The relative roles of environment and history in controlling large-scale species distributions are important not only theoretically, but also for forecasting range responses to climatic change. Here, we use atlas data to examine the extent to which 55 tree species fill their climatically determined potential ranges in Europe. Quantifying range filling (R/P) as realized/potential range size ratios using bioclimatic envelope modelling we find mean R/P ¼ 38.3% (±30.3% SD). Many European tree species naturalize extensively outside their native ranges, providing support for interpreting the many low R/Ps as primarily reflecting dispersal limitation. R/P increases strongly with latitudinal range centroid and secondarily with hardiness and decreases weakly with longitudinal range centroid. Hence, European tree species ranges appear strongly controlled by geographical dispersal constraints on post-glacial expansion as well as climate. Consequently, we expect European tree species to show only limited tracking of near-future climate changes.

The relative importance of contemporary climate and history as controls of geographical diversity patterns is intensely debated. A key example is the controversy over the extent to which temperate tree distributions and diversity patterns reflect postglacial dispersal limitation. Here, we focus on Central and Northern Europe, and show that recent estimates of tree migration rates < 100 m year(-1) imply that many species have probably not reached equilibrium with climate in this region. We then demonstrate that geographical accessibility from glacial refuges explains 78% of the geographical variation in the region's tree diversity and is a much stronger diversity predictor than climate. Finally, we show that realistic estimates of migration rates can be derived from the observed tree diversity pattern by assuming it to be purely dispersal driven. In conclusion, the tree diversity pattern in Central and Northern Europe could, to a large extent, be a result of postglacial dispersal limitation.

Aim This study uses a high-resolution simulation of the Last Glacial Maximum (LGM) climate to assess: (1) whether LGM climate still affects the geographical species richness patterns in the European tree flora and (2) the relative importance of modern and LGM climate as controls of tree species richness in Europe. LocationThe parts of Europe that were unglaciated during the LGM.Methods Atlas data on the distributions of 55 tree species were linked with data on modern and LGM climate and climatic heterogeneity in a geographical information system with a 60-km grid. Four measures of species richness were computed: total richness, and richness of the 18 most restricted species, 19 species of medium incidence (intermediate species) and 18 most widespread species. We used ordinary least-squares regression and spatial autoregressive modelling to test and estimate the richness-climate relationships. ResultsLGM climate constituted the best single set of explanatory variables for richness of restricted species, while modern climate and climatic heterogeneity was best for total and widespread species richness and richness of intermediate species, respectively. The autoregressive model with all climatic predictors was supported for all richness measures using an information-theoretic approach, albeit only weakly so for total species richness. Among the strongest relationships were increases in total and intermediate richness with climatic heterogeneity and in restricted richness with LGM growing-degree-days. Partial regression showed that climatic heterogeneity accounted for the largest unique variation fraction for intermediate richness, while LGM climate was particularly important for restricted richness. Main conclusionsLGM climate appears to still affect geographical patterns of tree species richness in Europe, albeit the relative importance of modern and LGM climate depends on range size. Notably, LGM climate is a strong richness control for species with a restricted range, which appear to still be associated with their glacial refugia.

The influence of dispersal limitation on species ranges remains controversial. Considering the dramatic impacts of the last glaciation in Europe, species might not have tracked climate changes through time and, as a consequence, their present-day ranges might be in disequilibrium with current climate. For 1016 European plant species, we assessed the relative importance of current climate and limited postglacial migration in determining species ranges using regression modelling and explanatory variables representing climate, and a novel species-specific hind-casting-based measure of accessibility to postglacial colonization. Climate was important for all species, while postglacial colonization also constrained the ranges of more than 50 per cent of the species. On average, climate explained five times more variation in species ranges than accessibility, but accessibility was the strongest determinant for one-sixth of the species. Accessibility was particularly important for species with limited long-distance dispersal ability, with southern glacial ranges, seed plants compared with ferns, and small-range species in southern Europe. In addition, accessibility explained one-third of the variation in species' disequilibrium with climate as measured by the realized/potential range size ratio computed with niche modelling. In conclusion, we show that although climate is the dominant broad-scale determinant of European plant species ranges, constrained dispersal plays an important supplementary role.

Climate is often singled out as the primary range limiting factor at large scales, while other environmental factors, notably soil, are thought to predominate at smaller scales. However, the postglacial migrational lag hypothesis controversially suggests that many species are strongly dispersal-limited and still expanding from their ice age refugia. We investigated the importance of postglacial migrational lag, climate, and soil as range determinants for 47 widespread forest plant species across nemoral Europe (47.0Á60.08N, west of 24.08E) using regression modeling, information-theoretic model selection, multi-model inference, and variation partitioning. Migrational lag was represented by a measure of accessibility to recolonization from ice age refugia.Twelve species were largely ubiquitous and not analyzed further. For the remaining species, there was strong support for climate, soil, and accessibility to postglacial recolonization. Accessibility accounted for a small to moderate amount of variation, but its model-averaged regression coefficient was stronger than those for climate or soil for 11 species and the second or third strongest coefficient for an additional 10 species. The resulting odds ratios were greater than one for 33 out of 35 species, i.e. adjusted for climate and soil, prevalence for the far majority of the 35 non-ubiquitous species increased with increasing accessibility. There were no differences among growth forms in the importance of accessibility. In contrast, compared to non-boreal species, accessibility had little importance for species with wide boreal distributions, as expected from their more widespread, northern glacial distributions.In conclusion, even the ranges of many widespread forest plant species are probably still moderately to strongly limited by postglacial migrational lag. Therefore, although species ranges are also strongly influenced by climate, we cannot expect most forest plant species to closely track the expected 21st century climatic changes.

The aim of this study is to evaluate the possible consequences of climate change on a representative sample of forest herbs in Europe. A fuzzy climatic envelope was used to predict the location of suitable climatic conditions under two climatic change scenarios. Expected consequences in terms of lost and gained range size and shift in distribution for 26 forest herbs were estimated. These results were combined in an Index of Predicted Range Change for each species. Finally, the effects of habitat fragmentation for potential dispersal routes were evaluated and options for management on a European scale are discussed.Generally, a good agreement of the estimated suitability under the present climate and the observed current distribution was observed. However, species vary a lot in the degree to which they occupy the presently climatically suitable areas in Europe. Many species are absent from large areas with suitable climate and thus could be said to have poor range-filling capacity.A general change in location (range centroid) of the total suitable area was observed: The total suitable area will on average move strongly northwards and moderately eastwards under the relatively mild B1 scenario and more strongly so under the A2 scenario. The required average minimum migration rate per year to track the potential range shift is 2.1 km under the B1 scenario and 3.9 km under the A2 scenario.Moderate losses in the total suitable area in Europe are predicted for most species under both scenarios. However, the predicted changes are very variable, with one species (Actaea erythrocarpa ) experiencing total range elimination in Europe (A2 scenario) while the total suitable area is predicted to show large increases for other species. The species that are predicted to experience the greatest proportional losses in their climatically suitable area within their presently realised range tend to have northern or eastern range centroids.

The Intergovernmental Panel on Climate Change (IPCC) predicts an increase in global temperatures of between 1.48C and 5.88C during the 21st century, as a result of elevated CO 2 levels. Using bioclimatic envelope models, we evaluate the potential impact of climate change on the distributions and species richness of 120 native terrestrial nonvolant European mammals under two of IPCC's future climatic scenarios. Assuming unlimited and no migration, respectively, our model predicts that 1% or 5-9% of European mammals risk extinction, while 32-46% or 70-78% may be severely threatened (lose > 30% of their current distribution) under the two scenarios. Under the no migration assumption endemic species were predicted to be strongly negatively affected by future climatic changes, while widely distributed species would be more mildly affected. Finally, potential mammalian species richness is predicted to become dramatically reduced in the Mediterranean region but increase towards the northeast and for higher elevations. Bioclimatic envelope models do not account for non-climatic factors such as land-use, biotic interactions, human interference, dispersal or history, and our results should therefore be seen as first approximations of the potential magnitude of future climatic changes.

Aim This study investigates the determinants of European-scale patterns in tree species composition and richness, addressing the following questions: (1) What is the relative importance of environment and history? History refers to lasting effects of past large-scale events and time-dependent cumulative effects of ongoing processes, notably dispersal limited range dynamics. (2) Among the environmental determinants, what is the relative importance of climate, soils, and forest cover? (3) Do the answers to questions 1 and 2 differ between conifers and Fagales, the two major monophyletic groups of European trees?Location The study area comprises most of Europe (34°N-72°N and 11°W-32°E).Methods Atlas data on native distributions of 54 large tree species at 50 · 50 km resolution were linked with climatic, edaphic, and forest cover maps in a geographical information system. Unconstrained (principal components analysis using Hellinger distance transformation and detrended correspondence analysis) and constrained ordinations (redundancy analysis using Hellinger distance transformation and canonical correspondence analysis) and multiple linear regressions were used to investigate the determinants of species composition and species richness, respectively. History is expected to leave its mark as broad spatial patterns and was represented by the nine spatial terms of a cubic trend surface polynomial. ResultsThe main floristic pattern identified by all ordinations was a latitudetemperature gradient, while the lower axes corresponded mostly to spatial variables. Partitioning the floristic variation using constrained ordinations showed the mixed spatial-environmental and pure spatial fractions to be much greater than the pure environmental fraction. Biplots, forward variable selection, and partial analyses all suggested climatic variables as more important floristic determinants than forest cover or soil variables. Tree species richness peaked in the mountainous regions of East-Central and Southern Europe, except the Far West. Variation partitioning of species richness found the mixed spatialenvironmental and pure spatial fractions to be much greater than the pure environmental fraction for all species combined and Fagales, but not for conifers. The scaled regression coefficients indicated climate as a stronger determinant of richness than soils or forest cover. While the dominant patterns were similar for conifers and Fagales, conifers exhibited less predictable patterns overall, a smaller pure spatial variation fraction relative to pure environmental fraction, and a greater relative importance of climate; all differences being more pronounced for species richness than for species composition.Main conclusions The analyses suggest that history is at least as important as current environment in controlling species composition and richness of European