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deadtrees.earth — An open-access and interactive database for centimeter-scale aerial imagery to uncover global tree mortality dynamics

Excessive tree mortality is a global concern and remains poorly understood as it is a complex phenomenon. We lack global and temporally continuous coverage on tree mortality data. Ground-based observations on tree mortality, e.g., derived from national inventories, are very sparse, and may not be standardized or spatially explicit. Earth observation data, combined with supervised machine learning, offer a promising approach to map overstory tree mortality in a consistent manner over space and time. However, global-scale machine learning requires broad training data covering a wide range of environmental settings and forest types. Low altitude observation platforms (e.g., drones or airplanes) provide a cost-effective source of training data by capturing high-resolution orthophotos of overstory tree mortality events at centimeter-scale resolution. Here, we introduce deadtrees.earth, an open-access platform hosting more than two thousand centimeter-resolution orthophotos, covering more than 1,000,000 ha, of which more than 58,000 ha are manually annotated with live/dead tree classifications. This community-sourced and rigorously curated dataset can serve as a comprehensive reference dataset to uncover tree mortality patterns from local to global scales using space-based Earth observation data and machine learning models. This will provide the basis to attribute tree mortality patterns to environmental changes or project tree mortality dynamics to the future. The open nature of deadtrees.earth, together with its curation of high-quality, spatially representative, and ecologically diverse data will continuously increase our capacity to uncover and understand tree mortality dynamics.

Citizen science↗

Sensitivity of Optical Satellites to Estimate Windthrow Tree-Mortality in a Central Amazon Forest

Windthrow (i.e., trees broken and uprooted by wind) is a major natural disturbance in Amazon forests. Images from medium-resolution optical satellites combined with extensive field data have allowed researchers to assess patterns of windthrow tree-mortality and to monitor forest recovery over decades of succession in different regions. Although satellites with high spatial-resolution have become available in the last decade, they have not yet been employed for the quantification of windthrow tree-mortality. Here, we address how increasing the spatial resolution of satellites affects plot-to-landscape estimates of windthrow tree-mortality. We combined forest inventory data with Landsat 8 (30 m pixel), Sentinel 2 (10 m), and WorldView 2 (2 m) imagery over an old-growth forest in the Central Amazon that was disturbed by a single windthrow event in November 2015. Remote sensing estimates of windthrow tree-mortality were produced from Spectral Mixture Analysis and evaluated with forest inventory data (i.e., ground true) by using Generalized Linear Models. Field measured windthrow tree-mortality (3 transects and 30 subplots) crossing the entire disturbance gradient was 26.9 ± 11.1% (mean ± 95% CI). Although the three satellites produced reliable and statistically similar estimates (from 26.5% to 30.3%, p < 0.001), Landsat 8 had the most accurate results and efficiently captured field-observed variations in windthrow tree-mortality across the entire gradient of disturbance (Sentinel 2 and WorldView 2 produced the second and third best results, respectively). As expected, mean-associated uncertainties decreased systematically with increasing spatial resolution (i.e., from Landsat 8 to Sentinel 2 and WorldView 2). However, the overall quality of model fits showed the opposite pattern. We suggest that this reflects the influence of a relatively minor disturbance, such as defoliation and crown damage, and the fast growth of natural regeneration, which were not measured in the field nor can be captured by coarser resolution imagery. Our results validate the reliability of Landsat imagery for assessing plot-to-landscape patterns of windthrow tree-mortality in dense and heterogeneous tropical forests. Satellites with high spatial resolution can improve estimates of windthrow severity by allowing the quantification of crown damage and mortality of lower canopy and understory trees. However, this requires the validation of remote sensing metrics using field data at compatible scales.

54 ENVIRONMENTAL SCIENCES↗

Climate Change Risks to Global Forest Health: Emergence of Unexpected Events of Elevated Tree Mortality Worldwide

Recent observations of elevated tree mortality following climate extremes, like heat and drought, raise concerns about climate change risks to global forest health. We currently lack both sufficient data and understanding to identify whether these observations represent a global trend toward increasing tree mortality. In this work, we document events of sudden and unexpected elevated tree mortality following heat and drought events in ecosystems that previously were considered tolerant or not at risk of exposure. These events underscore the fact that climate change may affect forests with unexpected force in the future. We use the events as examples to highlight current difficulties and challenges for realistically predicting such tree mortality events and the uncertainties about future forest condition. Advances in remote sensing technology and greater availably of high-resolution data, from both field assessments and satellites, are needed to improve both understanding and prediction of forest responses to future climate change.

59 BASIC BIOLOGICAL SCIENCES↗

Estimating Individual Tree Mortality in the Sierra Nevada Using Lidar and Multispectral Reflectance Data

Widespread tree mortality events occur during periods of severe drought in temperate conifer forests and are expected to become more frequent in many areas due to climate change. Improved mapping of individual tree mortality is needed to identify risk factors and design effective conservation strategies. In this study, we used National Ecological Observatory Network (NEON) lidar and multispectral reflectance airborne observations to map individual tree mortality over a 160 km 2 area during and after the 2012–2016 drought for two sites in California's Sierra National Forest. We used NEON lidar to derive tree locations and crown perimeters and multispectral data to map tree mortality for more than 1 million trees. We found that 25.4% of the trees in our study area died between 2013 and 2017, with considerably higher mortality at the lower-elevation Soaproot Saddle site. Between 2017 and 2019, an additional 2.0%–2.8% of the trees died each year. Two wildfires in 2020 and 2021 increased tree mortality within burned area perimeters by 49%–89% between 2019 and 2021. Consistent with previous work, we found that tree mortality risk increased as a function of tree height. Tree mortality was positively associated with distance from rivers, trees per hectare, and decreasing slope at the lower elevation site. In contrast, increasing slope was positively associated with tree mortality at the higher elevation site. Our approach and dataset provide a means to study the combined effects of drought and wildfire on tree mortality and may improve projections of forest resilience under a changing climate.

54 ENVIRONMENTAL SCIENCES↗

Tree mortality in a warming world: causes, patterns, and implications

This ERL focus collection has published 17 papers that have advanced our understanding of different dimensions of warming-induced tree mortality. Here we summarize these focus collection papers, organized by four topics related to tree mortality: pathogens, droughts/heat waves, fire/bark beetles, and teleconnections/air pollution. This focus collection illustrates a variety of methods in measuring and modeling tree-mortality, and adds significant new research findings into the scientific literature on tree mortality from hotter droughts. Some of these results also are useful for policymakers and forest managers in addressing amplified forest stress and tree mortality as a result of increasingly severe warming-induced climate and weather extremes.

54 ENVIRONMENTAL SCIENCES↗

Tree mortality during long-term droughts is lower in structurally complex forest stands

Increasing drought frequency and severity in a warming climate threaten forest ecosystems with widespread tree deaths. Canopy structure is important in regulating tree mortality during drought, but how it functions remains controversial. Here, we show that the interplay between tree size and forest structure explains drought-induced tree mortality during the 2012-2016 California drought. Through an analysis of over one million trees, we find that tree mortality rate follows a “negative-positive-negative” piecewise relationship with tree height, and maintains a consistent negative relationship with neighborhood canopy structure (a measure of tree competition). Trees overshadowed by tall neighboring trees experienced lower mortality, likely due to reduced exposure to solar radiation load and lower water demand from evapotranspiration. Our findings demonstrate the significance of neighborhood canopy structure in influencing tree mortality and suggest that re-establishing heterogeneity in canopy structure could improve drought resiliency. Our study also indicates the potential of advances in remote-sensing technologies for silvicultural design, supporting the transition to multi-benefit forest management.

59 BASIC BIOLOGICAL SCIENCES↗

Drought–induced increase in tree mortality and corresponding decrease in the carbon sink capacity of Canada's boreal forests from 1970 to 2020

Canada's boreal forests, which occupy approximately 30% of boreal forests world wide, play an important role in the global carbon budget. However, there is lit tle quantitative information available regarding the spatiotemporal changes in the drought-induced tree mortality of Canada's boreal forests overall and their associ ated impacts on biomass carbon dynamics. Here, we develop spatiotemporally ex plicit estimates of drought-induced tree mortality and corresponding biomass carbon sink capacity changes in Canada's boreal forests from 1970 to 2020. We show that the average annual tree mortality rate is approximately 2.7%. Approximately 43% of Canada's boreal forests have experienced significantly increasing tree mortality trends (71% of which are located in the western region of the country), and these trends have accelerated since 2002. This increase in tree mortality has resulted in sig nificant biomass carbon losses at an approximate rate of 1.51±0.29 MgC ha -1 year -1 (95% confidence interval) with an approximate total loss of 0.46±0.09 PgC year -1 (95% confidence interval). Under the drought condition increases predicted for this century, the capacity of Canada's boreal forests to act as a carbon sink will be further reduced, potentially leading to a significant positive climate feedback effect

59 BASIC BIOLOGICAL SCIENCES↗

Understanding the impacts of elevated CO2 and vapor pressure deficit on tree mortality - CRADA 599 (Abstract)

Many studies have examined the effects of rising CO2 on gas exchange and growth of plants. However, concurrent with the rise in CO2 has been the rise in temperature and associated drying of the atmosphere, or vapor pressure deficit (VPD), which increases tree mortality. Little attention has been given to the net outcome of these antagonistic drivers of plant performance. The net balance of rising CO2 and VPD upon plant and ecosystem function is possibly the largest uncertainty in model projections of the terrestrial carbon and water cycles. The purpose of this CRADA is to improve our predictive understanding of tree mortality under our changing environment. We will collaborate with WSU to provide a mechanistic understanding of the relative impacts of CO2 and VPD on tree mortality that scales across the globe. Ultimately, we will understand the potential impacts and mechanisms underlying CO2 and VPD responses and will enable improved prediction of future tree mortality.

54 ENVIRONMENTAL SCIENCES↗

Fire, insect and disease‐caused tree mortalities increased in forests of greater structural diversity during drought

Abstract Structural diversity is an emerging dimension of biodiversity that accounts for size variations in organs among individuals in a community. Previous studies show significant effects of structural diversity on forest growth, but its effects on forest mortality are not known, particularly at a large scale. To address this knowledge gap, we quantified structural diversity using stem structural diversity (SSD) based on both tree diameter and height. We obtained U.S. Forest Service Forest Inventory and Analysis (FIA) data from over 2400 plots across southcentral U.S. forests that have suffered a recent drought. Using data from multiple sampling times, we calculated SSD and compared the relative importance of SSD, species diversity, functional diversity and other stand attributes in determining tree mortalities caused by fire, insects and diseases. We also used FIRETEC, a physics‐based fire model, to test the effect of SSD on canopy consumption by fire. Our results showed that (1) SSD was positively associated with tree mortalities caused by all three disturbances; (2) species richness was negatively associated with insect‐ and disease‐caused mortalities; (3) functional diversity was negatively associated with fire‐ and disease‐caused mortalities and (4) more phylogenetically related species had more similar mortality rates by insect and disease but not fire. Moreover, the FIRETEC model showed increasing canopy consumption by fire in stands with greater SSD. Together, the different tree mortalities during drought associated with SSD more consistently than the other biodiversity metrics were evaluated. Synthesis . Our results suggest that SSD could be considered in modelling forest dynamics and planning management to sustain forest health under disturbances.

54 ENVIRONMENTAL SCIENCES↗

Context matters: Natural tree mortality can lead to neighbor growth release or suppression

Where competition suppresses tree growth, mortality of adjacent trees can release the surviving individuals, leading to a growth increase. However, primary forests are complicated systems, where trees interact in both competitive and facilitative ways mediated by their size, species, and the broad ecological context in which they grow. Thus, the magnitude and even direction of growth responses to the mortality of nearby trees may vary, which has implications for our understanding of community- and ecosystem-level dynamics following mortality events. Unfortunately, although many studies focus on the impacts of light availability and general crowding on tree growth, we know relatively little about the effects of naturally occurring mortality events on the growth of neighboring trees. To address this issue, we used 40 years of data from 15 permanent forest-monitoring plots in Mt Rainier old-growth forests, comparing observed to expected radial growth of individual trees following the death of their nearest neighbor. Although we found evidence of a general growth-release response, this was not universal among all trees, with small trees in particular exhibiting growth suppression (rather than release) following neighboring tree mortality. In addition to small size, growth-suppression was more likely if the dead neighbor was the same species, consistent with facilitative effects as mediated through belowground networks. At the stand level, the average growth release after nearest neighbor mortality was greatest in low-density stands with large trees, with elevation and community composition also playing a role. Decades more monitoring could reveal how long growth release (or suppression) is sustained by individual trees following neighboring mortality events, as well as potential response lags and the role of species identity in determining whether interactions with neighbors are competitive or facilitative. Nonetheless, our results suggest that although mature trees have competitive effects on their larger neighbors, they also have an important role in supporting the ingrowth of small trees. More broadly, we demonstrate that the nature of interactions between individual neighboring trees is highly context dependent.

54 ENVIRONMENTAL SCIENCES↗

Recent California tree mortality portends future increase in drought-driven forest die-off

Vegetation tolerance to drought depends on an array of site-specific environmental and plant physiological factors. This tolerance is poorly understood for many forest types despite its importance for predicting and managing vegetation stress. We analyzed the relationships between precipitation variability and forest die-off in California's Sierra Nevada and introduce a new measure of drought tolerance that emphasizes plant access to subsurface moisture buffers. We applied this metric to California's severe 2012-2015 drought, and show that it predicted the patterns of tree mortality. We then examined future climate scenarios, and found that the probability of droughts that lead to widespread die-off increases threefold by the end of the 21st century. Our analysis shows that tree mortality in the Sierra Nevada will likely accelerate in the coming decades and that forests in the Central and Northern Sierra Nevada that largely escaped mortality in 2012-2015 are vulnerable to die-off.

54 ENVIRONMENTAL SCIENCES↗

Canopy tree mortality depends on the proportion of crown exposed to sunlight, but this effect varies with species' wood density

Abstract Understanding what drives changes in tree mortality as well as the covariates influencing trees' response is a research priority to predict forest responses to global change. Here, we combined drone photogrammetry and ground‐based data to assess the influence of crown exposure to light (relative to total crown area), growth deviations (relative to conspecifics), tree size, and species' wood density (as a surrogate for light‐demanding and shade‐tolerant life‐history strategies) on the mortality of 984 canopy trees in an Amazonterra firmeforest. Trees with lower wood density were less prone to die when their proportion of crown was more exposed to sunlight, but this relationship with relative crown exposure weakened and slightly reversed as wood density increased. Trees growing less than their species average had higher mortality, especially when the species' wood density decreased. The role of wood density in determining the survival of canopy trees under varying light conditions indicates differential responses of light‐demanding versus shade‐tolerant species. Our results highlight the importance of accounting for life‐history strategies, via plant functional types, in vegetation dynamic models aiming to predict forest demography under a rapidly changing climate. Abstract in Spanish is available with online material.

Environmental Sciences & Ecology↗

The influence of increasing atmospheric CO 2 , temperature, and vapor pressure deficit on seawater-induced tree mortality

We report increasing seawater exposure is killing coastal trees globally, with expectations of accelerating mortality with rising sea levels. However, the impact of concomitant changes in atmospheric CO 2 concentration, temperature, and vapor pressure deficit (VPD) on seawater-induced tree mortality is uncertain. We examined the mechanisms of seawater-induced mortality under varying climate scenarios using a photosynthetic gain and hydraulic cost optimization model validated against observations in a mature stand of Sitka-spruce (Picea sitchensis) trees in the Pacific Northwest, USA, that were dying from recent seawater exposure. The simulations matched well with observations of photosynthesis, transpiration, nonstructural carbohydrates concentrations, leaf water potential, the percentage loss of xylem conductivity, and stand-level mortality rates. The simulations suggest that seawater-induced mortality could decrease by ~16.7% with increasing atmospheric CO 2 levels due to reduced risk of carbon starvation. Conversely, rising VPD could increase mortality by ~5.6% because of increasing risk of hydraulic failure. Across all scenarios, seawater-induced mortality was driven by hydraulic failure in the first two years after seawater exposure began, with carbon starvation becoming more important in subsequent years. Changing CO 2 and climate appear unlikely to have a significant impact on coastal tree mortality under rising sea levels.

54 ENVIRONMENTAL SCIENCES↗

Turbulence regimes in the nocturnal roughness sublayer: Interaction with deep convection and tree mortality in the Amazon

We investigated the influence of seasonality and proximity to the forest canopy on nocturnal turbulence regimes in the roughness sublayer of a Central Amazon forest. Since convective systems of different scales are common in this region, we also analyzed the effect of extreme wind gusts (propagated from convective downdrafts) on the organization of the turbulence regimes, and their potential to cause the mortality of canopy trees. Our data include high-frequency winds, temperature and ozone concentration at different heights during the dry and wet seasons of 2014. In addition, we used critical wind-speed data derived from a tree-winching experiment and a modeling study conducted in the same study site. Two different turbulence regimes were identified at three heights above the canopy: a weakly stable (WS) and a very stable regime (VS). The threshold wind speeds that mark the transition between turbulence regimes were larger during the dry season and increased as a function of the height above the canopy. The turbulent fluxes of sensible heat and momentum during the WS accounted for 88% of the entire nighttime flux. Downdrafts occurred only in the WS and favored a fully coupled state of wind flow along the canopy profile. The destructive potential of winds was four times higher than on nights without downdrafts.

54 ENVIRONMENTAL SCIENCES↗

Sensitivity of Pyrocumulus Convection to Tree Mortality During the 2020 Creek Fire in California

Abstract This study investigated the sensitivity of pyrocumulonimbus (PyroCb) induced by the California Creek fire of 2020 to the amount and type of surface fuels, within the WRF‐SFIRE modeling system. Satellite data were used to derive fire arrival times to constrain fire progression, and to augment the fuel characterization with better estimates of combustible vegetation accounting for tree mortality. Machine learning was employed to classify standing dead vegetation from aerial imagery, which was then added as a custom fuel class along with the standard Anderson fuel categories. Simulations using this new fuel class produced a larger and more vigorous PyroCb than the control run, however, still under‐predicted the cloud top. Additional augmentation of fuel mass to represent the accumulation of dead vegetation on the forest floor further improved the simulations, demonstrating the efficacy of representing both dead standing and fallen vegetation to produce more realistic PyroCb and smoke simulations.

54 ENVIRONMENTAL SCIENCES↗

Canopy tree mortality and crown exposure data from the Amacayacu Forest Dynamics Plot, Northwestern Amazon

Data on the mortality of 984 canopy trees, their crown exposure to light (relative to total crown area), growth deviations (relative to conspecifics), tree size, and species’ wood density collected between 2013 and 2019 in 18 ha of the Amacayacu Forest Dynamics Plot, Northwestern Amazon. This dataset contains a single CSV data file. Variable definitions: 1. Species: [character] species identification 2. Family: [character] family of the species 3. tag: [character] unique consecutive for the tree 4. status: [character] status of the tree in the third census (ALIVE or DEAD) 5. wsg: [numeric]: species’ wood density (g cm-3) 6. growth_r1: [numeric]: annual growth rate of the tree between the first and second census (cm y-1) 7. growth_r2: [numeric]: annual growth rate of the tree between the second and third census (cm y-1) 8. gt1: [numeric] modulus transformed growth rate with a lambda of 0.4 for the annual growth rate of the tree between the first and second census (cm y-1) 9. gt2: [numeric]: modulus transformed growth rate with a lambda of 0.4 for the annual growth rate of the tree between the second and third census (cm y-1) 10. rgr1: [numeric] relative growth rate between the first and second census (cm y-1) 11. rgr2: [numeric] relative growth rate between the second and third census (cm y-1) 12. sa_gt: [numeric] species-adjusted modulus transformed growth rate 13. sa_rgr: [numeric] species-adjusted relative growth rate 14. gr_n: [numeric] number of individuals of the species used to calculate the mean and species modulus transformed growth rate 15. dbh_flight: [numeric] diameter at breast height (1.3 m) estimated at the time of the drone flight (cm) 16. eca: [numeric] exposed crown area calculated as the area of the crown polygon delineated in the orthomosaic (m2) 17. total_ca: [numeric] total crown area estimated from a crown area model (m2) 18. rcel: [numeric] relative crown exposure to light (m2) 19. time_flight_census3: [numeric] time in years from the drone flight date to the third census for that tree (yr)

54 ENVIRONMENTAL SCIENCES↗

Characterizing post-fire delayed tree mortality with remote sensing: sizing up the elephant in the room

Abstract Background Despite recent advances in understanding the drivers of tree-level delayed mortality, we lack a method for mapping delayed mortality at landscape and regional scales. Consequently, the extent, magnitude, and effects of delayed mortality on post-fire landscape patterns of burn severity are unknown. We introduce a remote sensing approach for mapping delayed mortality based on post-fire decline in the normalized burn ratio (NBR). NBR decline is defined as the change in NBR between the first post-fire measurement and the minimum NBR value up to 5 years post-fire for each pixel. We validate the method with high-resolution aerial photography from six wildfires in California, Oregon, and Washington, USA, and then compare the extent, magnitude, and effects of delayed mortality on landscape patterns of burn severity among fires and forest types. Results NBR decline was significantly correlated with post-fire canopy mortality (r 2 = 0.50) and predicted the presence of delayed mortality with 83% accuracy based on a threshold of 105 NBR decline. Plots with NBR decline greater than 105 were 23 times more likely to experience delayed mortality than those below the threshold (p < 0.001). Delayed mortality occurred across 6–38% of fire perimeters not affected by stand-replacing fire, generally affecting more areas in cold (22–41%) and wet (30%) forest types than in dry (1.7–19%) types. The total area initially mapped as unburned/very low-severity declined an average of 38.1% and generally persisted in smaller, more fragmented patches when considering delayed mortality. The total area initially mapped as high-severity increased an average of 16.2% and shifted towards larger, more contiguous patches. Conclusions Differences between 1- and 5-year post-fire burn severity maps depict dynamic post-fire mosaics resulting from delayed mortality, with variability among fires reflecting a range of potential drivers. We demonstrate that tree-level delayed mortality scales up to alter higher-level landscape patterns of burn severity with important implications for forest resilience and a range of fire-driven ecological outcomes. Our method can complement existing tree-level studies on drivers of delayed mortality, refine mapping of fire refugia, inform estimates of habitat and carbon losses, and provide a more comprehensive assessment of landscape and regional scale fire effects and trends.

Environmental Sciences & Ecology↗