Scale-dependent patterns and drivers of vascular plant, bryophyte and lichen diversity in dry grasslands of the Swiss inneralpine valleys

The inner-alpine dry valleys of the Swiss Alps are characterized by subcontinental climate, leading to many peculiarities in dry grassland species composition. Despite their well-known uniqueness, comprehensive studies on biodiversity patterns of the dry grasslands in these valleys were still missing. To close this gap, we sampled 161 10-m2 vegetation plots in the Rhône, Rhine and Inn valleys, recording vascular plants, terricolous bryophyte and lichen species, as well as environmental data. Additionally, we tested the scale-dependence of environmental drivers using 34 nested-plot series with seven grain sizes (0.0001–100 m2). We analysed the effects of environmental drivers related to productivity/stress, disturbance and within-plot heterogeneity on species richness. Mean species richness ranged from 2.3 species in 0.0001 m2 to 58.8 species in 100 m2. For all taxa combined, the most relevant drivers at the grain size of 10 m2 were southing (negative), litter (negative), mean annual precipitation (unimodal), gravel cover (negative), inclination (unimodal) and mean annual precipitation (unimodal). For vascular plants the pattern was similar, while bryophyte and lichen richness differed by the opposite relationship to mean annual precipitation as well as negative influences of mean herb layer height, grazing and mowing. The explained variance of the multiple regression model increased with grain size, with very low values for the smallest two grain sizes. While southing and litter had high importance for the fiver larger grain sizes, pH and gravel cover were particularly important at the intermediate grain sizes, and inclination and mean annual precipitation for the two largest grain sizes. The findings emphasize the importance of taxonomic group and grain size for patterns and drivers of species richness in vegetation, consistent with ecological theory. Differences in the diversity–environment relationships among the three taxonomic groups can partly be explained by asymmetric competition that leads to low bryophyte and lichen diversity where vascular plants do well and vice versa. The relatively low alpha diversity of vascular plants in dry grasslands in Swiss inner-alpine valleys compared to similar communities in other parts of the Palaearctic remains puzzling, especially because Swiss stands are often large and well-preserved.