The structure of maturity: immature trees may drive the productivity of mature forests

Relating forest productivity to local variations in forest structure has been a long-standing challenge. Previous studies often focused on the connection between forest structure and stand-level photosynthesis (GPP). However, biomass production (NPP) and net ecosystem exchange (NEE) are also subject to respiration and other carbon losses, which vary with local conditions and life history traits. Here, we use a simulation approach to study how these losses impact forest productivity and reveal themselves in forest structure. We fit the process-based forest model Formind to a 25ha inventory of an old-growth temperate forest in China and classify trees as "mature" (full-grown) or "immature" based on their intrinsic carbon use efficiency. Our results reveal a strong negative connection between the stand-level carbon use efficiency and the prevalence of mature trees: GPP increases with the total basal area, whereas NPP and NEE are driven by the basal area of immature trees. Accordingly, the basal area entropy - a structural proxy for the prevalence of immature trees - correlated well with NPP and NEE and had higher predictive power than other structural characteristics such as Shannon diversity and height standard deviation. Our results were robust across spatial scales (0.04-1ha) and yield promising hypotheses field studies and new theoretical work.