Friend or foe? The role of biotic agents in drought-induced plant mortality

Plant mortality is a complex process influenced by both biotic and abiotic factors. In recent decades, widespread mortality events have been attributed to increasing drought severity, which has motivated research to examine the physiological mechanisms of drought-induced mortality, particularly hydraulic failure. Drought-based mortality mechanisms are further influenced by plant interactions with biota such as neighboring plants, insect pests, and microbes. In this review, we highlight some of the most influential papers addressing these biotic interactions and their influence on plant mortality. Plant???plant interactions can be positive (facilitation), neutral, or negative (competition), depending on drought intensity and neighbor identity. For example, stand-scale mortality likely increases with basal area (an index of competition). However, the diversity of forest stands matters, as more diverse forests suffer less mortality from drought than species-poor forests. Dense forest stands also increase bark beetle attack frequency, which can exacerbate drought stress and mortality, particularly for fast-growing species with lower defense allocation. In some cases, however, drought stress can alleviate biotic attack, depending on feedbacks between plant and pest physiology. Finally, plant interactions with beneficial microorganisms can increase drought tolerance, reduce the likelihood of mortality, and even extend plant distributions into drier habitats. Our review suggests more work is needed in natural herbaceous plant communities as well as dry tropical ecosystems where mortality mechanisms are less understood. Overall, relatively few studies directly link biotic interactions with the physiological mechanisms of mortality. Simultaneous manipulations of biotic interactions and measurements of physiological thresholds (e.g., xylem cavitation) are needed to fully represent biotic interactions in predictive models of plant mortality.