Species Richness, Community Specialization and Soil-Vegetation Relationships of Managed Grasslands in a Geologically Heterogeneous Landscape

Abstract
The increasing importance of the conservation value of managed grasslands has led to many studies exploring edaphic determinants of grassland biodiversity. Most studies, however, come either from very large areas, where biogeographical factors such as dispersal limitation may play a role, or from small, but ecologically rather uniform, regions. In addition, few studies further distinguish between plant specialists and generalists in the interpretation of the observed patterns. Here we studied species richness in semi-natural, managed grasslands in the Strážovské vrchy Mountains in the West Carpathians, Slovakia, where there is a matrix of different bedrocks (crystalline, sandstone, claystone, limestone) on a steep altitudinal gradient. In 89 vegetation plots we sampled the species composition of vascular plants and bryophytes and measured soil chemistry, slope angle, heat index, altitude and soil depth. We further applied Ellenberg indicator values and classified species into community specialists or generalists based on the analysis of a large phytosociological database. Using cluster analysis, we delimited five vegetation types that clearly differed in response to soil characteristics. Species richness varied between 19 and 64 species per 16 m2. The main compositional gradient correlated with measured soil pH and calcium, but species richness was not significantly correlated with these factors. Soil available phosphorus was not associated with species composition as has been found elsewhere, but it did correlate negatively with species richness and the richness of specialists. Overall, species richness was largely driven by the number of specialists in the plot and particular vegetation types differed conspicuously in their number. We further found significant effects of iron, potassium and sodium on species richness, species composition and the representation of specialists and generalists. Our results provide new insights into the determinants of diversity in managed grasslands as well as to the theoretical species pool concept, explaining species richness variation along a pH gradient.