Soil fertility controls on tropical forest productivity and mortality: synthesis and roadmap.

Tropical forests are highly diverse and productive ecosystems and store nearly 60% of vegetation biomass. Across the tropics, forests span large gradients of soil fertility, from vast regions situated on highly weathered, ancient geological formations to others on young, nutrient-rich landscapes. Tropical forest productivity, mortality, and biomass all vary systematically across these gradients in soil fertility. Aboveground forest productivity tends to increase with higher soil fertility, while counterintuitively, aboveground biomass does not increase proportionally. This disconnect is likely due to coinciding increases in mortality with higher soil fertility. However, we know relatively little about the mechanisms underlying how soil fertility regulates productivity or mortality - two critical determinants of forest biomass - and even less about how these relationships will shape tropical forest responses to global change. Here, we present a mechanistic framework for understanding how soil fertility affects productivity through photosynthesis, carbon allocation, and carbon-use efficiency; how it modulates mortality via direct and indirect interactions with various drivers of mortality; and how global change may alter these relationships. Based on this synthesis, we outline a roadmap to advance our understanding of tropical forests and improve predictions of their responses to global change.