Linking vegetation and soil functions during secondary forest succession in the Atlantic forest

Secondary forest succession can be an effective and low-cost strategy to increase forest cover and the associated biodiversity and soil functions. However, little is known about how soil functions develop during succession, and how vegetation attributes influence soil functions, especially in highly biodiverse and fragmented landscapes in the tropics. Here we assessed a wide range of indicators of taxonomic (e.g. number of tree species), structural (e.g. basal area, canopy openness) and functional diversity (e.g. community weighted means of functional traits) of tree species, as well as indicators for soil functions related to soil organic matter accumulation, nutrient cycling and soil cover in secondary forest patches ranging from 5 to 80 years. Two recently abandoned agricultural fields were included as the starting point of forest succession and two primary forest patches served as references for the end point of forest succession. Four ecological hypotheses, centred around the role of functional diversity, structural diversity and biomass, were tested to explore mechanisms in which forest vegetation may influence soil functions. Most measures of structural, taxonomic and functional diversity converged to values found in primary forests after 25-50 years of succession, whereas functional composition changed from acquisitive to conservative species. Soil carbon and nutrient cycling showed a quick recovery to the levels of primary forests after 15 years of succession. Although soil cover also increased during succession, levels of primary forests were not reached within 80 years. Variation in tree height and trait dominance were identified as aboveground drivers of carbon and nutrient cycling, while aboveground biomass was the main driver of litter accumulation, and the associated soil cover and water retention. Our results indicate that secondary forest succession can lead to a relative fast recovery of nutrient and carbon cycling functions, but not of soil cover. Our findings highlight the essential role of secondary forests in providing multiple ecosystem services. These results can be used to inform management and reforestation programmes targeted at strengthening soil functions, such as soil cover, nutrient and carbon cycling.