An individual-based forest model links canopy dynamics and shade tolerances along a soil moisture gradient

Understanding how forested ecosystems respond to climatic changes is a challenging problem as forest self-organization occurs simultaneously across multiple scales. Here, we explore the hypothesis that soil water availability shapes above-ground competition and gap dynamics, and ultimately alters the dominance of shade tolerant and intolerant species along the moisture gradient. We adapt a spatially explicit individual-based model with simultaneous crown and root competitions. Simulations show that the transition from xeric to mesic soils is accompanied by an increase in shade-tolerant species similar to the patterns documented in the North American forests. This transition is accompanied by a change from water to sunlight competitions, and happens at three successive stages: (i) mostly water-limited parkland, (ii) simultaneously water- and sunlight-limited closed canopy forests featuring a very sparse understory, and (iii) mostly sunlight-limited forests with a populated understory. This pattern is caused by contrasting successional dynamics that favour either shade-tolerant or shade-intolerant species, depending on soil moisture and understory density. This work demonstrates that forest patterns along environmental gradients can emerge from spatial competition without physiological trade-offs between shade and growth tolerance. Mechanistic understanding of population processes involved in the forest–parkland–desert transition will improve our ability to explain species distributions and predict forest responses to climatic changes.