A mechanistic model of tree competition and facilitation for Mediterranean forests: Scaling from leaf physiology to stand dynamics

Mechanistic theories of plant competition developed to explain changes in community structure and dynamics along resource availability gradients have been mostly applied to temperate forests and grasslands where light and nutrients are the two main limiting resources. In contrast, the mechanisms underlying the structure and dynamics of water-limited plant communities have been little explored. Also previous mechanistic models rely either on complex simulators, which are difficult to interpret or on simple conceptual models, which ignore too many critical details. In this study, we develop a model of stand dynamics for light and water-limited forests of intermediate complexity and we provide an analytical framework for its analyses. The model is an individual-based simulator that describes the feedback between transpiration, stomatal function and soil water dynamics with asymmetrical competition for light and water. Trees allocate carbon to three main compartments: shoot, stem and roots. We use the model to explore general patterns that may emerge across levels of biological organization from the leaf to the stand. Model predictions are consistent with a number of features of Mediterranean forests structure and dynamics. At the plant-level the leaf-based tradeoff between carbon gain and water loss expresses as a tradeoff between mortality and growth. This tradeoff explains plant morphological changes in above-ground biomass and root to shoot allocation along a water availability gradient. At the community-level, tradeoffs among carbon acquisition and water loss govern the sign of plant interactions along the gradient. Coexistence among morphological types was not observed for the range of parameters and environmental conditions explored. Overall the model provides an unifying explanation for the observed changes in the sign of plant to plant interactions along environmental gradients as well as a process-based formulation that can be linked to empirical studies.