Three decades of post-logging tree community recovery in naturally regenerating and actively restored dipterocarp forest in Borneo

Selective logging has affected large areas of tropical forests and there is increasing interest in how to manage selectively logged forests to enhance recovery. However, the impacts of logging and active restoration, by liberation cutting and enrichment planting, on tree community composition are poorly understood compared to trajectories of biomass recovery. Here, we assess the long-term impacts of selective logging and active restoration for biomass recovery on tree species diversity, community composition, and forest structure. We censused all stems >= 2 cm diameter at breast height (DBH) on 46 permanent plots in unlogged, primary forest in the Danum Valley Conservation Area (DVCA; 12 plots, totalling 0.6 ha) and in sites logged 23-35 years prior to the census in the Ulu Segama Forest Reserve adjacent to DVCA (34 plots, totalling 1.7 ha) in Sabah, Malaysian Borneo. Active restoration treatments, including enrichment planting and climber cutting, were implemented on 17 of the logged forest plots 12-24 years prior to the census. Total plot-level basal area and pole (5-10 cm DBH) stem density were lower in logged than unlogged forests, however no difference was found in stem density amongst saplings (2-5 cm DBH) or established trees (>= 10 cm DBH). Neither basal area, nor plot-level stem density varied with time since logging at any size class, although sapling and pole stem densities were lower in actively restored than naturally regenerating logged forest. Sapling species diversity was lower in logged than unlogged forest, however there were no other significant effects of logging on tree species richness or diversity indices. Tree species composition, however, differed between logged and unlogged forests across all stem size classes (PER-MANOVA), reflected by 23 significant indicator species that were only present in unlogged forest. PER-MANOVA tests revealed no evidence that overall species composition changed with time since logging or with active restoration treatments at any size class. However, when naturally regenerating and actively restored communities were compared, two indicator species were identified in naturally regenerating forest and three in actively restored forests. Together our results suggest that selective logging has a lasting effect on tree community composition regardless of active restoration treatments and, even when species richness and diversity are stable, species composition remains distinct from unlogged forest for more than two decades post-harvest. Active restoration efforts should be targeted, monitored, and refined to try to ensure positive outcomes for multiple metrics of forest recovery.