The post-glacial migration of European beech Fagus sylvatica has been addressed by many studies using either genetic or fossil data or a combination of both. In contrast to this, only little is known about the migration history of beech forest understorey species. In a review of phytosociological literature, we identified 110 plant species which are closely associated with beech forest. We divided the distribution range of European beech forests into 40 geographical regions, and the presence or absence of each species was recorded for each region. We compared overall species numbers per region and numbers of narrow-range species (species present in <10 regions). A multiple regression model was used to test for the explanatory value of three potential diversity controls: range in elevation, soil type diversity, and distance to the nearest potential refuge area. A hierarchical cluster analysis of the narrow-range species was performed.
The frequency of range sizes shows a U-shaped distribution, with 42 species occurring in <10 regions. The highest number of beech forest species is found in the southern Alps and adjacent regions, and species numbers decrease with increasing distance from these regions. With only narrow-range species taken into consideration, secondary maxima are found in Spain, the southern Apennines, the Carpathians, and Greece. Distance to the nearest potential refuge area is the strongest predictor of beech forest species richness, while altitudinal range and soil type diversity had little or no predictive value. The clusters of narrow-range species are in good concordance with the glacial refuge areas of beech and other temperate tree species as estimated in recent studies. These findings support the hypothesis that the distribution of many beech forest species is limited by post-glacial dispersal rather than by their environmental requirements.