Mechanisms of a coniferous woodland persistence under drought and heat

Predictions of warmer droughts causing increasing forest mortality are becoming abundant, yet few studies have investigated the mechanisms of forest persistence. To examine the resistance of forests to warmer droughts, we used a five-year precipitation reduction (~45% removal), heat (+4 °C above ambient) and combined drought and heat experiment in an isolated stand of mature Pinus edulis-Juniperus monosperma. Despite severe experimental drought and heating, no trees died, and we observed only minor evidence of hydraulic failure or carbon starvation. Two mechanisms promoting survival were supported. First, access to bedrock water, or 'hydraulic refugia' aided trees in their resistance to the experimental conditions. Second, the isolation of this stand amongst a landscape of dead trees precluded ingress by Ips confusus, frequently the ultimate biotic mortality agent of piñon. These combined abiotic and biotic landscape-scale processes can moderate the impacts of future droughts on tree mortality by enabling tree avoidance of hydraulic failure, carbon starvation, and exposure to attacking abiotic agents.This project was supported by the Department of Energy, Office of Science, and Pacific Northwest National Lab’s LDRD program. DDB participation was supported via NSF EF-1340624; EF-1550756, and EAR-1331408, DEB-1824796 and DEB-1833502. CG was supported by a Director’s Fellowship from the Los Alamos National Laboratory and by the Swiss National Science Foundation SNF (PZ00P3_174068). AV was supported by a fellowship from Generalitat Valenciana (BEST/2016/289) and the project Survive-2 (CGL2015-69773-C2-2-P MINECO/FEDER) from the Spanish Government. DSM was supported via NSF IOS-1450679, IOS-1444571, and IOS-1547796

A multi-species synthesis of physiological mechanisms in drought-induced tree mortality

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere–atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function