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.

Tree mortality from drought, insects, and their interactions in a changing climate.

Climate change is expected to drive increased tree mortality through drought, heat stress, and insect attacks, with manifold impacts on forest ecosystems. Yet, climate-induced tree mortality and biotic disturbance agents are largely absent from process-based ecosystem models. Using data sets from the western USA and associated studies, we present a framework for determining the relative contribution of drought stress, insect attack, and their interactions, which is critical for modeling mortality in future climates. We outline a simple approach that identifies the mechanisms associated with two guilds of insects - bark beetles and defoliators - which are responsible for substantial tree mortality. We then discuss cross-biome patterns of insect-driven tree mortality and draw upon available evidence contrasting the prevalence of insect outbreaks in temperate and tropical regions. We conclude with an overview of tools and promising avenues to address major challenges. Ultimately, a multitrophic approach that captures tree physiology, insect populations, and tree-insect interactions will better inform projections of forest ecosystem responses to climate change.

Patterns and causes of observed piñon pine mortality in the southwestern United States.

Recently, widespread piñon pine die-off occurred in the southwestern United States. Here we synthesize observational studies of this event and compare findings to expected relationships with biotic and abiotic factors. Agreement exists on the occurrence of drought, presence of bark beetles and increased mortality of larger trees. However, studies disagree about the influences of stem density, elevation and other factors, perhaps related to study design, location and impact of extreme drought. Detailed information about bark beetles is seldom reported and their role is poorly understood. Our analysis reveals substantial limits to our knowledge regarding the processes that produce mortality patterns across space and time, indicating a poor ability to forecast mortality in response to expected increases in future droughts.

Growth-mortality relationships in piñon pine (Pinus edulis) during severe droughts of the past century: shifting processes in space and time.

The processes leading to drought-associated tree mortality are poorly understood, particularly long-term predisposing factors, memory effects, and variability in mortality processes and thresholds in space and time. We use tree rings from four sites to investigate Pinus edulis mortality during two drought periods in the southwestern USA. We draw on recent sampling and archived collections to (1) analyze P. edulis growth patterns and mortality during the 1950s and 2000s droughts; (2) determine the influence of climate and competition on growth in trees that died and survived; and (3) derive regression models of growth-mortality risk and evaluate their performance across space and time. Recent growth was 53% higher in surviving vs. dying trees, with some sites exhibiting decades-long growth divergences associated with previous drought. Differential growth response to climate partly explained growth differences between live and dead trees, with responses wet/cool conditions most influencing eventual tree status. Competition constrained tree growth, and reduced trees' ability to respond to favorable climate. The best predictors in growth-mortality models included long-term (15-30 year) average growth rate combined with a metric of growth variability and the number of abrupt growth increases over 15 and 10 years, respectively. The most parsimonious models had high discriminatory power (ROC>0.84) and correctly classified ∼ 70% of trees, suggesting that aspects of tree growth, especially over decades, can be powerful predictors of widespread drought-associated die-off. However, model discrimination varied across sites and drought events. Weaker growth-mortality relationships and higher growth at lower survival probabilities for some sites during the 2000s event suggest a shift in mortality processes from longer-term growth-related constraints to shorter-term processes, such as rapid metabolic decline even in vigorous trees due to acute drought stress, and/or increases in the attack rate of both chronically stressed and more vigorous trees by bark beetles.

Drought predisposes piñon-juniper woodlands to insect attacks and mortality.

To test the hypothesis that drought predisposes trees to insect attacks, we quantified the effects of water availability on insect attacks, tree resistance mechanisms, and mortality of mature piñon pine (Pinus edulis) and one-seed juniper (Juniperus monosperma) using an experimental drought study in New Mexico, USA. The study had four replicated treatments (40 × 40 m plot/replicate): removal of 45% of ambient annual precipitation (H2 O-); irrigation to produce 125% of ambient annual precipitation (H2 O+); a drought control (C) to quantify the impact of the drought infrastructure; and ambient precipitation (A). Piñon began dying 1 yr after drought initiation, with higher mortality in the H2 O- treatment relative to other treatments. Beetles (bark/twig) were present in 92% of dead trees. Resin duct density and area were more strongly affected by treatments and more strongly associated with piñon mortality than direct measurements of resin flow. For juniper, treatments had no effect on insect resistance or attacks, but needle browning was highest in the H2 O- treatment. Our results provide strong evidence that ≥ 1 yr of severe drought predisposes piñon to insect attacks and increases mortality, whereas 3 yr of the same drought causes partial canopy loss in juniper.