There is widespread concern regarding declines in bee populations given their importance for the functioning of both natural and managed ecosystems. An increasing number of studies find negative relations between bee species richness and simplification of agricultural landscapes, but the role of land-use intensity and its relative importance compared to landscape simplification remain less clear. We compared the relative effects of nitrogen inputs, as a proxy for land-use intensity, and proportion of natural and semi-natural habitat, as a measure of landscape complexity on total bee species richness, rare species richness and dominant crop-visiting species richness. We used data from 282 grasslands across five countries, covering the entire range of low intensity, no-input systems, to high-input sites (>400 kg N/ha/year). We found consistent negative impacts of increasing land-use intensity at a regional scale on total bee species richness and dominant crop-visiting species across Europe, but no such effects of landscape complexity. In contrast, the richness of rare bee species was not significantly related to increasing land-use intensity. Nevertheless, based on species accumulation curves, grasslands with no nitrogen inputs had higher total bee richness and higher shares of rare species compared with sites with high nitrogen inputs (>125 kg N/ha/year). Our results highlight the importance of retaining grasslands characterised by low land-use intensity across agricultural landscapes to promote bee diversity.
