Drought-Associated Tree Mortality: Global patterns and insights from tree-ring studies in the Southwestern U.S.A.

Forests play an important role in the earth system, regulating climate, maintaining biodiversity, and provisioning human communities with water, food and fuel. Interactions between climate and forest dynamics are not well constrained, and high uncertainty characterizes projections of global warming impacts on forests and associated ecosystem services. Recently observed tree mortality and forest die-off forewarn an acceleration of forest change with rising temperature and increased drought. However, the processes leading to tree death during drought are poorly understood, limiting our ability to anticipate future forest dynamics.The objective of this dissertation was to improve understanding of drought-associated tree mortality through literature synthesis and tree-ring studies on trees that survived and died during droughts in the southwestern USA. Specifically, this dissertation, 1) documented global tree mortality patterns and identified emerging trends and research gaps; 2) quantified relationships between growth, climate, competition and mortality of piñon pine during droughts in New Mexico; 3) investigated tree defense anatomy as a potentially key element in piñon avoidance of death; and, 4) characterized the climate sensitivity of piñon resin ducts in order to gain insight into potential trends in tree defenses with climate variability and change.There has been an increase in studies reporting tree mortality linked to drought, heat, and the associated activity of insects and pathogens. Cases span the forested continents and occurred in water, light and temperature-limited forests. We hypothesized that increased tree mortality may be an emerging global phenomenon related to rising temperatures and drought. Recent radial growth was 53% higher on average in piñon that survived versus died during two episodes of drought-associated mortality, and statistical models of mortality risk based average growth, growth variability, and abrupt growth changes correctly classified the status of ~70% of trees. Climate responses and competitive interactions partly explained growth differences between dying and surviving trees, with muted response to wet/cool conditions and enhanced sensitivity to competition from congeners linked to growth patterns associated with death. Discrimination and validation of models of mortality risk varied widely across sites and drought events, indicating shifting growth-mortality relationships and differences in mortality processes across space and time. Pre-formed defense anatomy is strongly associated with piñon survivorship over a range of sites and stand conditions. Models of mortality risk that account for both growth and resin duct attributes had ≈1019 more support than models that contained only growth. The greatest improvement in classification was among trees from the 2000s drought, suggesting an enhanced role for tree defense allocation and/or bark beetle activity during recent warm versus historic cool drought. Accounting for defense characteristics and growth-defense allocation is likely to be important for improving representation of drought-associated mortality. Piñon resin duct chronologies contain climate responses that are coherent and distinct from those of radial growth. Growth responds positively and strongly to previous fall and current winter precipitation, and negatively to late spring and early summer temperature. A relatively equal positive resin duct response to winter precipitation and positive response to mid-to-late summer drought suggests that changes in climate will affect tree defense anatomy in complex ways, with the outcome determined by seasonal changes in precipitation and temperature (Appendix D).