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Abstract Structural overshoots, where biomass is overallocated to tree leaf area compared to sapwood area, could result in lethal stress during droughts. Climate change may alter climatic cues that drive leaf area production, such as temperature and precipitation, as well as seasonal dynamics that underlie summer rainfall due to the North American Monsoon (NAM). Combined, this could lead to temporal mismatches between leaf area‐driven water demand and availability, and increased drought‐induced mortality events.We used leaf area to sapwood area ratios to investigate the prevalence of overshoots and whether overshoots increase drought‐induced mortality. We measured populations of aspen spanning the northern transition zone of the NAM during and following severe droughts.We observed increased overshoots and drought‐induced mortality in southern latitude populations that rely more on summer monsoon rainfall. Changes in convective activity from low snowpack the preceding winter may be a climatic driver of heightened summer monsoon rainfall in the region and therefore may also trigger increased production of leaf area during wetter summers.Our results suggest that an overshoot of leaf area to sapwood area (A_L:A_S) ratios is associated with drought‐induced tree mortality and highlight that climate‐change driven alterations to the NAM could have major consequences for tree species' acclimation to environmental change. Read the freePlain Language Summaryfor this article on the Journal blog. 

Abstract Trees are arguably the most diverse and complex macro-organisms on Earth. The equally diverse functions of trees directly impact fluxes of carbon, water and energy from the land surface. A number of recent studies have shed light on the substantial within-species variability across plant traits, including aspects of leaf morphology and plant allocation of photosynthates to leaf biomass. Yet, within-tree variability in leaf traits due to microclimatic variations, leaf hydraulic coordination across traits at different physiological scales and variations in leaf traits over a growing season remain poorly studied. This knowledge gap is stymieing the fundamental understanding of what drives trait variation and covariation from tissues to trees to landscapes. Here, we present an extensive dataset measuring within-tree heterogeneity in leaf traits in California’s blue oak (Quercus douglasii) across an edaphic gradient and over the course of a growing season at an oak–grass savanna in Southern CA, USA. We found a high level of within-tree crown leaf area:sapwood area variation that was not attributable to sample height or aspect. We also found a higher level of trait integration at the tree level, rather than branch level, suggesting that trees optimize water use at the organismal level. Despite the large variance in traits within a tree crown and across trees, we did not find strong evidence for adaptive plasticity or acclimation in leaf morphological traits (e.g., changes to phenotype which increased fitness) across temporal and spatial water availability gradients. Collectively, our results highlight strong variation in drought-related physiology, but limited evidence for adaptive trait plasticity over shorter time scales. 

Summary Observational evidence indicates that tree leaf area may acclimate in response to changes in water availability to alleviate hydraulic stress. However, the underlying mechanisms driving leaf area changes and consequences of different leaf area allocation strategies remain unknown.Here, we use a trait‐based hydraulically enabled tree model with two endmember leaf area allocation strategies, aimed at either maximizing carbon gain or moderating hydraulic stress. We examined the impacts of these strategies on future plant stress and productivity.Allocating leaf area to maximize carbon gain increased productivity with high CO₂, but systematically increased hydraulic stress. Following an allocation strategy to avoid increased future hydraulic stress missed out on 26% of the potential future net primary productivity in some geographies. Both endmember leaf area allocation strategies resulted in leaf area decreases under future climate scenarios, contrary to Earth system model (ESM) predictions.Leaf area acclimation to avoid increased hydraulic stress (and potentially the risk of accelerated mortality) was possible, but led to reduced carbon gain. Accounting for plant hydraulic effects on canopy acclimation in ESMs could limit or reverse current projections of future increases in leaf area, with consequences for the carbon and water cycles, and surface energy budgets. 

Abstract Climate change is stressing many forests around the globe, yet some tree species may be able to persist through acclimation and adaptation to new environmental conditions. The ability of a tree to acclimate during its lifetime through changes in physiology and functional traits, defined here as its acclimation potential, is not well known.We investigated the acclimation potential of trembling aspenPopulus tremuloidesand ponderosa pinePinus ponderosatrees by examining within‐species variation in drought response functional traits across both space and time, and how trait variation influences drought‐induced tree mortality. We measured xylem tension, morphological traits and physiological traits on mature trees in southwestern Colorado, USA across a climate gradient that spanned the distribution limits of each species and 3 years with large differences in climate.Trembling aspen functional traits showed high within‐species variation, and osmotic adjustment and carbon isotope discrimination were key determinants for increased drought tolerance in dry sites and in dry years. However, trembling aspen trees at low elevation were pushed past their drought tolerance limit during the severe 2018 drought year, as elevated mortality occurred. Higher specific leaf area during drought was correlated with higher percentages of canopy dieback the following year. Ponderosa pine functional traits showed less within‐species variation, though osmotic adjustment was also a key mechanism for increased drought tolerance. Remarkably, almost all traits varied more year‐to‐year than across elevation in both species.Our results shed light on the scope and limits of intraspecific trait variation for mediating drought responses in key southwestern US tree species and will help improve our ability to model and predict forest responses to climate change. Read the freePlain Language Summaryfor this article on the Journal blog. 

Understanding the driving mechanisms behind existing patterns of vegetation hydraulic traits and community trait diversity is critical for advancing predictions of the terrestrial carbon cycle because hydraulic traits affect both ecosystem and Earth system responses to changing water availability. Here, we leverage an extensive trait database and a long-term continental forest plot network to map changes in community trait distributions and quantify “trait velocities” (the rate of change in community-weighted traits) for different regions and different forest types across the United States from 2000 to the present. We show that diversity in hydraulic traits and photosynthetic characteristics is more related to local water availability than overall species diversity. Finally, we find evidence for coordinated shifts toward communities with more drought-tolerant traits driven by tree mortality, but the magnitude of responses differs depending on forest type. The hydraulic trait distribution maps provide a publicly available platform to fundamentally advance understanding of community trait change in response to climate change and predictive abilities of mechanistic vegetation models. 

Abstract A central challenge in global change research is the projection of the future behavior of a system based upon past observations. Tree‐ring data have been used increasingly over the last decade to project tree growth and forest ecosystem vulnerability under future climate conditions. But how can the response of tree growth to past climate variation predict the future, when the future does not look like the past? Space‐for‐time substitution (SFTS) is one way to overcome the problem of extrapolation: the response at a given location in a warmer future is assumed to follow the response at a warmer location today. Here we evaluated an SFTS approach to projecting future growth of Douglas‐fir (Pseudotsuga menziesii), a species that occupies an exceptionally large environmental space in North America. We fit a hierarchical mixed‐effects model to capture ring‐width variability in response to spatial and temporal variation in climate. We found opposing gradients for productivity and climate sensitivity with highest growth rates and weakest response to interannual climate variation in the mesic coastal part of Douglas‐fir's range; narrower rings and stronger climate sensitivity occurred across the semi‐arid interior. Ring‐width response to spatial versus temporal temperature variation was opposite in sign, suggesting that spatial variation in productivity, caused by local adaptation and other slow processes, cannot be used to anticipate changes in productivity caused by rapid climate change. We thus substituted only climate sensitivities when projecting future tree growth. Growth declines were projected across much of Douglas‐fir's distribution, with largest relative decreases in the semiarid U.S. Interior West and smallest in the mesic Pacific Northwest. We further highlight the strengths of mixed‐effects modeling for reviving a conceptual cornerstone of dendroecology, Cook's 1987 aggregate growth model, and the great potential to use tree‐ring networks and results as a calibration target for next‐generation vegetation models.