Drivers of biomass stocks and productivity of tropical secondary forests.

Young tropical secondary forests play an important role in the local and global carbon cycles because of their large area and rapid biomass accumulation rates. This study examines how environmental conditions and forest attributes shape biomass compartments and the productivity of young tropical secondary forests. We compared 36 young secondary forest stands that differed in the time since agricultural land abandonment (2.3-3.6 years) from dry and wet regions in Ghana. We quantified biomass stocks in living and dead stems, roots, and soil, and aboveground biomass and litter productivity. We used structural equation models to evaluate how macroclimate, soil nutrients (N, P), and forest attributes (structure, diversity, and functional composition) affect ecosystem functioning. After three years of succession, tropical wet forests stored on average 115 t biomass ha (the sum of aboveground living and dead biomass, belowground fine root biomass, and soil organic matter), and dry forests stored 99 t ha. These values represent 31% (in the wet forest) and 39% (in the dry forest) of the biomass compared with neighboring old-growth forests. The majority of forest ecosystem biomass was stored in the soil (70%) and aboveground living vegetation (25%). Macroclimate strongly shaped forest attributes, which in turn determined biomass stocks and productivity. Soil phosphorus strongly increased litter production and soil organic matter, confirming that it is a limiting element in tropical ecosystems. Tree density and species diversity increased forest biomass stocks, suggesting crown packing and complementary resource use enhance forest functioning. A more conservative trait composition (high wood density) increased biomass stocks but reduced productivity, indicating that quantity, identity, and quality of species affect ecosystem functioning.