Effects of biotic and abiotic factors on resistance versus resilience of Douglas fir to drought.

Significant increases in tree mortality due to drought-induced physiological stress have been documented worldwide. This trend is likely to continue with increased frequency and severity of extreme drought events in the future. Therefore, understanding the factors that influence variability in drought responses among trees will be critical to predicting ecosystem responses to climate change and developing effective management actions. In this study, we used hierarchical mixed-effects models to analyze drought responses of Pseudotsuga menziesii in 20 unmanaged forests stands across a broad range of environmental conditions in northeastern Washington, USA. We aimed to 1) identify the biotic and abiotic attributes most closely associated with the responses of individual trees to drought and 2) quantify the variability in drought responses at different spatial scales. We found that growth rates and competition for resources significantly affected resistance to a severe drought event in 2001: slow-growing trees and trees growing in subordinate canopy positions and/or with more neighbors suffered greater declines in radial growth during the drought event. In contrast, the ability of a tree to return to normal growth when climatic conditions improved (resilience) was unaffected by competition or relative growth rates. Drought responses were significantly influenced by tree age: older trees were more resistant but less resilient than younger trees. Finally, we found differences between resistance and resilience in spatial scale: a significant proportion (approximately 50%) of the variability in drought resistance across the study area was at broad spatial scales (i.e. among different forest types), most likely due to differences in the total amount of precipitation received at different elevations; in contrast, variation in resilience was overwhelmingly (82%) at the level of individual trees within stands and there was no difference in drought resilience among forest types. Our results suggest that for Pseudotsuga menziesii resistance and resilience to drought are driven by different factors and vary at different spatial scales.