Ecological resilience of physical plant-soil feedback to chronic deer herbivory: Slow, partial, but functional recovery.

Ecological resilience to ungulate overbrowsing is an important issue in forest ecosystems. After chronic herbivory, the recovery rate of understory vegetation and its related functions can be slow even with decreasing grazing intensity; thus, detecting elasticity during alternative successional trajectories is fundamental to understanding state perturbations. In this context, we focused on physical plant-soil feedback (functional interactions between plant growth and soil physical conditions) and evaluated elasticity and recovery processes according to deer density. The effects of 40 years of chronic herbivory by sika deer (average density 14.7 individuals km ) on the recovery of understory plant communities and associated changes in soil physical properties in headwater catchments were assessed. Using 8 years of catchment-wide exclusion (fenced) and reduction (only culled; average 4.3 individuals km ) treatments, plot sampling was conducted in 2010 (before treatment) and 2018 (after treatment). The recovery of vegetation and soil physical properties were evaluated, and functional plant-soil relationships and spatial variability were assessed to detect recovery processes during alternative successional trajectory. Woody species increased only under the exclusion treatment and the average soil bulk density was lower than that under reduction treatments. Soil bulk density was negatively correlated with root biomass in the fenced catchment, and root biomass was positively associated with woody species richness. Reduced soil bulk density (~0.5 g cm ) was observed with greater root biomass and woody species richness on upper hillslopes in the deer-excluded catchment where plant coverage was minimal. Successional failure under the reduction treatment suggested slow recovery with a depressed threshold according to deer density, indicating a clockwise hysteretic response to deer density. Unlike plant coverage during the earlier period of overbrowsing, woody species root development led the recovery of functional physical plant-soil feedback; however, this was probably limited by the higher soil erosion rate in riparian areas and an under-developed herb layer. Our results highlight an alternative recovery trajectory of physical plant-soil feedback driven by an alternative plant element (woody roots) to recovery trajectory with increasing plant cover. However, riparian erosion and herb layer would still suppress recovery. Therefore, recovery might be slower at the landscape scale.