Land-use history, historical connectivity, and land management interact to determine longleaf pine woodland understory richness and composition

Restoration and management activities targeted at recovering biodiversity can lead to unexpected results. In part, this is due to a lack of understanding of how site-level characteristics, landscape factors, and land-use history interact with restoration and management practices to determine patterns of diversity. For plants, such factors may be particularly important since plant populations often exhibit lagged responses to habitat loss and degradation. Here, we assess the importance of site-level, landscape, and historical effects for understory plant species richness and composition across a set of 40 longleaf pine Pinus palustris woodlands undergoing restoration for the federally endangered red-cockaded woodpecker in the southeastern United States. Land-use history had an overarching effect on richness and composition. Relative to historically forested sites, sites with agricultural histories (i.e. former pastures or cultivated fields) supported lower species richness and an altered species composition due to fewer upland longleaf pine woodland community members. Landscape effects did not influence the total number of species in either historically forested or post-agricultural sites; however, understory species composition was affected by historical connectivity, but only for post-agricultural sites. The influences of management and restoration activities were only apparent once land-use history was accounted for. Prescribed burning and mechanical overstory thinning were key drivers of understory composition and promoted understory richness in post-agricultural sites. In historically forested sites these activities had no impact on richness and only prescribed fire influenced composition. Our findings reveal complex interplays between site-level, landscape, and historical effects, suggest fundamentally different controls over plant communities in longleaf pine woodlands with varying land-use history, and underscore the importance of considering land-use history and landscape effects during restoration.