The role of species interactions for forest resilience to drought.

Increasing durations and frequencies of droughts under climate change endanger the sustainable functioning of forests worldwide. The admixture of species with complementary resource use may increase the resilience of forests towards drought; however, little is known about modifications of species interactions (i.e. facilitation and competition) by increasing drought severity in mixed forests. In particular, knowledge on the regulation of central ecohydrological processes, such as tree water fluxes, is lacking. Therefore, we conducted a literature review to assess the impact of species interactions on tree resilience (resistance + recovery) under increasing drought severity. The classification of studies into three drought classes suggested that beneficial species interactions, i.e. through improved water relations, were prevalent under mild droughts. However, with increasing drought, negative effects, such as interspecific competition, occurred. These negative interactions were prominent under extreme droughts, where even trees with complementary resource-use strategies competed for water resources. Fewer data are available on recovery of water fluxes. The limited evidence supported the patterns observed for drought resistance, with facilitation and complementarity of species in mixtures enhancing tree recovery after moderate droughts. However, after extreme droughts, competition effects and reduced recovery for some species were observed, which can strongly compromise tree resilience. While we acknowledge the importance of mixed forests for biodiversity, ecosystem services or pest resistance, we caution that beneficial species interactions may shift under extreme droughts. Thus, there is an urgent need to investigate species interaction effects on resilience in more depth to adapt forest trees to increasing drought stress.