Community assembly influences plant trait economic spectra and functional trade-offs at ecosystem scales.

Plant functional traits hold the potential to greatly improve the understanding and prediction of climate impacts on ecosystems and carbon cycle feedback to climate change. Traits are commonly used to place species along a global conservative-acquisitive trade-off, yet how and if functional traits and conservative-acquisitive trade-offs scale up to mediate community and ecosystem fluxes is largely unknown. Here, we combine functional trait datasets and multibiome datasets of forest water and carbon fluxes at the species, community, and ecosystem-levels to quantify the scaling of the tradeoff between maximum flux and sensitivity to vapor pressure deficit. We find a strong conservative-acquisitive trade-off at the species scale, which weakens modestly at the community scale and largely disappears at the ecosystem scale. Functional traits, particularly plant water transport (hydraulic) traits, are strongly associated with the key dimensions of the conservative-acquisitive trade-off at community and ecosystem scales, highlighting that trait composition appears to influence community and ecosystem flux dynamics. Our findings provide a foundation for improving carbon cycle models by revealing i) that plant hydraulic traits are most strongly associated with community- and ecosystem scale flux dynamics and ii) community assembly dynamics likely need to be considered explicitly, as they give rise to ecosystem-level flux dynamics that differ substantially from trade-offs identified at the species-level.

Open science priorities for rigorous nature-based climate solutions.

Nature-based climate solutions (NbCS) hold promise, but must be based on the best available science to be successful. We outline key ingredients of open data and science crucial for robust and scalable nature-based climate solutions efforts, as an urgent call to action for academic researchers, nongovernmental organizations, government agencies, and private companies.

Dynamic regulation of water potential in Juniperus osteosperma mediates ecosystem carbon fluxes.

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.

A new empirical framework to quantify the hydraulic effects of soil and atmospheric drivers on plant water status.

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.

Hydraulic diversity of forests regulates ecosystem resilience during drought.

Plants influence the atmosphere through fluxes of carbon, water and energy1, and can intensify drought through land-atmosphere feedback effects2-4. The diversity of plant functional traits in forests, especially physiological traits related to water (hydraulic) transport, may have a critical role in land-atmosphere feedback, particularly during drought. Here we combine 352 site-years of eddy covariance measurements from 40 forest sites, remote-sensing observations of plant water content and plant functional-trait data to test whether the diversity in plant traits affects the response of the ecosystem to drought. We find evidence that higher hydraulic diversity buffers variation in ecosystem flux during dry periods across temperate and boreal forests. Hydraulic traits were the predominant significant predictors of cross-site patterns in drought response. By contrast, standard leaf and wood traits, such as specific leaf area and wood density, had little explanatory power. Our results demonstrate that diversity in the hydraulic traits of trees mediates ecosystem resilience to drought and is likely to have an important role in future ecosystem-atmosphere feedback effects in a changing climate.

Fire frequency drives decadal changes in soil carbon and nitrogen and ecosystem productivity.

Fire frequency is changing globally and is projected to affect the global carbon cycle and climate. However, uncertainty about how ecosystems respond to decadal changes in fire frequency makes it difficult to predict the effects of altered fire regimes on the carbon cycle; for instance, we do not fully understand the long-term effects of fire on soil carbon and nutrient storage, or whether fire-driven nutrient losses limit plant productivity. Here we analyse data from 48 sites in savanna grasslands, broadleaf forests and needleleaf forests spanning up to 65 years, during which time the frequency of fires was altered at each site. We find that frequently burned plots experienced a decline in surface soil carbon and nitrogen that was non-saturating through time, having 36 per cent (±13 per cent) less carbon and 38 per cent (±16 per cent) less nitrogen after 64 years than plots that were protected from fire. Fire-driven carbon and nitrogen losses were substantial in savanna grasslands and broadleaf forests, but not in temperate and boreal needleleaf forests. We also observe comparable soil carbon and nitrogen losses in an independent field dataset and in dynamic model simulations of global vegetation. The model study predicts that the long-term losses of soil nitrogen that result from more frequent burning may in turn decrease the carbon that is sequestered by net primary productivity by about 20 per cent of the total carbon that is emitted from burning biomass over the same period. Furthermore, we estimate that the effects of changes in fire frequency on ecosystem carbon storage may be 30 per cent too low if they do not include multidecadal changes in soil carbon, especially in drier savanna grasslands. Future changes in fire frequency may shift ecosystem carbon storage by changing soil carbon pools and nitrogen limitations on plant growth, altering the carbon sink capacity of frequently burning savanna grasslands and broadleaf forests.

Rapid increases in shrubland and forest intrinsic water-use efficiency during an ongoing megadrought.

Globally, intrinsic water-use efficiency (iWUE) has risen dramatically over the past century in concert with increases in atmospheric CO2 concentration. This increase could be further accelerated by long-term drought events, such as the ongoing multidecadal "megadrought" in the American Southwest. However, direct measurements of iWUE in this region are rare and largely constrained to trees, which may bias estimates of iWUE trends toward more mesic, high elevation areas and neglect the responses of other key plant functional types such as shrubs that are dominant across much of the region. Here, we found evidence that iWUE is increasing in the Southwest at one of the fastest rates documented due to the recent drying trend. These increases were particularly large across three common shrub species, which had a greater iWUE sensitivity to aridity than Pinus ponderosa, a common tree species in the western United States. The sensitivity of both shrub and tree iWUE to variability in atmospheric aridity exceeded their sensitivity to increasing atmospheric [CO2]. The shift to more water-efficient vegetation would be, all else being equal, a net positive for plant health. However, ongoing trends toward lower plant density, diminished growth, and increasing vegetation mortality across the Southwest indicate that this increase in iWUE is unlikely to offset the negative impacts of aridification.

Anthropogenic climate change is worsening North American pollen seasons.

Airborne pollen has major respiratory health impacts and anthropogenic climate change may increase pollen concentrations and extend pollen seasons. While greenhouse and field studies indicate that pollen concentrations are correlated with temperature, a formal detection and attribution of the role of anthropogenic climate change in continental pollen seasons is urgently needed. Here, we use long-term pollen data from 60 North American stations from 1990 to 2018, spanning 821 site-years of data, and Earth system model simulations to quantify the role of human-caused climate change in continental patterns in pollen concentrations. We find widespread advances and lengthening of pollen seasons (+20 d) and increases in pollen concentrations (+21%) across North America, which are strongly coupled to observed warming. Human forcing of the climate system contributed ∼50% (interquartile range: 19-84%) of the trend in pollen seasons and ∼8% (4-14%) of the trend in pollen concentrations. Our results reveal that anthropogenic climate change has already exacerbated pollen seasons in the past three decades with attendant deleterious effects on respiratory health.

Large volcanic eruptions elucidate physiological controls of tree growth and photosynthesis.

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.

Decoupling of functional traits from intraspecific patterns of growth and drought stress resistance.

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.

Linking remotely sensed ecosystem resilience with forest mortality across the continental United States.

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.

Declining tree growth resilience mediates subsequent forest mortality in the US Mountain West.

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.

Global patterns of drought recovery.

Drought, a recurring phenomenon with major impacts on both human and natural systems, is the most widespread climatic extreme that negatively affects the land carbon sink. Although twentieth-century trends in drought regimes are ambiguous, across many regions more frequent and severe droughts are expected in the twenty-first century. Recovery time-how long an ecosystem requires to revert to its pre-drought functional state-is a critical metric of drought impact. Yet the factors influencing drought recovery and its spatiotemporal patterns at the global scale are largely unknown. Here we analyse three independent datasets of gross primary productivity and show that, across diverse ecosystems, drought recovery times are strongly associated with climate and carbon cycle dynamics, with biodiversity and CO2 fertilization as secondary factors. Our analysis also provides two key insights into the spatiotemporal patterns of drought recovery time: first, that recovery is longest in the tropics and high northern latitudes (both vulnerable areas of Earth's climate system) and second, that drought impacts (assessed using the area of ecosystems actively recovering and time to recovery) have increased over the twentieth century. If droughts become more frequent, as expected, the time between droughts may become shorter than drought recovery time, leading to permanently damaged ecosystems and widespread degradation of the land carbon sink.

Trait velocities reveal that mortality has driven widespread coordinated shifts in forest hydraulic trait composition.

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

Forest and woodland replacement patterns following drought-related mortality.

Forest vulnerability to drought is expected to increase under anthropogenic climate change, and drought-induced mortality and community dynamics following drought have major ecological and societal impacts. Here, we show that tree mortality concomitant with drought has led to short-term (mean 5 y, range 1 to 23 y after mortality) vegetation-type conversion in multiple biomes across the world (131 sites). Self-replacement of the dominant tree species was only prevalent in 21% of the examined cases and forests and woodlands shifted to nonwoody vegetation in 10% of them. The ultimate temporal persistence of such changes remains unknown but, given the key role of biological legacies in long-term ecological succession, this emerging picture of postdrought ecological trajectories highlights the potential for major ecosystem reorganization in the coming decades. Community changes were less pronounced under wetter postmortality conditions. Replacement was also influenced by management intensity, and postdrought shrub dominance was higher when pathogens acted as codrivers of tree mortality. Early change in community composition indicates that forests dominated by mesic species generally shifted toward more xeric communities, with replacing tree and shrub species exhibiting drier bioclimatic optima and distribution ranges. However, shifts toward more mesic communities also occurred and multiple pathways of forest replacement were observed for some species. Drought characteristics, species-specific environmental preferences, plant traits, and ecosystem legacies govern postdrought species turnover and subsequent ecological trajectories, with potential far-reaching implications for forest biodiversity and ecosystem services.

Contemporary tree growth shows altered climate memory.

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.

Future climate risks from stress, insects and fire across US forests.

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.

Wood density and hydraulic traits influence species' growth response to drought across biomes.

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.