Restoration of plant diversity in permanent grassland by seeding: assessing the limiting factors along land-use gradients.
Land-use intensification has caused dramatic declines in grassland biodiversity, and in Europe this decline has not yet been halted. Therefore, a major challenge in ecological restoration is to effectively restore plant diversity in permanent grasslands, that is, long-term managed grassland not being used as cropland. As species richness is often limited by seed dispersal, introducing locally absent species is crucial. Seeding diverse seed mixtures is an appropriate tool, but restoration outcomes are highly variable. Considering constraints by land-use intensity and plant functional traits during restoration could improve the efficacy of restoration efforts. We established a full-factorial experiment with high-diversity seeding and topsoil disturbance in 73 grasslands along land-use intensity and productivity gradients and evaluated the restoration success over 5 years. In addition, after 5 years we assessed the number of established species and tested whether specific traits promoted the establishment of sown species in interaction with productivity. Plant diversity increased considerably when seeding and disturbance were combined while seeding alone only slightly increased species richness. Along land-use gradients, we found no direct effects of grazing or mowing on the establishment of sown species, but fertilization indirectly limited establishment in the combined treatment via a negative effect of productivity. Functional traits structured the establishment of sown species, especially in the combined treatments, but effects varied with productivity. Species with low seed mass and low specific leaf area had greater establishment success in low-productivity grasslands, whereas species with high seed mass had greater establishment in high-productivity grasslands.