Direct and productivity-mediated indirect effects of fertilization, mowing and grazing on grassland species richness

1. Recent declines in biodiversity have given new urgency to questions about the relationship between land-use change, biodiversity and ecosystem processes. Despite the existence of a large body of research on the effects of land use on species richness, it is unclear whether the effects of land use on species richness are principally direct or indirect, mediated by concomitant changes in ecosystem processes. Therefore, we compared the direct effects of land use (fertilization, mowing and grazing) on species richness with indirect ones (mediated via grassland productivity) for grasslands in central Europe. 2. We measured the richness and above-ground biomass in 150 grassland plots in 3 regions of Germany (the so-called Biodiversity Exploratories). We used univariate and structural equation models to examine direct and indirect land-use effects. 3. The direct effects of mowing (-0.37, effect size) and grazing (0.04) intensity on species richness were stronger compared with the indirect effects of mowing (-0.04) and grazing (-0.01). However, the strong negative effect of fertilization (-0.23) on species richness was mainly indirect, mediated by increased productivity compared with the weak direct negative effect (-0.07). 4. Differences between regions in land-use effects showed five times weaker negative effects of mowing (-0.13) in the region with organic soils (Schorfheide-Chorin), strong overall negative effects of grazing (-0.29) for the region with organic soils opposed to a similar strong positive effect (0.30) in the Hainich-Dün region, whereas the Schwäbische Alb region displayed a five times weaker positive effect (0.06) only. Further, fertilization effects on species richness were positive (0.03) for the region with organic soils compared to up to 25 times stronger negative effects in the other two regions. 5. Synthesis. Our results clearly show the importance of studying both direct and indirect effects of land-use intensity. They demonstrate the indirect nature, via productivity, of the negative effect of fertilization intensity on plant species richness in the real-world context of management-induced gradients of intensity of fertilization, mowing and grazing. Finally, they highlight that careful consideration of regional environments is necessary before attempting to generalize land-use effects on species diversity.