High-diversity seed additions promote herb-layer recovery during restoration of degraded oak woodland.
Seed limitation represents a fundamental constraint to the restoration of native plant communities, and practitioners often apply seed additions to overcome this barrier. However, surprisingly few studies have experimentally tested whether seed additions can increase diversity in herbaceous communities of oak woodlands, which have undergone large-scale transformation due to logging, altered fire regimes and invasion by non-native species. Previous studies suggest that structural (thinning of woody biomass) and process-based (prescribed fire) restoration treatments alone are unlikely to restore the full breadth of taxonomic and functional diversity in the herb layer, which accounts for most species in woodland ecosystems. To explore whether seed additions can improve restoration outcomes in an oak woodland, we sowed high-diversity seed mixes in paired transects (seeded vs. controls) along a topographic gradient in a degraded site undergoing restoration with non-native shrub removal, selective tree thinning and prescribed fire. Seed mixes contained native forbs, grasses and sedges from locally sourced material (n=169 total species) in the regional species pool, and were designed to match species' habitat affinity to appropriate locations along the topographic gradient. The herb flora was sampled pre-seeding, and for two consecutive years after additions. Seed additions significantly altered community and functional composition, and increased native species richness by 29% (43.0 vs 55.4), and floristic quality by 30% relative to controls. However, fewer than half of the sown species were established 2 years after planting, suggesting that dispersal and establishment limitation are both important barriers to the recovery of the herb flora in oak woodlands. We also tested if species' sown abundance, conservatism or functional group predicted establishment success. Species sown at high abundances and less conservative species recruited the most reliably. Grass and forb establishment rates were more dependent on seeding rate than sedges or legumes, and the mechanisms behind this trend merit further investigation. We found that adding high-diversity seed mixes in conjunction with non-native shrub removal, canopy thinning and burning, can accelerate recovery of herbaceous communities in a highly degraded woodland.