Natural regeneration triggers compositional and functional shifts in soil seed banks

Secondary forests emerging during traditional shifting cultivation practices are increasingly recognized for their fulfillment of ecosystem services and mitigation potential of climate change and biodiversity losses. The soil seed bank as a recruit reservoir is a limiting factor for natural forest regeneration of such secondary forests and is decisive for the formation and restitution of the post-disturbance community. The aim of this study was to compare the composition of the soil seed bank along a natural regeneration chronosequence from the Caxiuanã National Forest, eastern Amazon, including old-growth reference sites. We sampled standing vegetation, soil properties and soil seed banks and compared the density and species richness of different life forms among different regeneration stages. Using nonmetric, multiple scaling, we compared the composition of the soil seed bank among different regeneration stages and with standing vegetation composition. Furthermore, we outlined the influence of stand age, vegetation structure and soil properties on the density, richness and functional characterization of the soil seed bank using mixed effect models. The soil seed bank was dominated by herb seeds in all regeneration stages, and the density and richness of tree seeds increased with regeneration time and recovery of vegetation structure. Seed bank composition changed gradually with regeneration advance and differed from standing vegetation, containing a high amount of allochthonous seeds, especially in older stands. This observation highlights the importance of dispersal and habitat connectivity for the natural regeneration of these secondary forests. Shifts in soil seed bank composition towards slow-growing, animal-dispersed, non-pioneer species with larger, recalcitrant seeds in older regeneration stands indicate changes in vegetation composition along succession. Thus, our data indicate the importance of connectivity for forest regeneration and long fallow periods (> 40 years) to increase the performance of ecosystem services, resilience and stability of secondary forests arising during shifting cultivation practices.