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Summary: Some plants exhibit dynamic hydraulic regulation, in which the strictness of hydraulic regulation (i.e. iso/anisohydry) changes in response to environmental conditions. However, the environmental controls over iso/anisohydry and the implications of flexible hydraulic regulation for plant productivity remain unknown.In Juniperus osteosperma, a drought‐resistant dryland conifer, we collected a 5‐month growing season time series of in situ, high temporal‐resolution plant water potential (Ψ) and stand gross primary productivity (GPP). We quantified the stringency of hydraulic regulation associated with environmental covariates and evaluated how predawn water potential contributes to empirically predicting carbon uptake.Juniperus osteosperma showed less stringent hydraulic regulation (more anisohydric) after monsoon precipitation pulses, when soil moisture and atmospheric demand were high, and corresponded with GPP pulses. Predawn water potential matched the timing of GPP fluxes and improved estimates of GPP more strongly than soil and/or atmospheric moisture, notably resolving GPP underestimation before vegetation green‐up.Flexible hydraulic regulation appears to allow J. osteosperma to prolong soil water extraction and, therefore, the period of high carbon uptake following monsoon precipitation pulses. Water potential and its dynamic regulation may account for why process‐based and empirical models commonly underestimate the magnitude and temporal variability of dryland GPP. [ABSTRACT FROM AUTHOR]

Metrics to quantify regulation of plant water status at the daily as opposed to the seasonal scale do not presently exist. This gap is significant since plants are hypothesised to regulate their water potential not only with respect to slowly changing soil drought but also with respect to faster changes in air vapour pressure deficit (VPD), a variable whose importance for plant physiology is expected to grow because of higher temperatures in the coming decades. We present a metric, the stringency of water potential regulation, that can be employed at the daily scale and quantifies the effects exerted on plants by the separate and combined effect of soil and atmospheric drought. We test our theory using datasets from two experiments where air temperature and VPD were experimentally manipulated. In contrast to existing metrics based on soil drought that can only be applied at the seasonal scale, our metric successfully detects the impact of atmospheric warming on the regulation of plant water status. We show that the thermodynamic effect of VPD on plant water status can be isolated and compared against that exerted by soil drought and the covariation between VPD and soil drought. Furthermore, in three of three cases, VPD accounted for more than 5 MPa of potential effect on leaf water potential. We explore the significance of our findings in the context of potential future applications of this metric from plant to ecosystem scale. [ABSTRACT FROM AUTHOR]

Climate change‐triggered forest die‐off is an increasing threat to global forests and carbon sequestration but remains extremely challenging to predict. Tree growth resilience metrics have been proposed as measurable proxies of tree susceptibility to mortality. However, it remains unclear whether tree growth resilience can improve predictions of stand‐level mortality. Here, we use an extensive tree‐ring dataset collected at ~3000 permanent forest inventory plots, spanning 13 dominant species across the US Mountain West, where forests have experienced strong drought and extensive die‐off has been observed in the past two decades, to test the hypothesis that tree growth resilience to drought can explain and improve predictions of observed stand‐level mortality. We found substantial increases in growth variability and temporal autocorrelation as well declining drought resistance and resilience for a number of species over the second half of the 20th century. Declining resilience and low tree growth were strongly associated with cross‐ and within‐species patterns of mortality. Resilience metrics had similar explicative power compared to climate and stand structure, but the covariance structure among predictors implied that the effect of tree resilience on mortality could partially be explained by stand and climate variables. We conclude that tree growth resilience offers highly valuable insights on tree physiology by integrating the effect of stressors on forest mortality but may have only moderate potential to improve large‐scale projections of forest die‐off under climate change. [ABSTRACT FROM AUTHOR]

Summary: Intraspecific variation in functional traits may mediate tree species' drought resistance, yet whether trait variation is due to genotype (G), environment (E), or G×E interactions remains unknown. Understanding the drivers of intraspecific trait variation and whether variation mediates drought response can improve predictions of species' response to future drought.Using populations of quaking aspen spanning a climate gradient, we investigated intraspecific variation in functional traits in the field as well as the influence of G and E among propagules in a common garden. We also tested for trait‐mediated trade‐offs in growth and drought stress tolerance.We observed intraspecific trait variation among the populations, yet this variation did not necessarily translate to higher drought stress tolerance in hotter/drier populations. Additionally, plasticity in the common garden was low, especially in propagules derived from the hottest/driest population. We found no growth–drought stress tolerance trade‐offs and few traits exhibited significant relationships with mortality in the natural populations, suggesting that intraspecific trait variation among the traits measured did not strongly mediate responses to drought stress.Our results highlight the limits of trait‐mediated responses to drought stress and the complex G×E interactions that may underlie drought stress tolerance variation in forests in dry environments. [ABSTRACT FROM AUTHOR]

Episodes of forest mortality have been observed worldwide associated with climate change, impacting species composition and ecosystem services such as water resources and carbon sequestration. Yet our ability to predict forest mortality remains limited, especially across large scales. Time series of satellite imagery has been used to document ecosystem resilience globally, but it is not clear how well remotely sensed resilience can inform the prediction of forest mortality across continental, multi-biome scales. Here, we leverage forest inventories across the continental United States to systematically assess the potential of ecosystem resilience derived using different data sets and methods to predict forest mortality. We found high resilience was associated with low mortality in eastern forests but was associated with high mortality in western regions. The unexpected resilience–mortality relation in western United States may be due to several factors including plant trait acclimation, insect population dynamics, or resource competition. Overall, our results not only supported the opportunity to use remotely sensed ecosystem resilience to predict forest mortality but also highlighted that ecological factors may have crucial influences because they can reverse the sign of the resilience–mortality relationships. [ABSTRACT FROM AUTHOR]

Forest productivity projections remain highly uncertain, notably because underpinning physiological controls are delicate to disentangle. Transient perturbation of global climate by large volcanic eruptions provides a unique opportunity to retrospectively isolate underlying processes. Here, we use a multi‐proxy dataset of tree‐ring records distributed over the Northern Hemisphere to investigate the effect of eruptions on tree growth and photosynthesis and evaluate CMIP6 models. Tree‐ring isotope records denoted a widespread 2–4 years increase of photosynthesis following eruptions, likely as a result of diffuse light fertilization. We found evidence that enhanced photosynthesis transiently drove ring width, but the latter further exhibited a decadal anomaly that evidenced independent growth and photosynthesis responses. CMIP6 simulations reproduced overall tree growth decline but did not capture observed photosynthesis anomaly, its decoupling from tree growth or the climate sensitivities of either processes, highlighting key disconnects that deserve further attention to improve forest productivity projections under climate change. [ABSTRACT FROM AUTHOR]

Trees are long‐lived organisms, exhibiting temporally complex growth arising from strong climatic "memory." But conditions are becoming increasingly arid in the western USA. Using a century‐long tree‐ring network, we find altered climate memory across the entire range of a widespread western US conifer: growth is supported by precipitation falling further into the past (+15 months), while increasingly impacted by more recent temperature conditions (−8 months). Tree‐ring datasets can be biased, so we confirm altered climate memory in a second, ecologically‐sampled tree‐ring network. Predicted drought responses show trees may have also become more sensitive to repeat drought. Finally, plots near sites with relatively longer precipitation memory and shorter temperature memory had significantly lower recent mortality rates (R2 = 0.61). We argue that increased drought frequency has altered climate memory, demonstrate how non‐stationarity may arise from failure to account for memory, and suggest memory length may be predictive of future tree mortality. [ABSTRACT FROM AUTHOR]

Nature‐based climate solutions are a vital component of many climate mitigation strategies, including California's, which aims to achieve carbon neutrality by 2045. Most carbon offsets in California's cap‐and‐trade program come from improved forest management (IFM) projects. Since 2012, various landowners have set up IFM projects following the California Air Resources Board's IFM protocol. As many of these projects approach their 10th year, we now have the opportunity to assess their effectiveness, identify best practices, and suggest improvements toward future protocol revisions. In this study, we used remote sensing‐based datasets to evaluate the carbon trends and harvest histories of 37 IFM projects in California. Despite some current limitations and biases, these datasets can be used to quantify carbon accumulation and harvest rates in offset project lands relative to nearby similar "control" lands before and after the projects began. Five lines of evidence suggest that the carbon accumulated in offset projects to date has generally not been additional to what might have otherwise occurred: (1) most forests in northwestern California have been accumulating carbon since at least the mid‐1980s and continue to accumulate carbon, whether enrolled in offset projects or not; (2) harvest rates were high in large timber company project lands before IFM initiation, suggesting they are earning carbon credits for forests in recovery; (3) projects are often located on lands with higher densities of low‐timber‐value species; (4) carbon accumulation rates have not yet increased on lands that enroll as offset projects, relative to their pre‐enrollment levels; and (5) harvest rates have not decreased on most project lands since offset project initiation. These patterns suggest that the current protocol should be improved to robustly measure and reward additionality. In general, our framework of geospatial analyses offers an important and independent means to evaluate the effectiveness of the carbon offsets program, especially as these data products continue improving and as offsets receive attention as a climate mitigation strategy. [ABSTRACT FROM AUTHOR]

Mechanistic representations of biogeochemical processes in ecosystem models are rapidly advancing, requiring advancements in model evaluation approaches. Here we quantify multiple aspects of model functional performance to evaluate improved process representations in ecosystem models. We compare semi‐empirical stomatal models with hydraulic constraints against more mechanistic representations of stomatal and hydraulic functioning at a semi‐arid pine site using a suite of metrics and analytical tools. We find that models generally perform similarly under unstressed conditions, but performance diverges under atmospheric and soil drought. The more empirical models better capture synergistic information flows between soil water potential and vapor pressure deficit to transpiration, while the more mechanistic models are overly deterministic. Although models can be parameterized to yield similar functional performance, alternate parameterizations could not overcome structural model constraints that underestimate the unique information contained in soil water potential about transpiration. Additionally, both multilayer canopy and big‐leaf models were unable to capture the magnitude of canopy temperature divergence from air temperature, and we demonstrate that errors in leaf temperature can propagate to considerable error in simulated transpiration. This study demonstrates the value of merging underutilized observational data streams with emerging analytical tools to characterize ecosystem function and discriminate among model process representations. Plain Language Summary: Earth system models are an essential tool for understanding the consequences of changing climate conditions on forest ecosystems. Models are rapidly incorporating more realistic representations of how drought impacts ecosystem carbon and water cycling. These advancements need to be thoroughly evaluated to ensure that the models adequately capture the plant functional response to drought stress. Here we merge underutilized measurements with new analytical tools to evaluate several model representations of plant response to drought. These tools allow us to both better understand relationships among drought stress and ecosystem response, as well as quantify model accuracy. We find that models generally perform similarly under unstressed conditions, but performance diverges under drought. Key Points: We evaluate several model formulations for coupling plant hydraulic and stomatal response using functional performance metricsInformation flows from soil water potential and vapor pressure deficit to transpiration reflect structural differences among modelsConsiderable biases in modeled canopy temperature propagate to a 5% offset in cumulative growing season transpiration [ABSTRACT FROM AUTHOR]

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 aspen Populus tremuloides and ponderosa pine Pinus ponderosa trees 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 free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Tree species display a wide variety of water-use strategies, growth rates and capacity to tolerate drought. However, if we want to forecast species capacity to cope with increasing aridity and drought, we need to identify which measurable traits confer resilience to drought across species. Here, we use a global tree ring network (65 species; 1931 site series of ring-width indices--RWI) to evaluate the relationship of long-term growth-drought sensitivity (RWI-SPEI drought index relationship) and short-term growth response to extreme drought episodes (resistance, recovery and resilience indices) with functional traits related to leaf, wood and hydraulic properties. Furthermore, we assess the influence of climate (temperature, precipitation and climatic water deficit) on these trait-growth relationships. We found a close correspondence between the long-term relationship between RWI and SPEI and resistance and recovery of tree growth to severe drought episodes. Species displaying a stronger RWI-SPEI relationship to drought and low resistance and high recovery to extreme drought episodes tended to have a higher wood density (WD) and more negative leaf minimum water potential (Ψmin). Such associations were largely maintained when accounting for direct climate effects. Our results indicate that, at a cross-species level and global scale, wood and hydraulic functional traits explain species' growth responses to drought at short-and long-term scales. These trait-growth response relationships can improve our understanding of the cross-species capacity to withstand climate change and inform models to better predict drought effects on forest ecosystem dynamics. [ABSTRACT FROM AUTHOR]

Nature-based Climate Solutions (NbCS) are managed alterations to ecosystems designed to increase carbon sequestration or reduce greenhouse gas emissions. While they have growing public and private support, the realizable benefits and unintended consequences of NbCS are not well understood. At regional scales where policy decisions are often made, NbCS benefits are estimated from soil and tree survey data that can miss important carbon sources and sinks within an ecosystem, and do not reveal the biophysical impacts of NbCS for local water and energy cycles. The only direct observations of ecosystem-scale carbon fluxes, for example, by eddy covariance flux towers, have not yet been systematically assessed for what they can tell us about NbCS potentials, and state-of-the-art remote sensing products and land-surface models are not yet being widely used to inform NbCS policymaking or implementation. As a result, there is a critical mismatch between the point-and tree-scale data most often used to assess NbCS benefits and impacts, the ecosystem and landscape scales where NbCS projects are implemented, and the regional to continental scales most relevant to policymaking. Here, we propose a research agenda to confront these gaps using data and tools that have long been used to understand the mechanisms driving ecosystem carbon and energy cycling, but have not yet been widely applied to NbCS. We outline steps for creating robust NbCS assessments at both local to regional scales that are informed by ecosystem-scale observations, and which consider concurrent biophysical impacts, future climate feedbacks, and the need for equitable and inclusive NbCS implementation strategies. We contend that these research goals can largely be accomplished by shifting the scales at which pre-existing tools are applied and blended together, although we also highlight some opportunities for more radical shifts in approach. [ABSTRACT FROM AUTHOR]

Forests are currently a substantial carbon sink globally. Many climate change mitigation strategies leverage forest preservation and expansion, but rely on forests storing carbon for decades to centuries. Yet climate‐driven disturbances pose critical risks to the long‐term stability of forest carbon. We quantify the climate drivers that influence wildfire and climate stress‐driven tree mortality, including a separate insect‐driven tree mortality, for the contiguous United States for current (1984–2018) and project these future disturbance risks over the 21st century. We find that current risks are widespread and projected to increase across different emissions scenarios by a factor of >4 for fire and >1.3 for climate‐stress mortality. These forest disturbance risks highlight pervasive climate‐sensitive disturbance impacts on US forests and raise questions about the risk management approach taken by forest carbon offset policies. Our results provide US‐wide risk maps of key climate‐sensitive disturbances for improving carbon cycle modeling, conservation and climate policy. [ABSTRACT FROM AUTHOR]

Carbon offsets are widely used by individuals, corporations, and governments to mitigate their greenhouse gas emissions on the assumption that offsets reflect equivalent climate benefits achieved elsewhere. These climate‐equivalence claims depend on offsets providing real and additional climate benefits beyond what would have happened, counterfactually, without the offsets project. Here, we evaluate the design of California's prominent forest carbon offsets program and demonstrate that its climate‐equivalence claims fall far short on the basis of directly observable evidence. By design, California's program awards large volumes of offset credits to forest projects with carbon stocks that exceed regional averages. This paradigm allows for adverse selection, which could occur if project developers preferentially select forests that are ecologically distinct from unrepresentative regional averages. By digitizing and analyzing comprehensive offset project records alongside detailed forest inventory data, we provide direct evidence that comparing projects against coarse regional carbon averages has led to systematic over‐crediting of 30.0 million tCO2e (90% CI: 20.5–38.6 million tCO2e) or 29.4% of the credits we analyzed (90% CI: 20.1%–37.8%). These excess credits are worth an estimated $410 million (90% CI: $280–$528 million) at recent market prices. Rather than improve forest management to store additional carbon, California's forest offsets program creates incentives to generate offset credits that do not reflect real climate benefits. [ABSTRACT FROM AUTHOR]

Classifying the diverse ways that plants respond to hydrologic stress into generalizable 'water‐use strategies' has long been an eco‐physiological research goal. While many schemes for describing water‐use strategies have proven to be quite useful, they are also associated with uncertainties regarding their theoretical basis and their connection to plant carbon and water relations. In this review, we discuss the factors that shape plant water stress responses and assess the approaches used to classify a plant's water‐use strategy, paying particular attention to the popular but controversial concept of a continuum from isohydry to anisohydry.A generalizable and predictive framework for assessing plant water‐use strategies has been historically elusive, yet recent advances in plant physiology and hydraulics provide the field with a way past these obstacles. Specifically, we promote the idea that many metrics that quantify water‐use strategies are highly dynamic and emergent from the interaction between plant traits and environmental conditions, and that this complexity has historically hindered the development of a generalizable water‐use strategy framework.This idea is explored using a plant hydraulics model to identify: (a) distinct temporal phases in plant hydraulic regulation during drought that underpin dynamic water‐use responses, and (b) how variation in both traits and environmental forcings can significantly alter common metrics used to characterize plant water‐use strategies. This modelling exercise can bridge the divide between various conceptualizations of water‐use strategies and provide targeted hypotheses to advance the understanding and quantification of plant water status regulation across spatial and temporal scales.Finally, we describe research frontiers that are necessary to improve the predictive capacity of the plant water‐use strategy concept, including further investigation into the below‐ground determinants of plant water relations, targeted data collection efforts and the potential to scale these concepts from individuals to whole regions. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Droughts in a warming climate have become more common and more extreme, making understanding forest responses to water stress increasingly pressing. Analysis of water stress in trees has long focused on water potential in xylem and leaves, which influences stomatal closure and water flow through the soil‐plant‐atmosphere continuum. At the same time, changes of vegetation water content (VWC) are linked to a range of tree responses, including fluxes of water and carbon, mortality, flammability, and more. Unlike water potential, which requires demanding in situ measurements, VWC can be retrieved from remote sensing measurements, particularly at microwave frequencies using radar and radiometry. Here, we highlight key frontiers through which VWC has the potential to significantly increase our understanding of forest responses to water stress. To validate remote sensing observations of VWC at landscape scale and to better relate them to data assimilation model parameters, we introduce an ecosystem‐scale analog of the pressure–volume curve, the non‐linear relationship between average leaf or branch water potential and water content commonly used in plant hydraulics. The sources of variability in these ecosystem‐scale pressure‐volume curves and their relationship to forest response to water stress are discussed. We further show to what extent diel, seasonal, and decadal dynamics of VWC reflect variations in different processes relating the tree response to water stress. VWC can also be used for inferring belowground conditions—which are difficult to impossible to observe directly. Lastly, we discuss how a dedicated geostationary spaceborne observational system for VWC, when combined with existing datasets, can capture diel and seasonal water dynamics to advance the science and applications of global forest vulnerability to future droughts. [ABSTRACT FROM AUTHOR]

Summary: Tree water use is central to plant function and ecosystem fluxes. However, it is still unknown how organ‐level water‐relations traits are coordinated to determine whole‐tree water‐use strategies in response to drought, and whether this coordination depends on climate.Here we used a global sap flow database (SAPFLUXNET) to study the response of water use, in terms of whole‐tree canopy conductance (G), to vapour pressure deficit (VPD) and to soil water content (SWC) for 142 tree species. We investigated the individual and coordinated effect of six water‐relations traits (vulnerability to embolism, Huber value, hydraulic conductivity, turgor‐loss point, rooting depth and leaf size) on water‐use parameters, also accounting for the effect of tree height and climate (mean annual precipitation, MAP).Reference G and its sensitivity to VPD were tightly coordinated with water‐relations traits rather than with MAP. Species with efficient xylem transport had higher canopy conductance but also higher sensitivity to VPD. Moreover, we found that angiosperms had higher reference G and higher sensitivity to VPD than did gymnosperms.Our results highlight the need to consider trait integration and reveal the complications and challenges of defining a single, whole‐plant resource use spectrum ranging from 'acquisitive' to 'conservative'. [ABSTRACT FROM AUTHOR]

Summary: Trees partition biomass in response to resource limitation and physiological activity. It is presumed that these strategies evolved to optimize some measure of fitness. If the optimization criterion can be specified, then allometry can be modeled from first principles without prescribed parameterization.We present the Tree Hydraulics and Optimal Resource Partitioning (THORP) model, which optimizes allometry by estimating allocation fractions to organs as proportional to their ratio of marginal gain to marginal cost, where gain is net canopy photosynthesis rate, and costs are senescence rates. Root total biomass and profile shape are predicted simultaneously by a unified optimization. Optimal partitioning is solved by a numerically efficient analytical solution.THORP's predictions agree with reported tree biomass partitioning in response to size, water limitations, elevated CO2 and pruning. Roots were sensitive to soil moisture profiles and grew down to the groundwater table when present. Groundwater buffered against water stress regardless of meteorology, stabilizing allometry and root profiles as deep as c. 30 m.Much of plant allometry can be explained by hydraulic considerations. However, nutrient limitations cannot be fully ignored. Rooting mass and profiles were synchronized with hydrological conditions and groundwater even at considerable depths, illustrating that the below ground shapes whole‐tree allometry. [ABSTRACT FROM AUTHOR]

Summary: Global warming is expected to exacerbate the duration and intensity of droughts in the western United States, which may lead to increased tree mortality. A prevailing proximal mechanism of drought‐induced tree mortality is hydraulic damage, but predicting tree mortality from hydraulic theory and climate data still remains a major scientific challenge.We used forest inventory data and a plant hydraulic model (HM) to address three questions: can we capture regional patterns of drought‐induced tree mortality with HM‐predicted damage thresholds; do HM metrics improve predictions of mortality across broad spatial areas; and what are the dominant controls of forest mortality when considering stand characteristics, climate metrics, and simulated hydraulic stress?We found that the amount of variance explained by models predicting mortality was limited (R2 median = 0.10, R2 range: 0.00–0.52). HM outputs, including hydraulic damage and carbon assimilation diagnostics, moderately improve mortality prediction across the western US compared with models using stand and climate predictors alone.Among factors considered, metrics of stand density and tree size tended to be some of the most critical factors explaining mortality, probably highlighting the important roles of structural overshoot, stand development, and biotic agent host selection and outbreaks in mortality patterns. See also the Commentary on this article by Rowland et al., 230: 1685–1687. [ABSTRACT FROM AUTHOR]

Summary: Nocturnal transpiration is widely observed across species and biomes, and may significantly impact global water, carbon, and energy budgets. However, it remains elusive why plants lose water at night and how to model it at large scales.We hypothesized that plants optimize nighttime leaf diffusive conductance (gwn) to balance potential daytime photosynthetic benefits and nocturnal transpiration benefits. We quantified nighttime benefits from respiratory reductions due to evaporative leaf cooling. We described nighttime costs in terms of a reduced carbon gain during the day because of water use at night. We measured nighttime stomatal responses and tested our model with water birch (Betula occidentalis) saplings grown in a glasshouse.The gwn of water birch decreased with drier soil, higher atmospheric CO2, wetter air, lower leaf temperature, and lower leaf respiration rate. Our model predicted all these responses correctly, except for the response of gwn to air humidity. Our results also suggested that the slow decrease in gwn after sunset could be associated with decreasing leaf respiration.The optimality‐based nocturnal transpiration model smoothly integrates with daytime stomatal optimization approaches, and thus has the potential to quantitatively predict nocturnal transpiration across space and time. [ABSTRACT FROM AUTHOR]

For migratory species, seasonal movements complicate local climate adaptation, as it is unclear whether individuals track climate niches across the annual cycle. In the migratory songbird yellow warbler (Setophaga petechia), we find a correlation between individual‐level wintering and breeding precipitation, but not temperature. Birds wintering in the driest regions of the Neotropics breed in the driest regions of North America. Individuals from drier regions also possess distinct morphologies and population responses to varying rainfall. We find a positive association between bill size and breeding season precipitation which, given documented climate‐associated genomic variation, might reflect adaptation to local precipitation regimes. Relative abundance in the breeding range is linked to interannual fluctuations in precipitation, but the directionality of this response varies across geography. Together, our results suggest that variation in climate optima may exist across the breeding range of yellow warblers and provide a mechanism for selection across the annual cycle. [ABSTRACT FROM AUTHOR]

Carbon fluxes in terrestrial ecosystems and their response to environmental change are a major source of uncertainty in the modern carbon cycle. The National Ecological Observatory Network (NEON) presents the opportunity to merge eddy covariance (EC)‐derived fluxes with CO2 isotope ratio measurements to gain insights into carbon cycle processes. Collected continuously and consistently across >40 sites, NEON EC and isotope data facilitate novel integrative analyses. However, currently provisioned atmospheric isotope data are uncalibrated, greatly limiting ability to perform cross‐site analyses. Here, we present two approaches to calibrating NEON CO2 isotope ratios, along with an R package to calibrate NEON data. We find that calibrating CO2 isotopologues independently yields a lower δ13C bias (<0.05‰) and higher precision (<0.40‰) than directly correcting δ13C with linear regression (bias: <0.11‰, precision: 0.42‰), but with slightly higher error and lower precision in calibrated CO2 mole fraction. The magnitude of the corrections to δ13C and CO2 mole fractions vary substantially by site, underscoring the need for users to apply a consistent calibration framework to data in the NEON archive. Post‐calibration data sets show that site mean annual δ13C correlates negatively with precipitation, temperature, and aridity, but positively with elevation. Forested and agricultural ecosystems exhibit larger gradients in CO2 and δ13C than other sites, particularly during the summer and at night. The overview and analysis tools developed here will facilitate cross‐site analysis using NEON data, provide a model for other continental‐scale observational networks, and enable new advances leveraging the isotope ratios of specific carbon fluxes. Plain Language Summary: The magnitude and variability in the exchange of carbon between ecosystems and the atmosphere is a major source of uncertainty in carbon cycle models. These uncertainties have motivated substantial effort to measure these fluxes. Most observational methods provide an estimate of net carbon exchange, and efforts to break the net flux into its constituents requires a model, which without further constraints risks compounding observational uncertainties. Variations in the ratio of heavy‐to‐light carbon isotopes, 13C/12C, arise from photosynthetic responses to environmental conditions, and create a natural tracer to evaluate ecosystem fluxes. The NEON collects measurements of carbon isotope ratios of atmospheric carbon dioxide and plants and represents the largest network of consistently collected carbon isotope data associated with eddy covariance measurements to date. However, the accuracy of NEON's atmospheric carbon dioxide isotope ratios could be improved by using reference gas measurements to reduce bias and drift in these measurements. We develop and evaluate two correction methods and analyze continental‐scale patterns in the corrected atmospheric carbon dioxide isotope data. Further, we make these tools available as an R package to enable further cross‐site analyses using NEON's carbon isotope data. Key Points: Carbon isotope ratios record ecosystem assimilation and respiration processesWe present methods and tools for calibrating and drift‐correcting National Ecological Observatory Network CO2 isotope measurements, which enables cross‐site analysesCO2 isotope ratios vary with site precipitation, temperature, aridity, elevation, and ecosystem structure and composition [ABSTRACT FROM AUTHOR]

The Craig–Gordon type (C–G) leaf water isotope enrichment models assume a homogeneous distribution of enriched water across the leaf surface, despite observations that Δ18O can become increasingly enriched from leaf base to tip. Datasets of this 'progressive isotope enrichment' are limited, precluding a comprehensive understanding of (a) the magnitude and variability of progressive isotope enrichment, and (b) how progressive enrichment impacts the accuracy of C–G leaf water model predictions. Here, we present observations of progressive enrichment in two conifer species that capture seasonal and diurnal variability in environmental conditions. We further examine which leaf water isotope models best capture the influence of progressive enrichment on bulk needle water Δ18O. Observed progressive enrichment was large and equal in magnitude across both species. The magnitude of this effect fluctuated seasonally in concert with vapour pressure deficit, but was static in the face of diurnal cycles in meteorological conditions. Despite large progressive enrichment, three variants of the C–G model reasonably successfully predicted bulk needle Δ18O. Our results thus suggest that the presence of progressive enrichment does not impact the predictive success of C–G models, and instead yields new insight regarding the physiological and anatomical mechanisms that cause progressive isotope enrichment. Δ18O of needle water in two conifer species was drastically enriched down the length of the needle, the magnitude of which was driven by seasonal increases in VPD. Despite this progressive enrichment, traditional Craig‐Gordon model variants successfully predicted bulk needle water Δ18O. [ABSTRACT FROM AUTHOR]

Prediction of ecosystem responses to a changing climate is challenging at the landscape to regional scale, in part because topography creates various habitats and influences ecosystem productivity in complex ways. However, the effects of topography on ecosystem function remain poorly characterized and quantified. To address this knowledge gap, we developed a framework to systematically quantify and evaluate the effects of topographic convergence, elevation, aspect, and forest type on the long‐term (1986–2011) average and interannual variability of remotely sensed ecosystem productivity. In a forested watershed in the Rocky Mountains, spanning elevations from 1,800 to 4,000 m, we found a prevalent and positive influence of topographic convergence on long‐term productivity. Interannual growing season productivity was positively related to precipitation, with higher sensitivity in low elevation and highly productive areas and lower sensitivity in convergent areas. Our findings highlight the influence of topographic complexity on both long‐term and interannual variations of ecosystem productivity and have implications for understanding and prediction of ecosystem dynamics at hillslope to regional scales. Key Points: Remote sensing was used to quantify the effects of topographic complexity on ecosystem productivity and its temporal variationsHigher elevation, northeast aspects, and convergent valleys had higher long‐term ecosystem productivityEcosystem sensitivity to precipitation was higher in low elevation and highly productive areas and lower in convergent areas [ABSTRACT FROM AUTHOR]

Summary: Optimal stomatal control models have shown great potential in predicting stomatal behavior and improving carbon cycle modeling. Basic stomatal optimality theory posits that stomatal regulation maximizes the carbon gain relative to a penalty of stomatal opening. All models take a similar approach to calculate instantaneous carbon gain from stomatal opening (the gain function). Where the models diverge is in how they calculate the corresponding penalty (the penalty function). In this review, we compare and evaluate 10 different optimization models in how they quantify the penalty and how well they predict stomatal responses to the environment. We evaluate models in two ways. First, we compare their penalty functions against seven criteria that ensure a unique and qualitatively realistic solution. Second, we quantitatively test model against multiple leaf gas‐exchange datasets. The optimization models with better predictive skills have penalty functions that meet our seven criteria and use fitting parameters that are both few in number and physiology based. The most skilled models are those with a penalty function based on stress‐induced hydraulic failure. We conclude by proposing a new model that has a hydraulics‐based penalty function that meets all seven criteria and demonstrates a highly predictive skill against our test datasets. [ABSTRACT FROM AUTHOR]

This article is a Commentary on Eller et al., 226: 1622–1637, and Sabot et al., 226: 1638–1655. [ABSTRACT FROM AUTHOR]

Multi‐year lags in tree drought recovery, termed 'drought legacy effects', are important for understanding the impacts of drought on forest ecosystems, including carbon (C) cycle feedbacks to climate change. Despite the ubiquity of lags in drought recovery, large uncertainties remain regarding the mechanistic basis of legacy effects and their importance for the C cycle. In this review, we identify the approaches used to study legacy effects, from tree rings to whole forests. We then discuss key knowledge gaps pertaining to the causes of legacy effects, and how the various mechanisms that may contribute these lags in drought recovery could have contrasting implications for the C cycle. Furthermore, we conduct a novel data synthesis and find that legacy effects differ drastically in both size and length across the US depending on if they are identified in tree rings versus gross primary productivity. Finally, we highlight promising approaches for future research to improve our capacity to model legacy effects and predict their impact on forest health. We emphasise that a holistic view of legacy effects – from tissues to whole forests – will advance our understanding of legacy effects and stimulate efforts to investigate drought recovery via experimental, observational and modelling approaches. [ABSTRACT FROM AUTHOR]

Plant functional traits provide a link in process‐based vegetation models between plant‐level physiology and ecosystem‐level responses. Recent advances in physiological understanding and computational efficiency have allowed for the incorporation of plant hydraulic processes in large‐scale vegetation models. However, a more mechanistic representation of water limitation that determines ecosystem responses to plant water stress necessitates a re‐evaluation of trait‐based constraints for plant carbon allocation, particularly allocation to leaf area. In this review, we examine model representations of plant allocation to leaves, which is often empirically set by plant functional type‐specific allometric relationships. We analyze the evolution of the representation of leaf allocation in models of different scales and complexities. We show the impacts of leaf allocation strategy on plant carbon uptake in the context of recent advancements in modeling hydraulic processes. Finally, we posit that deriving allometry from first principles using mechanistic hydraulic processes is possible and should become standard practice, rather than using prescribed allometries. The representation of allocation as an emergent property of scarce resource constraints is likely to be critical to representing how global change processes impact future ecosystem dynamics and carbon fluxes and may reduce the number of poorly constrained parameters in vegetation models. [ABSTRACT FROM AUTHOR]

Drought‐induced tree mortality is projected to increase due to climate change, which will have manifold ecological and societal impacts including the potential to weaken or reverse the terrestrial carbon sink. Predictions of tree mortality remain limited, in large part because within‐species variations in ecophysiology due to plasticity or adaptation and ecosystem adjustments could buffer mortality in dry locations. Here, we conduct a meta‐analysis of 50 studies spanning >100 woody plant species globally to quantify how populations within species vary in vulnerability to drought mortality and whether functional traits or climate mediate mortality patterns. We find that mortality predominantly occurs in drier populations and this pattern is more pronounced in species with xylem that can tolerate highly negative water potentials, typically considered to be an adaptive trait for dry regions, and species that experience higher variability in water stress. Our results indicate that climate stress has exceeded physiological and ecosystem‐level tolerance or compensating mechanisms by triggering extensive mortality at dry range edges and provides a foundation for future mortality projections in empirical distribution and mechanistic vegetation models. [ABSTRACT FROM AUTHOR]

Forest leaf area has enormous leverage on the carbon cycle because it mediates both forest productivity and resilience to climate extremes. Despite widespread evidence that trees are capable of adjusting to changes in environment across both space and time through modifying carbon allocation to leaves, many vegetation models use fixed carbon allocation schemes independent of environment, which introduces large uncertainties into predictions of future forest responses to atmospheric CO2 fertilization and anthropogenic climate change. Here, we develop an optimization‐based model, whereby tree carbon allocation to leaves is an emergent property of environment and plant hydraulic traits. Using a combination of meta‐analysis, observational datasets, and model predictions, we find strong evidence that optimal hydraulic–carbon coupling explains observed patterns in leaf allocation across large environmental and CO2 concentration gradients. Furthermore, testing the sensitivity of leaf allocation strategy to a diversity in hydraulic and economic spectrum physiological traits, we show that plant hydraulic traits in particular have an enormous impact on the global change response of forest leaf area. Our results provide a rigorous theoretical underpinning for improving carbon cycle predictions through advancing model predictions of leaf area, and underscore that tree‐level carbon allocation to leaves should be derived from first principles using mechanistic plant hydraulic processes in the next generation of vegetation models. [ABSTRACT FROM AUTHOR]

Summary: Determining physiological mechanisms and thresholds for climate‐driven tree die‐off could help improve global predictions of future terrestrial carbon sinks. We directly tested for the lethal threshold in hydraulic failure – an inability to move water due to drought‐induced xylem embolism – in a pine sapling experiment.In a glasshouse experiment, we exposed loblolly pine (Pinus taeda) saplings (n = 83) to drought‐induced water stress ranging from mild to lethal. Before rewatering to relieve drought stress, we measured native hydraulic conductivity and foliar color change. We monitored all measured individuals for survival or mortality.We found a lethal threshold at 80% loss of hydraulic conductivity – a point of hydraulic failure beyond which it is more likely trees will die, than survive, and describe mortality risk across all levels of water stress. Foliar color changes lagged behind hydraulic failure – best predicting when trees had been dead for some time, rather than when they were dying.Our direct measurement of native conductivity, while monitoring the same individuals for survival or mortality, quantifies a continuous probability of mortality risk from hydraulic failure. Predicting tree die‐off events and understanding the mechanism involved requires knowledge not only of when trees are dead, but when they begin dying – having passed the point of no return. [ABSTRACT FROM AUTHOR]

Severe drought can cause lagged effects on tree physiology that negatively impact forest functioning for years. These "drought legacy effects" have been widely documented in tree‐ring records and could have important implications for our understanding of broader scale forest carbon cycling. However, legacy effects in tree‐ring increments may be decoupled from ecosystem fluxes due to (a) postdrought alterations in carbon allocation patterns; (b) temporal asynchrony between radial growth and carbon uptake; and (c) dendrochronological sampling biases. In order to link legacy effects from tree rings to whole forests, we leveraged a rich dataset from a Midwestern US forest that was severely impacted by a drought in 2012. At this site, we compiled tree‐ring records, leaf‐level gas exchange, eddy flux measurements, dendrometer band data, and satellite remote sensing estimates of greenness and leaf area before, during, and after the 2012 drought. After accounting for the relative abundance of tree species in the stand, we estimate that legacy effects led to ~10% reductions in tree‐ring width increments in the year following the severe drought. Despite this stand‐scale reduction in radial growth, we found that leaf‐level photosynthesis, gross primary productivity (GPP), and vegetation greenness were not suppressed in the year following the 2012 drought. Neither temporal asynchrony between radial growth and carbon uptake nor sampling biases could explain our observations of legacy effects in tree rings but not in GPP. Instead, elevated leaf‐level photosynthesis co‐occurred with reduced leaf area in early 2013, indicating that resources may have been allocated away from radial growth in conjunction with postdrought upregulation of photosynthesis and repair of canopy damage. Collectively, our results indicate that tree‐ring legacy effects were not observed in other canopy processes, and that postdrought canopy allocation could be an important mechanism that decouples tree‐ring signals from GPP. [ABSTRACT FROM AUTHOR]

Climate change‐driven drought stress has triggered numerous large‐scale tree mortality events in recent decades. Advances in mechanistic understanding and prediction are greatly limited by an inability to detect in situ where trees are likely to die in order to take timely measurements and actions. Thus, algorithms of early warning and detection of drought‐induced tree stress and mortality could have major scientific and societal benefits. Here, we leverage two consecutive droughts in the southwestern United States to develop and test a set of early warning metrics. Using Landsat satellite data, we constructed early warning metrics from the first drought event. We then tested these metrics' ability to predict spatial patterns in tree physiological stress and mortality from the second drought. To test the broader applicability of these metrics, we also examined a separate drought in the Amazon rainforest. The early warning metrics successfully explained subsequent tree mortality in the second drought in the southwestern US, as well as mortality in the independent drought in tropical forests. The metrics also strongly correlated with spatial patterns in tree hydraulic stress underlying mortality, which provides a strong link between tree physiological stress and remote sensing during the severe drought and indicates that the loss of hydraulic function during drought likely mediated subsequent mortality. Thus, early warning metrics provide a critical foundation for elucidating the physiological mechanisms underpinning tree mortality in mature forests and guiding management responses to these climate‐induced disturbances. [ABSTRACT FROM AUTHOR]

Summary: Drought‐induced tree mortality has major impacts on ecosystem carbon and water cycles, and is expected to increase in forests across the globe with climate change. A large body of research in the past decade has advanced our understanding of plant water and carbon relations under drought. However, despite intense research, we still lack generalizable, cross‐scale indicators of mortality risk. In this Viewpoint, we propose that a more explicit consideration of water pools could improve our ability to monitor and anticipate mortality risk. Specifically, we focus on the relative water content (RWC), a classic metric in plant water relations, as a potential indicator of mortality risk that is physiologically relevant and integrates different aspects related to hydraulics, stomatal responses and carbon economy under drought. Measures of plant water content are likely to have a strong mechanistic link with mortality and to be integrative, threshold‐prone and relatively easy to measure and monitor at large spatial scales, and may complement current mortality metrics based on water potential, loss of hydraulic conductivity and nonstructural carbohydrates. We discuss some of the potential advantages and limitations of these metrics to improve our capacity to monitor and predict drought‐induced tree mortality. [ABSTRACT FROM AUTHOR]

Summary: The widely documented phenomenon of nighttime stomatal conductance gsn could lead to substantial water loss with no carbon gain, and thus it remains unclear whether nighttime stomatal conductance confers a functional advantage. Given that studies of gsn have focused on controlled environments or small numbers of species in natural environments, a broad phylogenetic and biogeographic context could provide insights into potential adaptive benefits of gsn.We measured gsn on a diverse suite of species (n = 73) across various functional groups and climates‐of‐origin in a common garden to study the phylogenetic and biogeographic/climatic controls on gsn and further assessed the degree to which gsn co‐varied with leaf functional traits and daytime gas‐exchange rates.Closely related species were more similar in gsn than expected by chance. Herbaceous species had higher gsn than woody species. Species that typically grow in climates with lower mean annual precipitation – where the fitness cost of water loss should be the highest – generally had higher gsn.Our results reveal the highest gsn rates in species from environments where neighboring plants compete most strongly for water, suggesting a possible role for the competitive advantage of gsn. [ABSTRACT FROM AUTHOR]

Drought‐induced tree mortality is expected to increase in future climates with the potential for significant consequences to global carbon, water, and energy cycles. Xylem embolism can accumulate to lethal levels during drought, but species that can refill embolized xylem and recover hydraulic function may be able to avoid mortality. Yet the potential controls of embolism recovery, including cross‐biome patterns and plant traits such as nonstructural carbohydrates (NSCs), hydraulic traits, and nocturnal stomatal conductance, are unknown. We exposed eight plant species, originating from mesic (tropical and temperate) and semi‐arid environments, to drought under ambient and elevated CO2 levels, and assessed recovery from embolism following rewatering. We found a positive association between xylem recovery and NSCs, and, surprisingly, a positive relationship between xylem recovery and nocturnal stomatal conductance. Arid‐zone species exhibited greater embolism recovery than mesic zone species. Our results indicate that nighttime stomatal conductance often assumed to be a wasteful use of water, may in fact be a key part of plant drought responses, and contribute to drought survival. Findings suggested distinct biome‐specific responses that partially depended on species climate‐of‐origin precipitation or aridity index, which allowed some species to recover from xylem embolism. These findings provide improved understanding required to predict the response of diverse plant communities to drought. Our results provide a framework for predicting future vegetation shifts in response to climate change. [ABSTRACT FROM AUTHOR]

Summary: Empirical models of plant drought responses rely on parameters that are difficult to specify a priori. We test a trait‐ and process‐based model to predict environmental responses from an optimization of carbon gain vs hydraulic risk.We applied four drought treatments to aspen (Populus tremuloides) saplings in a research garden. First we tested the optimization algorithm by using predawn xylem pressure as an input. We then tested the full model which calculates root‐zone water budget and xylem pressure hourly throughout the growing season.The optimization algorithm performed well when run from measured predawn pressures. The per cent mean absolute error (MAE) averaged 27.7% for midday xylem pressure, transpiration, net assimilation, leaf temperature, sapflow, diffusive conductance and soil‐canopy hydraulic conductance. Average MAE was 31.2% for the same observations when the full model was run from irrigation and rain data. Saplings that died were projected to exceed 85% loss in soil‐canopy hydraulic conductance, whereas surviving plants never reached this threshold.The model fit was equivalent to that of an empirical model, but with the advantage that all inputs are specific traits. Prediction is empowered because knowing these traits allows knowing the response to climatic stress. [ABSTRACT FROM AUTHOR]

Understanding which species are able to recover from drought, under what conditions, and the mechanistic processes involved, will facilitate predictions of plant mortality in response to global change. In response to drought, some species die because of embolism-induced hydraulic failure, whilst others are able to avoid mortality and recover, following rehydration. Several tree species have evolved strategies to avoid embolism, whereas others tolerate high embolism rates but can recover their hydraulic functioning upon drought relief. Here, we focus on structures and processes that might allow some plants to recover from drought stress via embolism reversal. We provide insights into how embolism repair may have evolved, anatomical and physiological features that facilitate this process, and describe possible trade-offs and related costs. Recent controversies on methods used for estimating embolism formation/repair are also discussed, providing some methodological suggestions. Although controversial, embolism repair processes are apparently based on the activity of phloem and ray/axial parenchyma. The mechanism is energetically demanding, and the costs to plants include metabolism and transport of soluble sugars, water and inorganic ions. We propose that embolism repair should be considered as a possible component of a ‘hydraulic efficiency-safety’ spectrum. We also advance a framework for vegetation models, describing how vulnerability curves may change in hydrodynamic model formulations for plants that recover from embolism. [ABSTRACT FROM AUTHOR]

Abstract: Stomatal response to environmental conditions forms the backbone of all ecosystem and carbon cycle models, but is largely based on empirical relationships. Evolutionary theories of stomatal behaviour are critical for guarding against prediction errors of empirical models under future climates. Longstanding theory holds that stomata maximise fitness by acting to maintain constant marginal water use efficiency over a given time horizon, but a recent evolutionary theory proposes that stomata instead maximise carbon gain minus carbon costs/risk of hydraulic damage. Using data from 34 species that span global forest biomes, we find that the recent carbon‐maximisation optimisation theory is widely supported, revealing that the evolution of stomatal regulation has not been primarily driven by attainment of constant marginal water use efficiency. Optimal control of stomata to manage hydraulic risk is likely to have significant consequences for ecosystem fluxes during drought, which is critical given projected intensification of the global hydrological cycle. [ABSTRACT FROM AUTHOR]

Abstract: The mechanistic understanding of drought‐induced forest mortality hinges on improved models that incorporate the interactions between plant physiological responses and the spatiotemporal dynamics of water availability. We present a new framework integrating a three‐dimensional groundwater model, Parallel Flow, with a physiologically sophisticated plant model, Terrestrial Regional Ecosystem Exchange Simulator. The integrated model, Parallel Flow‐Terrestrial Regional Ecosystem Exchange Simulator, was demonstrated to quantify the susceptibility of riparian cottonwoods (Populus angustifolia, Populus deltoides, and native hybrids) in southwestern Canada to sustained atmospheric drought and variability in stream flow. The model reasonably captured the dynamics of soil moisture and evapotranspiration in both wet and dry years, including the resilience of cottonwoods despite their high vulnerability to xylem cavitation. Unrealistic predictions of mortality could be generated when ignoring lateral groundwater flow. Our results also illustrated a mechanistic linkage between streamflow and cottonwood health. In the absence of precipitation, normal streamflow could sustain 94% of cottonwoods, and higher streamflows would be required to sustain all of the floodplain cottonwoods. Further, the risk of mortality was mediated by plant hydraulic properties. These results underpin the importance of integrating groundwater processes and plant hydraulics in order to analyze the forest response to sustained severe drought, which could increase in the future due to climate change combined with increasing river water withdrawals. [ABSTRACT FROM AUTHOR]

Summary: Accumulating evidence highlights increased mortality risks for trees during severe drought, particularly under warmer temperatures and increasing vapour pressure deficit (VPD). Resulting forest die‐off events have severe consequences for ecosystem services, biophysical and biogeochemical land–atmosphere processes. Despite advances in monitoring, modelling and experimental studies of the causes and consequences of tree death from individual tree to ecosystem and global scale, a general mechanistic understanding and realistic predictions of drought mortality under future climate conditions are still lacking. We update a global tree mortality map and present a roadmap to a more holistic understanding of forest mortality across scales. We highlight priority research frontiers that promote: (1) new avenues for research on key tree ecophysiological responses to drought; (2) scaling from the tree/plot level to the ecosystem and region; (3) improvements of mortality risk predictions based on both empirical and mechanistic insights; and (4) a global monitoring network of forest mortality. In light of recent and anticipated large forest die‐off events such a research agenda is timely and needed to achieve scientific understanding for realistic predictions of drought‐induced tree mortality. The implementation of a sustainable network will require support by stakeholders and political authorities at the international level. [ABSTRACT FROM AUTHOR]

Abstract: Rapid warming and changes in water availability at high latitudes alter resource abundance, tree competition, and disturbance regimes. While these changes are expected to disrupt the functioning of boreal forests, their ultimate implications for forest composition are uncertain. In particular, recent site‐level studies of the Alaskan boreal forest have reported both increases and decreases in productivity over the past few decades. Here, we test the idea that variations in Alaskan forest growth and mortality rates are contingent on species composition. Using forest inventory measurements and climate data from plots located throughout interior and south‐central Alaska, we show significant growth and mortality responses associated with competition, midsummer vapor pressure deficit, and increased growing season length. The governing climate and competition processes differed substantially across species. Surprisingly, the most dramatic climate response occurred in the drought tolerant angiosperm species, trembling aspen, and linked high midsummer vapor pressure deficits to decreased growth and increased insect‐related mortality. Given that species composition in the Alaskan and western Canadian boreal forests is projected to shift toward early‐successional angiosperm species due to fire regime, these results underscore the potential for a reduction in boreal productivity stemming from increases in midsummer evaporative demand. [ABSTRACT FROM AUTHOR]

The article reports on the international and interdisciplinary meeting on tree mortality held in Hanover, Germany in 2018. Topics discussed in the meeting include global initiative for global forest monitoring, forest growth assessments, and models of global tree mortality. Also being discussed are the increasing amount of data on forest ecosystems.

Numerous current efforts seek to improve the representation of ecosystem ecology and vegetation demographic processes within Earth System Models ( ESMs). These developments are widely viewed as an important step in developing greater realism in predictions of future ecosystem states and fluxes. Increased realism, however, leads to increased model complexity, with new features raising a suite of ecological questions that require empirical constraints. Here, we review the developments that permit the representation of plant demographics in ESMs, and identify issues raised by these developments that highlight important gaps in ecological understanding. These issues inevitably translate into uncertainty in model projections but also allow models to be applied to new processes and questions concerning the dynamics of real-world ecosystems. We argue that stronger and more innovative connections to data, across the range of scales considered, are required to address these gaps in understanding. The development of first-generation land surface models as a unifying framework for ecophysiological understanding stimulated much research into plant physiological traits and gas exchange. Constraining predictions at ecologically relevant spatial and temporal scales will require a similar investment of effort and intensified inter-disciplinary communication. [ABSTRACT FROM AUTHOR]

Abstract: The concept of isohydry/anisohydry describes the degree to which plants regulate their water status, operating from isohydric with strict regulation to anisohydric with less regulation. Though some species level measures of isohydry/anisohydry exist at a few locations, ecosystem‐scale information is still largely unavailable. In this study, we use diurnal observations from active (Ku‐Band backscatter from QuikSCAT) and passive (X‐band vegetation optical depth (VOD) from Advanced Microwave Scanning Radiometer on EOS Aqua) microwave satellite data to estimate global ecosystem isohydry/anisohydry. Here diurnal observations from both satellites approximate predawn and midday plant canopy water contents, which are used to estimate isohydry/anisohydry. The two independent estimates from radar backscatter and VOD show reasonable agreement at low and middle latitudes but diverge at high latitudes. Grasslands, croplands, wetlands, and open shrublands are more anisohydric, whereas evergreen broadleaf and deciduous broadleaf forests are more isohydric. The direct validation with upscaled in situ species isohydry/anisohydry estimates indicates that the VOD‐based estimates have much better agreement than the backscatter‐based estimates. The indirect validation with prior knowledge suggests that both estimates are generally consistent in that vegetation water status of anisohydric ecosystems more closely tracks environmental fluctuations of water availability and demand than their isohydric counterparts. However, uncertainties still exist in the isohydry/anisohydry estimate, primarily arising from the remote sensing data and, to a lesser extent, from the methodology. The comprehensive assessment in this study can help us better understand the robustness, limitation, and uncertainties of the satellite‐derived isohydry/anisohydry estimates. The ecosystem isohydry/anisohydry has the potential to reveal new insights into spatiotemporal ecosystem response to droughts. [ABSTRACT FROM AUTHOR]

Stomatal regulation presumably evolved to optimize CO2 for H2O exchange in response to changing conditions. If the optimization criterion can be readily measured or calculated, then stomatal responses can be efficiently modelled without recourse to empirical models or underlying mechanism. Previous efforts have been challenged by the lack of a transparent index for the cost of losing water. Yet it is accepted that stomata control water loss to avoid excessive loss of hydraulic conductance from cavitation and soil drying. Proximity to hydraulic failure and desiccation can represent the cost of water loss. If at any given instant, the stomatal aperture adjusts to maximize the instantaneous difference between photosynthetic gain and hydraulic cost, then a model can predict the trajectory of stomatal responses to changes in environment across time. Results of this optimization model are consistent with the widely used Ball-Berry-Leuning empirical model ( r2 > 0.99) across a wide range of vapour pressure deficits and ambient CO2 concentrations for wet soil. The advantage of the optimization approach is the absence of empirical coefficients, applicability to dry as well as wet soil and prediction of plant hydraulic status along with gas exchange. [ABSTRACT FROM AUTHOR]

Fire regimes in savannas and forests are changing over much of the world. Anticipating the impact of these changes requires understanding how plants are adapted to fire. In this study, we test whether fire imposes a broad selective force on a key fire-tolerance trait, bark thickness, across 572 tree species distributed worldwide. We show that investment in thick bark is a pervasive adaptation in frequently burned areas across savannas and forests in both temperate and tropical regions where surface fires occur. Geographic variability in bark thickness is largely explained by annual burned area and precipitation seasonality. Combining environmental and species distribution data allowed us to assess vulnerability to future climate and fire conditions: tropical rainforests are especially vulnerable, whereas seasonal forests and savannas are more robust. The strong link between fire and bark thickness provides an avenue for assessing the vulnerability of tree communities to fire and demands inclusion in global models. [ABSTRACT FROM AUTHOR]

Elevated forest mortality has been attributed to climate change-induced droughts, but prediction of spatial mortality patterns remains challenging. We evaluated whether introducing plant hydraulics and topographic convergence-induced soil moisture variation to land surface models ( LSM) can help explain spatial patterns of mortality., A scheme predicting plant hydraulic safety loss from soil moisture was developed using field measurements and a plant physiology-hydraulics model, TREES. The scheme was upscaled to Populus tremuloides forests across Colorado, USA, using LSM-modeled and topography-mediated soil moisture, respectively. The spatial patterns of hydraulic safety loss were compared against aerial surveyed mortality., Incorporating hydraulic safety loss raised the explanatory power of mortality by 40% compared to LSM-modeled soil moisture. Topographic convergence was mostly influential in suppressing mortality in low and concave areas, explaining an additional 10% of the variations in mortality for those regions., Plant hydraulics integrated water stress along the soil-plant continuum and was more closely tied to plant physiological response to drought. In addition to the well-recognized topo-climate influence due to elevation and aspect, we found evidence that topographic convergence mediates tree mortality in certain parts of the landscape that are low and convergent, likely through influences on plant-available water. [ABSTRACT FROM AUTHOR]

Ecosystem models have difficulty predicting plant drought responses, partially from uncertainty in the stomatal response to water deficits in soil and atmosphere. We evaluate a 'supply-demand' theory for water-limited stomatal behavior that avoids the typical scaffold of empirical response functions. The premise is that canopy water demand is regulated in proportion to threat to supply posed by xylem cavitation and soil drying., The theory was implemented in a trait-based soil-plant-atmosphere model. The model predicted canopy transpiration ( E), canopy diffusive conductance ( G), and canopy xylem pressure ( Pcanopy) from soil water potential ( Psoil) and vapor pressure deficit ( D)., Modeled responses to D and Psoil were consistent with empirical response functions, but controlling parameters were hydraulic traits rather than coefficients. Maximum hydraulic and diffusive conductances and vulnerability to loss in hydraulic conductance dictated stomatal sensitivity and hence the iso- to anisohydric spectrum of regulation. The model matched wide fluctuations in G and Pcanopy across nine data sets from seasonally dry tropical forest and piñon-juniper woodland with < 26% mean error., Promising initial performance suggests the theory could be useful in improving ecosystem models. Better understanding of the variation in hydraulic properties along the root-stem-leaf continuum will simplify parameterization. [ABSTRACT FROM AUTHOR]

Aim The differential responses of plant species to climate change are of great interest and grave concern for scientists and conservationists. One underexploited resource for better understanding these changes are the records held by herbaria. Using these records to assess the responses of different groups of species across the entire flora of California, we sought to quantify the magnitude of species elevational shifts, to measure differences in shifts among functional groups and between native and introduced species, and to evaluate whether these shifts were related to the conservation of thermal niches. Location California. Methods To characterize these shifts in California, we used 681,609 georeferenced herbarium records to estimate mean shifts in elevational and climatic space of 4426 plant taxa. We developed and employed a statistical method to robustly analyse the data represented in these records. Results We found that 15% of all taxa in California have ranges that have shifted upward over the past century. There are significant differences between range shifts of taxa with different naturalization statuses: 12% of endemic taxa show significant upward range shifts, while a greater proportion (27%) of introduced taxa have shifted upward. We found significant differences between the proportion of significant range shifts across taxa with different seed sizes, but did not find evidence for differences in shift based on life-form (annual versus perennial, herbaceous versus woody). Main conclusions Our analyses suggest that introduced species have disproportionately expanded their ranges upward in elevation over the past century when compared with native species.While these shifts in introduced species may not be exclusively driven by climate, they highlight the importance of considering the interacting factors of climate-driven range shifts and invasion to understand how floras are responding in the face of anthropogenic change. [ABSTRACT FROM AUTHOR]