Nucleation increases understory species and functional diversity in early tropical forest restoration

Degraded tropical forests that were converted into pastures dominated by exotic grasses are a challenge for restoration due to arrested succession. Nucleation is a suitable strategy to restore such abandoned pastures as it can overcome seed and site limitation, and would foster the recovery of structural, functional, and species diversity. To explore the outcomes of different nucleation techniques in terms of richness, functional diversity, resource offer, and species and trait composition during the first year after implementation in an abandoned pasture in the tropical Andes, we conducted a field experiment with four nucleation treatments established in 1m(2) subplots, i.e. natural regeneration (control), seed rain transfer, seed bank transfer, and pre-grown plant mats. Number of species, functional diversity, proportion of natives, flowering and fruiting were response variables of the nucleation treatments and time since restoration using linear mixed-effects models. Species and trait composition trajectories over time were depicted with NMDS. Species richness and functional diversity of understory vegetation increased over time and with nucleation, being significantly higher by the final survey. Seed bank transfer introduced most species to the area in comparison to natural regeneration. Functional diversity, proportion of native, flowering, and fruiting species changed with time albeit with no differences among treatments. Species and trait composition showed convergence over time, which stresses the strong influence of environmental filtering in early restoration of abandoned pastures. Although positive aspects of planting trees and shrubs cannot be neglected, we argue that the incorporation of (several) more sophisticated nucleation techniques is beneficial for restoration of tropical forests.