Nitrogen enrichment and vascular plant richness loss reduce bryophyte richness

Grasslands' high diversity is threatened by land-use changes, such as nitrogen fertilization, leading to productive but low-richness, fast-growing plant communities. Bryophytes are a key component of grassland diversity and react strongly to land use. However, it is unclear whether land-use effects are direct or mediated by changes in vascular plants. Increases in vascular plant cover are likely to decrease bryophyte abundance through light competition. Whether changes in vascular plant composition and richness also play a role remains unclear. We sampled bryophytes in a factorial grassland experiment manipulating nitrogen fertilization, fungicide, species richness, and functional composition of vascular plants crossed with moderate disturbances by weeding. Disturbance increased bryophyte richness and modulated treatment effects. In contrast to previous studies reporting indirect negative fertilization effects via increasing vascular plant productivity and reduced light levels, nitrogen fertilization directly reduced bryophyte cover and species richness, possibly because of toxic effects. Low vascular plant richness and dominance of fast-growing species reduced bryophyte richness. This might be because of decreased structural and resource niche heterogeneity in species-poor communities. Our results highlight novel mechanisms by which land-use intensification can affect bryophytes and suggest that a loss of vascular plant richness might have cascading effects on other taxonomic groups.