At least it is a dry cold: the global distribution of freeze-thaw and drought stress and the traits that may impart poly-tolerance in conifers.

Conifers inhabit some of the most challenging landscapes where multiple abiotic stressors (e.g., aridity, freezing temperatures) often co-occur. Physiological tolerance to multiple stressors ('poly-tolerance') is thought to be rare because exposure to one stress generally limits responses to another through functional trade-offs. However, the capacity to exhibit poly-tolerance may be greater when combined abiotic stressors have similar physiological impacts, such as the disruption of hydraulic function imposed by drought or freezing. Here, we reviewed empirical data in light of theoretical expectations for conifer adaptations to drought and freeze-thaw cycles with particular attention to hydraulic traits of the stem and leaf. Additionally, we examined the commonality and spatial distribution of poly-stress along indices of these combined stressors. We found that locations with the highest values of our poly-stress index (PSi) are characterized by moderate drought and moderate freeze-thaw, and most of the global conifer distribution occupies areas of moderate poly-stress. Among traits examined, we found diverse responses to the stressors. Turgor loss point did not correlate with freeze-thaw or drought stress individually, but did with the PSi, albeit inverse to what was hypothesized. Leaf mass per area was more strongly linked with drought stress than the poly-stress and not at all with freeze-thaw stress. In stems, the water potential causing 50% loss of hydraulic conductivity became more negative with increasing drought stress and poly-stress but did not correlate with freeze-thaw stress. For these traits, we identified a striking lack of coverage for substantial portions of species ranges, particularly at the upper boundaries of their respective PSis, demonstrating a critical gap in our understanding of trait prevalence and plasticity along these stress gradients. Future research should investigate traits that confer tolerance to both freeze-thaw and drought stress in a wide range of species across broad geographic scales.

Poor acclimation to experimental field drought in subalpine forest tree seedlings.

The ability of tree species to acclimate and tolerate projected increases in drought frequency and intensity has fundamental implications for future forest dynamics with climate change. Inquiries to date on the drought tolerance capacities of tree species, however, have focused almost exclusively on mature trees with scant in situ work on seedlings, despite the central role that regeneration dynamics play in forest responses to changing conditions. We subjected naturally established seedlings of co-dominant subalpine conifer species (Abies lasiocarpa and Picea engelmannii) in the southern Rocky Mountains to 2 years of in situ summer precipitation exclusion, simulating summer drought conditions similar to a failure of the North American monsoon. We compared the morphological and physiological responses of seedlings growing in drought vs. ambient conditions to assess the relative changes in drought tolerance traits as a function of seedling size. Drought treatments had a marked impact on soil moisture: volumetric water content averaged ≈5-8 % in drought treatments and ≈8-12 % in ambient controls. We detected no significant shifts in morphology (e.g. root biomass, leaf:stem area ratio) in response to drought for either species, but net photosynthesis in drought treatments was 78 % lower for spruce and 37 % lower for fir. Greater stomatal control associated with increasing stem diameter conferred greater water use efficiencies in larger seedlings in both species but was not significantly different between drought and ambient conditions, suggesting an overall lack of responsivity to water stress and a prioritization of carbon gain over investment in drought mitigation traits. These results indicate a canonization of traits that, while useful for early seedling establishment, may portend substantial vulnerability of subalpine seedling populations to prolonged or recurrent droughts, especially for spruce.