Moderate grazing weakens legacy effects of grazing history on primary productivity and biodiversity in a meadow grassland

Herbivore grazing has long-lasting effects on the structure and functions of grassland ecosystems, even after the grazing pressure has ceased. These lasting effects are known as grazing legacy effects and significantly affect plant productivity and biodiversity, with the magnitude increased with grazing intensity and further led to grassland degradation. Fencing is a common approach used for restoring degraded grasslands by excluding grazing. Moderate grazing representing a controlled level of grazing intensity is believed to be a useful way to restore ecological functions and thus mitigate grazing legacies. However, how fencing and ongoing moderate grazing affect grazing legacy effects remain unclear and rarely tested with field experiments, particularly with different grazing histories. Here, we investigated aboveground net primary productivity (ANPP), species richness, litter, soil organic carbon (SOC), soil dissolved nitrogen (DON) and hyphal length density (HLD) in 2015 (fencing for 1 year) and 2018 (fencing vs moderate grazing for the subsequent 3 years) after 0, 2, 5, 7 and 12 years of free grazing in a meadow steppe to explore how moderate grazing and fencing affect grazing legacy effects with different histories. We observed a decline in ANPP and species richness with increasing grazing histories, and the magnitude of grazing legacy effects increased accordingly. In most grazing histories, moderate grazing had a greater impact than fencing in reducing the grazing legacy effects on ANPP and species richness. However, with a grazing history of 12 years, moderate grazing led to a smaller increase in ANPP compared to fencing. Additionally, the grazing legacy effects on ANPP and species richness weakened with longer fencing duration for excluding grazing. Litter, SOC, DON and HLD exhibited consistent patterns with the dynamics of ANPP and species richness. Our results indicate that grazing legacy effects on ANPP and species richness intensify with prolonged grazing histories, suggesting that grazing history have stronger impacts on grassland ecosystem functions and structure. The study implies that moderate grazing is an effective strategy for restoring degraded grasslands, particularly for species richness and with short-term grazing histories, instead of relying solely on fencing. The changes of soil variables underlie the grazing legacy effects and the interactions with fencing and moderate grazing.