Half a century of herb layer changes in Quercus-dominated forests of the Western Carpathians

Abstract

Species composition of European temperate Quercus forests have undergone dramatic changes in the past century. Observed changes have largely been attributed to anthropogenic environmental changes that drove the processes of thermophilization, eutrophication, or biotic homogenization. However, ecological processes behind these changes can be difficult to explain considering the variability among regions: their history, management, landscape patterns, site properties, and vegetation types. To reveal vegetation change trends and their drivers in the subcontinental Western Carpathians, we analysed vegetation and environmental data from 158 quasi-permanent plots established between 1961 and 1980 in Quercus petraea- dominated forests on the andesite bedrock of the Slovenské stredohorie Mountains in Slovakia (Central Europe), and we resurveyed them again between 2005 and 2020.Among alpha and gamma diversity measures, we found only slight species richness decreases. A significant shift in species composition associated with very high species turnover (64%) and a decrease in herb layer cover (by 24%) resulted in higher spatial heterogeneity of herb understoreys in the resurvey. Winning species were more thermophilous, eutrophic, and mesic/oceanic plants of lower latitudes, replacing original species of open Quercus forests. Increased temperatures was the most significant environmental driver. Climate warming corresponded not only to thermophilization, but, after interaction with management, it was also related to eutrophication of the herb layer. Eutrophication was identified as the main trend of the species composition changes (mean EIV for nutrients of winners was 5.9 compared to 4.1 of losers) even though the effect of anthropic nitrogen depositions was negligible. Successive undergrowing of Quercus canopies by more shade-casting trees (Acer campestre, Carpinus betulus, and Fagus sylvatica) in the period of decreasing management intensity was another major driver. Their increase by 8.6% in the resurveyed plots contributed to decreased species richness and they supported shade-tolerant eutrophic understorey species typical of mesic Carpinus betulus and Fagus sylvatica forests.We concluded that Quercus forests in the Western Carpathians underwent thermophilization and eutrophication, similar to the forests in densely inhabited Western Europe; however, their plant understoreys were not homogenized, and eutrophication was not driven by nitrogen depositions. Eutrophication should be recognised as the next important negative impact of global warming on ecosystems. Dramatic changes in species composition and loss of original characteristic species can only be partly halted by reducing shade-casting trees from lower tree layers in managed stands.