Patterns of fine-scale plant species richness in dry grasslands across the eastern Balkan Peninsula

Abstract

Fine-scale plant species richness varies across habitats, climatic and biogeographic regions, but the large-scale context of this variation is insufficiently explored. The patterns at the borders between biomes harbouring rich but different floras are of special interest. Dry grasslands of the eastern Balkan Peninsula, situated in the Eurasian forest-steppe zone and developed under Mediterranean influence, are a specific case of such biome transition. However, there are no studies assessing the patterns of fine-scale species richness and their underlying factors across the eastern Balkans. To explore these patterns, we sampled dry and semi-dry grasslands (phytosociological class Festuco-Brometea) across Bulgaria and SE Romania. In total, 172 vegetation plots of 10 × 10 m2 were sampled, in which all vascular plant species were recorded, soil depth was measured, and soil samples were collected and analysed in a laboratory for pH and plant-available nutrients. Geographic coordinates were used to extract selected climatic variables. Regression trees and linear regressions were used to quantify the relationships between species richness and environmental variables. Climatic factors were identified as the main drivers of species richness: (1) Species richness was strongly positively correlated with the mean temperature of the coldest month: sub-Mediterranean areas of S and E Bulgaria, characterized by warmer winters, were more species-rich. (2) Outside the sub-Mediterranean areas, species richness strongly increased with annual precipitation, which was primarily controlled by altitude. (3) Bedrock type and soil pH also significantly affected dry grassland richness outside the sub-Mediterranean areas. These results suggest that fine-scale species richness of dry grasslands over large areas is driven by processes at the regional level, especially by the difference in the species pools of large regions, in our case the Continental and Mediterranean biogeographic regions. Local environmental factors are of secondary importance over broad extents, but their effect on fine-scale species richness increases within climatically and biogeographically homogeneous regions.