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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. 

Synopsis Classic debates in community ecology focused on the complexities of considering an ecosystem as a super-organ or organism. New consideration of such perspectives could clarify mechanisms underlying the dynamics of forest carbon dioxide (CO2) uptake and water vapor loss, important for predicting and managing the future of Earth’s ecosystems and climate system. Here, we provide a rubric for considering ecosystem traits as aggregated, systemic, or emergent, i.e., representing the ecosystem as an aggregate of its individuals or as a metaphorical or literal super-organ or organism. We review recent approaches to scaling-up plant water relations (hydraulics) concepts developed for organs and organisms to enable and interpret measurements at ecosystem-level. We focus on three community-scale versions of water relations traits that have potential to provide mechanistic insight into climate change responses of forest CO2 and H2O gas exchange and productivity: leaf water potential (Ψcanopy), pressure volume curves (eco-PV), and hydraulic conductance (Keco). These analyses can reveal additional ecosystem-scale parameters analogous to those typically quantified for leaves or plants (e.g., wilting point and hydraulic vulnerability) that may act as thresholds in forest responses to drought, including growth cessation, mortality, and flammability. We unite these concepts in a novel framework to predict Ψcanopy and its approaching of critical thresholds during drought, using measurements of Keco and eco-PV curves. We thus delineate how the extension of water relations concepts from organ- and organism-scales can reveal the hydraulic constraints on the interaction of vegetation and climate and provide new mechanistic understanding and prediction of forest water use and productivity. 

Abstract Forests are a large carbon sink and could serve as natural climate solutions that help moderate future warming. Thus, establishing forest carbon baselines is essential for tracking climate‐mitigation targets. Western US forests are natural climate solution hotspots but are profoundly threatened by drought and altered disturbance regimes. How these factors shape spatial patterns of carbon storage and carbon change over time is poorly resolved. Here, we estimate live and dead forest carbon density in 19 forested western US ecoregions with national inventory data (2005–2019) to determine: (a) current carbon distributions, (b) underpinning drivers, and (c) recent trends. Potential drivers of current carbon included harvest, wildfire, insect and disease, topography, and climate. Using random forests, we evaluated driver importance and relationships with current live and dead carbon within ecoregions. We assessed trends using linear models. Pacific Northwest (PNW) and Southwest (SW) ecoregions were most and least carbon dense, respectively. Climate was an important carbon driver in the SW and Lower Rockies. Fire reduced live and increased dead carbon, and was most important in the Upper Rockies and California. No ecoregion was unaffected by fire. Harvest and private ownership reduced carbon, particularly in the PNW. Since 2005, live carbon declined across much of the western US, likely from drought and fire. Carbon has increased in PNW ecoregions, likely recovering from past harvest, but recent record fire years may alter trajectories. Our results provide insight into western US forest carbon function and future vulnerabilities, which is vital for effective climate change mitigation strategies. 

Abstract Spatiotemporal patterns of plant water uptake, loss, and storage exert a first‐order control on photosynthesis and evapotranspiration. Many studies of plant responses to water stress have focused on differences between species because of their different stomatal closure, xylem conductance, and root traits. However, several other ecohydrological factors are also relevant, including soil hydraulics, topographically driven redistribution of water, plant adaptation to local climatic variations, and changes in vegetation density. Here, we seek to understand the relative importance of the dominant species for regional‐scale variations in woody plant responses to water stress. We map plant water sensitivity (PWS) based on the response of remotely sensed live fuel moisture content to variations in hydrometeorology using an auto‐regressive model. Live fuel moisture content dynamics are informative of PWS because they directly reflect vegetation water content and therefore patterns of plant water uptake and evapotranspiration. The PWS is studied using 21,455 wooded locations containing U.S. Forest Service Forest Inventory and Analysis plots across the western United States, where species cover is known and where a single species is locally dominant. Using a species‐specific mean PWS value explains 23% of observed PWS variability. By contrast, a random forest driven by mean vegetation density, mean climate, soil properties, and topographic descriptors explains 43% of observed PWS variability. Thus, the dominant species explains only 53% (23% compared to 43%) of explainable variations in PWS. Mean climate and mean NDVI also exert significant influence on PWS. Our results suggest that studies of differences between species should explicitly consider the environments (climate, soil, topography) in which observations for each species are made, and whether those environments are representative of the entire species range. 

Abstract Forest fire frequency, extent, and severity have rapidly increased in recent decades across the western United States (US) due to climate change and suppression‐oriented wildfire management. Fuels reduction treatments are an increasingly popular management tool, as evidenced by California's plan to treat 1 million acres annually by 2050. However, the aggregate efficacy of fuels treatments in dry forests at regional and multi‐decadal scales is unknown. We develop a novel fuels treatment module within a coupled dynamic vegetation and fire model to study the effects of dead biomass removal from forests in the Sierra Nevada region of California. We ask how annual treatment extent, stand‐level treatment intensiveness, and spatial treatment placement alter fire severity and live carbon loss. We find that a ∼30% reduction in stand‐replacing fire was achieved under our baseline treatment scenario of 1,000 km² year⁻¹after a 100‐year treatment period. Prioritizing the most fuel‐heavy stands based on precise fuel distributions yielded cumulative reductions in pyrogenic stand‐replacement of up to 50%. Both removing constraints on treatment location due to remoteness, topography, and management jurisdiction and prioritizing the most fuel‐heavy stands yielded the highest stand‐replacement rate reduction of ∼90%. Even treatments that succeeded in lowering aggregate fire severity often took multiple decades to yield measurable effects, and avoided live carbon loss remained negligible across scenarios. Our results suggest that strategically placed fuels treatments are a promising tool for controlling forest fire severity at regional, multi‐decadal scales, but may be less effective for mitigating live carbon losses. 

Summary Predictive relationships between plant traits and environmental factors can be derived at global and regional scales, informing efforts to reorient ecological models around functional traits. However, in a changing climate, the environmental variables used as predictors in such relationships are far from stationary. This could yield errors in trait–environment model predictions if timescale is not accounted for.Here, the timescale dependence of trait–environment relationships is investigated by regressingin situtrait measurements of specific leaf area, leaf nitrogen content, and wood density on local climate characteristics summarized across several increasingly long timescales.We identify contrasting responses of leaf and wood traits to climate timescale. Leaf traits are best predicted by recent climate timescales, while wood density is a longer term memory trait. The use of sub‐optimal climate timescales reduces the accuracy of the resulting trait–environment relationships.This study concludes that plant traits respond to climate conditions on the timescale of tissue lifespans rather than long‐term climate normals, even at large spatial scales where multiple ecological and physiological mechanisms drive trait change. Thus, determining trait–environment relationships with temporally relevant climate variables may be critical for predicting trait change in a nonstationary climate system. 

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. 

Abstract In semi‐arid regions where drought and wildfire events often co‐occur, such as in Southern California chaparral, relationships between plant hydration, drought‐ and fire‐adapted traits may explain landscape‐scale wildfire dynamics. To examine these patterns, fire scientists and plant physiologists quantify hydration in plants via mass‐based metrics of water content, including live fuel moisture, or pressure‐based metrics of physiological status, such as xylem water potential; however, relationships across these metrics, plant traits and flammability remain unresolved.To determine the impact of hydration on tissue‐level flammability (leaves and stems), we conducted laboratory dehydration tests across wet and dry seasons in which we simultaneously measured xylem water potential, live fuel moisture and flammability. We tested two widespread chaparral shrubs,Adenostoma fasciculatumandCeanothus megacarpus.Live fuel moisture showed a threshold‐type relationship with tissue flammability (increased ignitability and combustibility at specific hydration levels) that aligned with drought‐response traits (turgor loss point) and fire behaviour (increased fire likelihood and spread) identified at the landscape scale. Water potential was the better predictor of flammability in linear statistical models.A. fasciculatumwas more flammable thanC. megacarpus, and both species were more flammable during the wet growing season, suggesting seasonal growth or drought‐related tissue characteristics other than moisture content, such as lignin or chemical content, are critical for determining flammability.Our results suggest a mechanism for landscape‐scale increases in flammability at specific levels of drought stress. Integration of drought‐related traits, such as the turgor loss point, might improve models of wildfire risk in drought‐ and fire‐prone systems. Read the freePlain Language Summaryfor this article on the Journal blog. 

Abstract The extent to which future climate change will increase forest stress and the amount to which species and forest ecosystems can acclimate or adapt to increased stress is a major unknown. We used high‐resolution maps of hydraulic traits representing the diversity in tree drought tolerance across the United States, a hydraulically enabled tree model, and forest inventory observations of demographic shifts to quantify the ability for within‐species acclimation and between‐species range shifts to mediate climate stress. We found that forests are likely to experience increases in both acute and chronic hydraulic stress with climate change. Based on current species distributions, regional hydraulic trait diversity was sufficient to buffer against increased stress in 88% of forested areas. However, observed trait velocities in 81% of forested areas are not keeping up with the rate required to ameliorate projected future stress without leaf area acclimation. 

Abstract Forests sequester ∼25% of anthropogenic carbon (C) emissions annually and are of increasing interest for their potential as Nature‐based Climate Solutions (NbCS). Emergent from the need to assess terrestrial ecosystem health and quantify C storage and fluxes, several gridded products documenting terrestrial C and changes in C stocks over time have been developed. However, researchers have not yet systematically compared C distributions across products, or developed a clear path forward for investigating and leveraging this cross‐product uncertainty in estimates of terrestrial C. Alaniz et al. (2022,https://doi.org/10.1029/2021EF002560) synthesize multiple published products to constrain the distribution of forest C stocks and fluxes globally. Building off of their results, we comment on opportunities for advancing both basic science and NbCS policy recommendations through systematic product cross‐comparisons and targeting of areas with differing levels of uncertainties in the terrestrial C sink.