Grassland diversity related to the Late Iron Age human population density

1 Species-rich semi-natural grasslands in Europe developed during prehistoric times and have endured due to human activity. At the same time, intensive grassland management or changes in land use may result in species extinction. As a consequence, plant diversity in semi-natural calcareous grasslands may be related to both historical and current human population density. 2 We hypothesize that current vascular plant diversity in semi-natural calcareous grasslands is positively correlated with the Late Iron Age (c. 800-1000 years ago) density of human settlements (indicated by Late Iron Age fortresses and villages) due to enhancement of grassland extent and species dispersal, and negatively correlated with current human population density due to habitat loss and deterioration. 3 We described the size of the community vascular plant species pool, species richness per 1 m(2) and the relative richness (richness divided by the size of the species pool) in 45 thin soil, calcareous (alvar) grasslands in Estonia. In addition to historical and current human population density we considered simultaneously the effects of grassland area, connectivity to other alvar grasslands, elevation above sea level (indicating grassland age), soil pH, soil N, soil P, soil depth, soil depth heterogeneity, geographical east-west gradient, precipitation and spatial autocorrelation. 4 Both the size of the community species pool and the species richness are significantly correlated with the Late Iron Age human population density. In addition, species richness was unimodally related to the current human population density. The relative richness (species 'packing density') was highest in the intermediate current human population densities, indicative of moderate land-use intensity. 5 Community species pool size decreased non-linearly with increasing soil N, and was highest at intermediate elevation. Small-scale richness was greater when sites were well connected and when the elevation was intermediate. Spatial autocorrelation was also significant for both species pool size and small-scale richness. 6 In summary, human land-use legacy from prehistoric times is an important aspect in plant ecology, which could be an important contributor to the current variation in biodiversity.