Soil moisture and a legacy of prehistoric human activities have contributed to the extraordinary plant species diversity of grasslands in the White Carpathians

The factors that determine the unique species richness and composition of some temperate grasslands are poorly known. Uniqueness of the extraordinarily species-rich grasslands in the White Carpathian mountain range (Czech Republic, Slovakia), with many disjunct occurrences of species, have been previously attributed to intermittently wet deep soils, which facilitate the co-occurrence of steppe and wet-grassland species, and Holocene continuity of open land, resulted in large species pools. Based on a detailed investigation of 23 well-preserved regularly mown grasslands differing in their vegetation composition and species richness, we tested the relative importance of the hypothetical determinants of compositional variability within semi-dry grasslands. For the first time we included measurements of seasonal moisture at different soil depths and landscape differences in the intensity of the effect of prehistoric humans. Soil moisture was measured using the frequency-domain reflectometry based Profile Probe PR2 sensor at depths of 10, 20, 30, 40 and 60 cm in the soil, repeatedly every four weeks from April to November. Soil samples were taken from the same depths in order to determine soil basicity and nutrient content. As a proxy of the intensity of the effect of prehistoric human activities, mean distances between sampled localities and the five nearest Neolithic or Aeneolithic settlements were used. The species richness was best explained by mean soil moisture, which increased towards the most species-rich grasslands, whereas the distance from prehistoric settlements had no effect. Basicity, moisture and the distance from prehistoric settlements had significant conditional effects on species composition. There was a high frequency of species of intermittently wet soils, thermophilous oak forests and forest fringes in the most valuable grasslands, which are located mostly in the south-western part of the area where the soils were moister and closer to prehistoric settlements, but the seasonal variation in moisture was not high. We conclude that coexistence of species from different habitats is dependent on regular management and high soil moisture throughout the growing season. Due to intermediate moisture conditions that are tolerated by multiple ecological groups of species, many species may locally coexist and form a species-rich grassland community of unique species composition. In addition, prehistoric human activities contributed to landscape openness and helped maintain a forest-steppe species pool during the Holocene forest optimum. Our results indicate that although this historical determinant shapes predominantly the species composition pattern, the extraordinarily species-rich spots are determined rather by abiotic factors, such as intermediate moisture and biotic interactions resulting from the type and duration of management practices.