Disentangling the Ecological Determinants of Species and Functional Trait Diversity in Herb-Layer Plant Communities in European Temperate Forests

Abstract

Forest herb-layer vegetation responds sensitively to environmental conditions. This paper compares drivers of both taxonomic, i.e., species richness, cover and evenness, and functional herb-layer diversity, i.e., the diversity of clonal, bud bank and leaf-height-seed plant traits. We investigated the dependence of herb-layer diversity on ecological determinants related to soil properties, climatic parameters, forest stand characteristics, and topographic and abiotic and biotic factors associated with forest floor structure. The study was conducted in different forest types in Slovenia, using vegetation and environmental data from 50 monitoring plots (400 m2 each) belonging to the ICP Forests Level I and II network. The main objective was to first identify significant ecological predictors and then quantify their relative importance. Species richness was strongly determined by forest stand characteristics, such as richness of the shrub layer, tree layer shade-casting ability as a proxy for light availability and tree species composition. It showed a clear positive relation to soil pH. Variation in herb-layer cover was also best explained by forest stand characteristics and, to a lesser extent, by structural factors such as moss cover. Species evenness was associated with tree species composition, shrub layer cover and soil pH. Various ecological determinants were decisive for the diversity of below-ground traits, i.e., clonal and bud bank traits. For these two trait groups we observed a substantial climatic signal that was completely absent for taxonomy-based measures of diversity. In contrast, above-ground leaf-height-seed (LHS) traits were driven exclusively by soil reaction and nitrogen availability. In synthesis, local stand characteristics and soil properties acted as the main controlling factors for both species and trait diversity in herb-layer communities across Slovenia, confirming many previous studies. Our findings suggest that the taxonomic and functional facets of herb-layer vegetation are mainly influenced by a similar set of ecological determinants. However, their relative importance varies among individual taxonomy- and functional trait-based diversity measures. Integrating multi-faceted approaches can provide complementary information on patterns of herb-layer diversity in European forest plant communities.