Does economic optimisation explain LAI and leaf trait distributions across an Amazon soil moisture gradient?

Abstract

Leaf area index (LAI) underpins terrestrial ecosystem functioning, yet our ability to predict LAI remains limited. Across Amazon forests, mean LAI, LAI seasonal dynamics and leaf traits vary with soil moisture stress. We hypothesise that LAI variation can be predicted via an optimality‐based approach, using net canopy C export (NCE, photosynthesis minus the C cost of leaf growth and maintenance) as a fitness proxy. We applied a process‐based terrestrial ecosystem model to seven plots across a moisture stress gradient with detailed in situ measurements, to determine nominal plant C budgets. For each plot, we then compared observations and simulations of the nominal (i.e. observed) C budget to simulations of alternative, experimental budgets. Experimental budgets were generated by forcing the model with synthetic LAI timeseries (across a range of mean LAI and LAI seasonality) and different leaf trait combinations (leaf mass per unit area, lifespan, photosynthetic capacity and respiration rate) operating along the leaf economic spectrum. Observed mean LAI and LAI seasonality across the soil moisture stress gradient maximised NCE, and were therefore consistent with optimality‐based predictions. Yet, the predictive power of an optimality‐based approach was limited due to the asymptotic response of simulated NCE to mean LAI and LAI seasonality. Leaf traits fundamentally shaped the C budget, determining simulated optimal LAI and total NCE. Long‐lived leaves with lower maximum photosynthetic capacity maximised simulated NCE under aseasonal high mean LAI, with the reverse found for short‐lived leaves and higher maximum photosynthetic capacity. The simulated leaf trait LAI trade‐offs were consistent with observed distributions. We suggest that a range of LAI strategies could be equally economically viable at local level, though we note several ecological limitations to this interpretation (e.g. between‐plant competition). In addition, we show how leaf trait trade‐offs enable divergence in canopy strategies. Our results also allow an assessment of the usefulness of optimality‐based approaches in simulating primary tropical forest functioning, evaluated against in situ data.