Habitat specialists and generalists drive homogenization and differentiation of temperate forest plant communities at the regional scale

In European temperate forests, changes in the compositional similarity among local plant communities (beta diversity) have rarely been studied due to the lack of adequate baseline data. Several studies, however, report the spread of common, generalist species or a decline in specialized, rare species. Both processes may lead to increased similarity among communities, i.e., biotic homogenization.To quantify changes in beta diversity, we resampled the herb layer composition of ancient broadleaf forests at 175 semi-permanent plots distributed across the Weser-Elbe region in NW Germany 20 years after first sampling. We hypothesized that beta diversity would have decreased on average as a result of a spread of habitat generalists and a decline in habitat specialists.After two decades, the forest communities did not yet exhibit severe biotic impoverishment, although there was a broad trend towards homogenization. The actual magnitude of change depended on which beta diversity measure was applied. The downward trend was primarily the result of the spread of native species that are able to tolerate broad pH and moisture ranges. A distinction between forest specialists (closely tied to forest habitats) and generalists (also found in open habitats) did not help explain changes in beta diversity. The study shows that on the regional scale and in habitats not yet threatened by the invasion of alien species, shifts in native species can promote biotic homogenization.