 6% (Red‐eyed Vireo) of temporal variation in vital rates. Our results showing a significant correlation between initial occupancy and environmental covariates are consistent with biological expectation and previous studies. The weak correlation between vital rates and temporal changes in covariates indicated that we have yet to identify the most relevant components of global change influencing the distributions of these species and, more importantly, that spatially significant covariates are not necessarily driving temporal shifts in avian distributions.     

Integrating anthropogenic factors into regional‐scale species distribution models—A novel application in the imperiled sagebrush biome

Species distribution models (SDMs) that rely on regional‐scale environmental variables will play a key role in forecasting species occurrence in the face of climate change. However, in the Anthropocene, a number of local‐scale anthropogenic variables, including wildfire history, land‐use change, invasive species, and ecological restoration practices can override regional‐scale variables to drive patterns of species distribution. Incorporating these human‐induced factors into SDMs remains a major research challenge, in part because spatial variability in these factors occurs at fine scales, rendering prediction over regional extents problematic. Here, we used big sagebrush (Artemisia tridentata Nutt.) as a model species to explore whether including human‐induced factors improves the fit of the SDM. We applied a Bayesian hurdle spatial approach using 21,753 data points of field‐sampled vegetation obtained from the LANDFIRE program to model sagebrush occurrence and cover by incorporating fire history metrics and restoration treatments from 1980 to 2015 throughout the Great Basin of North America. Models including fire attributes and restoration treatments performed better than those including only climate and topographic variables. Number of fires and fire occurrence had the strongest relative effects on big sagebrush occurrence and cover, respectively. The models predicted that the probability of big sagebrush occurrence decreases by 1.2% (95% CI: ?6.9%, 0.6%) when one fire occurs and cover decreases by 44.7% (95% CI: ?47.9%, ?41.3%) if at least one fire occurred over the 36 year period of record. Restoration practices increased the probability of big sagebrush occurrence but had minimal effect on cover. Our results demonstrate the potential value of including disturbance and land management along with climate in models to predict species distributions. As an increasing number of datasets representing land‐use history become available, we anticipate that our modeling framework will have broad relevance across a range of biomes and species.