Long-lasting negative effects of past forestation on grassland pollination networks and their function

O_LIGrasslands are facing a rapid decline worldwide. Among the primary threats to these ecosystems are changes in land use, such as land abandonment and forestation, which promote forest vegetation in detriment of grassland plant diversity. To support the conservation and restoration of grasslands, it is key to understand what ecological processes limit the recovery of their biodiversity and functions after perturbations. However, we still know little about the legacy effects of forestation on the ecological mechanisms involved in the recovery of grasslands, especially concerning long-lasting impacts on plant-pollinator interaction networks and plant reproduction. C_LIO_LIHere, we aim to fill this gap of knowledge by comparing the plant and pollinator diversity, the degree of network generalization, the pollination success and pollen limitation of native plant species in 30 plant-pollinator networks of old-growth and restored grasslands of different ages since recovery (from 2 to 84 years). We hypothesized that past forestation would exert long-lasting legacy negative effects on plant richness and plant-pollinator networks, increasing pollen limitation for native plants, thereby delaying community recovery in restored grasslands. C_LIO_LIResults showed that restored grasslands exhibited significantly lower plant richness, less specialized (more generalized) interaction networks, lower pollination success and pollen-limited seed reproduction of native plants compared to old-growth grasslands. Meanwhile, the degree of network specialization and pollination success gradually increased with time after grassland restoration initiated. Overall, network generalization, which was caused by low plant richness, reduced pollination and reproduction success in native grassland plants, and degraded pollination networks and functions could recover in restored grasslands with continuous management. These findings imply that plant diversity restoration was slow because of the negative feedbacks associated with low plant richness and consequently, generalized plant-pollinator interaction networks, which diminished native plant reproduction in restored grasslands. C_LIO_LISynthesis and applications Our findings suggest that the recovery of specialized plant-pollinator networks by enhancing plant diversity is essential for restoring pollination function. For quicker grassland restoration, it may be effective to facilitate the establishment of highly specialized pollination networks by seeding or planting diverse native plants collected from neighbouring areas while avoiding genetic contamination. C_LI