Changes in Species and Functional Diversity of the Herb Layer of Riparian Forest despite Six Decades of Strict Protection

Abstract

The herb layer of temperate forests contributes to long-term forest ecosystem functioning and provisioning of ecosystem services. Therefore, a thorough understanding of its dynamics in the face of environmental changes is essential. This paper focuses on the species and functional diversity of the herb layer of riparian forests to verify how these two community components changed over time and under strict protection. The understory vegetation was surveyed on 42 semi-permanent plots in three time periods between 1960 and 2020. The overall pattern in vegetation changes that related to species richness and diversity, functional structure, and habitat conditions was analyzed using ordination and permutation techniques. We found significant changes in species composition and the functional structure of herbaceous vegetation over the last six decades. Forests were enriched with nutrient-demanding and alien species. A significant increase in functional diversity and the proportion of species with high SLA and canopy height was also observed, whereas changes in habitat conditions were insignificant. The observed trends indicate that the strict protection of forest communities within small and isolated reserves does not fully protect their species composition. Forest reserves should be surrounded by unmanaged forests and spatially connected to allow species mobility.