Interactions between Climate and Stand Conditions Predict Pine Mortality during a Bark Beetle Outbreak

In temperate coniferous forests, biotic disturbances such as bark beetle outbreaks can result in widespread tree mortality. The characteristics of individual trees and stands, such as tree diameter and stand density, often influence the probability of tree mortality during a bark beetle outbreak. However, it is unclear if these relationships are mediated by climate. To test this, we assembled tree mortality data for over 3800 ponderosa pine trees from Forest Inventory and Analysis (FIA) plots measured before and after a mountain pine beetle outbreak in the Black Hills, South Dakota, USA. Logistic models were used to determine which tree, stand, and climate characteristics were associated with the probability of mortality. Interactions were tested between significant climate variables and significant tree/stand variables. Our analysis revealed that mortality rates were lower in trees with higher live crown ratios. Mortality rates rose in response to increasing tree diameter, stand basal area (both from ponderosa pine and non-ponderosa pine), and elevation. Below 1500 m, the mortality rate was ~1%, while above 1700 m, the rate increased to ~30%. However, the association between elevation and mortality risk was buffered by precipitation, such that relatively moist high-elevation stands experienced less mortality than relatively dry high-elevation stands. Tree diameter, crown ratio, and stand density affected tree mortality independent of precipitation. This study demonstrates that while stand characteristics affect tree susceptibility to bark beetles, these relationships may be mediated by climate. Thus, both site and stand level characteristics should be considered when implementing management treatments to reduce bark beetle susceptibility.