Patterns and mechanisms of spatial variation in tropical forest productivity, woody residence time, and biomass.

Tropical forests vary widely in biomass carbon (C) stocks and fluxes even after controlling for forest age. A mechanistic understanding of this variation is critical to accurately predicting responses to global change. We review empirical studies of spatial variation in tropical forest biomass, productivity and woody residence time, focusing on mature forests. Woody productivity and biomass decrease from wet to dry forests and with elevation. Within lowland forests, productivity and biomass increase with temperature in wet forests, but decrease with temperature where water becomes limiting. Woody productivity increases with soil fertility, whereas residence time decreases, and biomass responses are variable, consistent with an overall unimodal relationship. Areas with higher disturbance rates and intensities have lower woody residence time and biomass. These environmental gradients all involve both direct effects of changing environments on forest C fluxes and shifts in functional composition - including changing abundances of lianas - that substantially mitigate or exacerbate direct effects. Biogeographic realms differ significantly and importantly in productivity and biomass, even after controlling for climate and biogeochemistry, further demonstrating the importance of plant species composition. Capturing these patterns in global vegetation models requires better mechanistic representation of water and nutrient limitation, plant compositional shifts and tree mortality.