Neighbourhood species richness and drought-tolerance traits modulate tree growth and δ13C responses to drought

Mixed-species forests are promoted as a forest management strategy for climate change mitigation and adaptation because they are more productive and can be more resistant and resilient than monospecific forests under drought stress. However, the trait-based mechanisms driving these properties remain elusive, making it difficult to predict which functional identities of species best improve tree growth and decrease tree physiological water stress under drought. We investigated tree growth and physiological stress responses (i.e. increase in wood carbon isotopic ratio; δ13C) to changes in climate-induced water availability (wet-to-dry years) along gradients in neighbourhood tree species richness and drought-tolerance traits. Using tree cores from a large-scale biodiversity experiment, we tested the overarching hypothesis that neighbourhood species richness increases growth and decreases δ13C. We further hypothesized that the abiotic (i.e. climatic conditions) and the biotic context modulate these biodiversity-ecosystem functioning relationships. We characterized the biotic context using drought-tolerance traits of focal trees and their neighbours. These traits are related to cavitation resistance vs. resource acquisition and stomatal control. We found that tree growth increased with neighbourhood species richness. However, we did not observe a universal relief of water stress in species-rich neighbourhoods, nor an increase in the strength of the relationship between richness and growth and between richness and δ13C from wet-to-dry years. Instead, these relationships depended on both the traits of the focal trees and their neighbours. At either end of each drought-tolerance gradient, species responded in opposing directions during drought and non-drought years. Synthesis. We report that the biotic context can determine the strength and nature of biodiversity-ecosystem functioning relationships in experimental tree communities. We derive two key conclusions: (1) drought-tolerance traits of focal trees and their neighbours can explain divergent tree responses to drought and diversity, and (2) contrasting, trait-driven responses of tree species to wet vs dry climatic conditions can promote forest community stability. Mixing tree species with a range of drought-tolerance traits may therefore increase forest productivity and stability