Responses of grassland species richness to local and landscape factors depend on spatial scale and habitat specialization

Questions: To what extent is species richness in semi-natural grasslands related to local environmental factors and (present/ past) surrounding landscape structure? Do responses of species richness depend on degree of habitat specialization (specialists vs generalists) and/ or scale of the study? Location: O land, Sweden. Methods: Richness of herbaceous vascular plants (subdivided into richness of grassland specialists and generalists) was recorded within 50 9 50 cm plots and 0.1-4.8 ha grassland polygons. Generalized linearmodels and hierarchical partitioning were used to identify local factors (habitat area and heterogeneity, grazing intensity, habitat continuity) and landscape factors (proportion of surrounding grassland in 2004, 1938 and 1800, and landscape diversity in 2004) associated with the richness estimates. Results: At the polygon scale, both specialist and generalist richness was positively associated with local habitat area and heterogeneity and, independently of area and heterogeneity, with grazing intensity, habitat continuity and amount of surrounding grassland in 1800. At the plot scale, specialist species richness was positively associated with habitat heterogeneity, amount of surrounding grassland in 2004 and landscape diversity. Plot-scale generalist richness was negatively associated with surrounding grassland in 1938 and positively associated with local grazing intensity. Conclusions: Because both habitat specialization and study scale influence conclusions about relationships between species richness and local and landscape factors, the study highlights the need to consider species diversity at multiple spatial scales when making decisions about grassland management. Large-scale (polygon) species richness is influenced by immigration processes, with both specialists and generalists accumulating in old grasslands over centuries of grazing management. Habitat heterogeneity increased specialist species richness at both scales, suggesting that management policies should favour maintenance of a heterogeneous mosaic of open areas, trees and shrubs in temperate grazed grasslands. Although grassland specialists are sensitive to grassland isolation, in extensively managed landscapes with high landscape diversity input of grassland species from the landscape matrix may buffer negative effects of habitat fragmentation on grassland communities.