Relationship between extreme species richness and Holocene persistence of forest-steppe grasslands in Transylvania, Romania

The most species-rich grasslands worldwide are known from the Carpathian Mts and their periphery in East-Central Europe. They occur in forest-steppe regions, transitional between temperate forest and arid steppe biomes. Their climate, largely suitable for forests, raises questions about the origin of these grasslands. Have they been forested in the past, or locally maintained through a disturbance regime? We addressed these questions to contribute to the broader understanding of Holocene dynamics of open habitats in temperate Europe. We employed soil charcoal analysis and soil morphology to reconstruct past representation of woody species with fine spatial resolution. Our study area was Romanian Transylvania, a region renowned for a well-developed forest-steppe. Six soil profiles along a climatic gradient were assessed: four in forest-steppe grasslands, two in grasslands in adjacent forest region (forest grasslands). The results revealed profound differences between forest-steppe and forest grasslands. Forest-steppe profiles showed Phaeozems with low specific anthracomass of woody species and continuous dominance by Juniperus, suggesting a long-term presence of grasslands. Forest grasslands showed Luvisols with higher anthracomass and abundant charcoal of broad-leaved trees, indicating establishment after deforestation. The high radiocarbon ages of charcoals in basal soil horizons point to a glacial origin of soils and the link of forest-steppe grasslands to glacial forests. Siberian hemiboreal forests and related grasslands may be modern analogues of the reconstructed ecosystems, sharing many species with present day forest-steppe. We suggest that disturbances such as fire, herbivore grazing, and human activities have played an important role in shaping the forest-steppe over time, contributing to the formation of today’s richest grasslands.