The relative roles of environment and history as controls of tree species composition and richness in Europe
Authors: Svenning Jens‐Christian , Skov Flemming 
Published: May 2005

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.

Results The main floristic pattern identified by all ordinations was a latitude‐temperature 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 spatial‐environmental 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 trees, with the exception of conifer species richness. Strong support for interpreting the spatial patterns as outcomes of historical processes, notably dispersal limitation, came from the observation that many European tree species naturalize extensively outside their native ranges. Furthermore, it was confirmed that climate predominates among environmental determinants of distribution and diversity patterns at large spatial scales. Finally, the particular patterns exhibited by conifers probably reflect greater environmental specialization and greater human impact. These findings warn against expecting the European tree flora to be able track fast future climate changes on its own.