Drought-induced Forest Mortality Research Articles

Impact of drought-induced forest mortality on terrestrial carbon cycle from remote sensing perspective

Impact of drought-induced forest mortality on terrestrial carbon cycle from remote sensing perspective

Higher site productivity and stand age enhance forest susceptibility to drought-induced mortality

Warmer and drier conditions increase forest mortality worldwide. At the same time, nitrogen deposition, longer growing seasons and higher atmospheric CO2 concentrations may increase site productivity accelerating forest growth. However, tree physiological studies suggest that increased site productivity can also have adverse effects, reducing adaptation to drought. Understanding such intricate interactions that might foster tree mortality is essential for designing activities and policies aimed at preserving forests and the ecosystem services they provide. This study shows how site factors and stand features affect the susceptibility of Scots pine to drought-induced stand-level mortality. We use extensive forest data covering 750,000 ha, including 47,450 managed Scots pine stands, of which 2,547 were affected by mortality during the drought in 2015–2019. We found that the oldest and most dense stands growing on the most productive sites showed the highest susceptibility to enhanced mortality during drought. Our findings suggest that increasing site productivity may accelerate the intensity and prevalence of drought-induced forest mortality. Therefore, climate change may increase mortality, particularly in old and high-productive forests. Such exacerbated susceptibility to mortality should be considered in forest carbon sink projections, forest management, and policies designed to increase resilience and protect forest ecosystems.

Does competition management improve the ecophysiological response to water shortage of mixed woodland species of North Patagonia?

Globally, the increasing forests vulnerability and drought-induced forest mortality events, extended to a larger scale, may have the potential to rapidly alter forest functioning and the provision of ecosystem services. Patagonian forests in Southern South America constitute an extensive reservoir of wildlife, but at the same time they have high productive potential. These low-mixed forests are composed of co-dominant species that are highly disturbed by local population, both for cattle raising and wood extraction; and different efforts are being dedicated to improve their management. Very little is known about the response of their different woody species to climatic variation, particularly to severe drought in particular, that may guide management strategies to improve productivity and resilience in context of drought stress and climate change. The present study aimed at characterizing the physiological strategies of response to drought to identify the most vulnerable and most resilient species to drought and how they respond under different competition levels. The response of four species of the Andean Patagonian low mixed forest was quantified at three moments of the 2018–2019 growing season in three sites through measurements of the net carbon exchange, vulnerability to xylem cavitation, tissue water relations, and damage of the photosynthetic apparatus. Additionally, records of average monthly values of precipitation and air temperature were used to calculate a standardized precipitation-evapotranspiration index. The normal climatic conditions of NW Patagonia region are characterized by water deficits during the spring-summer growing season, but the studied season presented lower values of standardized precipitation-evapotranspiration index respect to the historical mean values, indicating drier than normal conditions. Analysis of the tissue water relations showed that, in all cases, competition had no effect over the different studied parameters. There were species-specific differences in the models fitted to the vulnerability to cavitation curves; as a whole, all the species differed from each other. Overall, for all the physiological response traits studied, the Site, Species, and Season factors were important predictors in the minimum adequate model. Although there is an effect of competition on the individual annual growth, that effect is not evidenced in physiological variables of punctual measurement. Our results suggest that the effect of summer drought cannot be modulated by density management. The productivity of the species of interest can be improved, but not their adaptability, at least to severe events such as the one of the year of study.

Age-related patterns and climatic driving factors of drought-induced forest mortality in Northeast China

Given the significantly increasing frequency and intensity of droughts in Northeast China under climate change, it is necessary to understand drought-induced mortality in forest ecosystems. The magnitude of drought-induced forest mortality strongly depends on climate variability and forest age, which are both important driving factors of tree mortality. However, the age-related patterns and climatic driving factors of drought-induced forest mortality at large scales are poorly understood. This study identified the age-related pattern and the key climatic driving factors of drought-induced mortality for 17 dominant tree species in Northeast China using rain-use efficiency (RUE), standardized precipitation evapotranspiration index (SPEI), and tree-list information (species and diameter for every tree). Considering that climate variability and forest age are important factors of drought-induced mortality, species-specific analysis was conducted for describing age-dependent mortality patterns. The results showed that the mean annual rate of the drought-induced mortality of forests in Northeast China was 0.49%, with relatively high mortality rates in the Changbai Mountains and the eastern part of Liaoning Province. The sensitivity of age-dependent mortality patterns to climatic drivers of drought exhibited considerable variability. Resolving the contributions of precipitation deficits and heatwaves on the drought-induced mortality of trees, almost 91.60% of the forest region in Northeast China was found to be primarily affected by heatwaves (temperature anomalies), suggesting the important role of temperature extremes in forest mortality. The findings provide deeper insight into the mechanisms behind the species-specific formation of age-dependent patterns in Northeast China. This study provides a basis for the formulation of drought adaptation measures for forest species across successional stages and highlights the potential of remote sensing indices in identifying the patterns and climatic drivers of large-scale drought-induced forest mortality.

Conduit position and connectivity affect the likelihood of xylem embolism during natural drought in evergreen woodland species.

Hydraulic failure is considered a main cause of drought-induced forest mortality. Yet, we have a limited understanding of how the varying intensities and long time scales of natural droughts induce and propagate embolism within the xylem. X-ray computed tomography (microCT) images were obtained from different aged branch xylem to study the number, size and spatial distribution of in situ embolized conduits among three dominant tree species growing in a woodland community. Among the three studied tree species, those with a higher xylem vulnerability to embolism (higher water potential at 50 % loss of hydraulic conductance; P50) were more embolized than species with lower P50. Within individual stems, the probability of embolism was independent of conduit diameter but associated with conduit position. Rather than the occurrence of random or radial embolism, we observed circumferential clustering of high and low embolism density, suggesting that embolism spreads preferentially among conduits of the same age. Older xylem also appeared more likely to accumulate embolisms than young xylem, but there was no pattern suggesting that branch tips were more vulnerable to cavitation than basal regions. The spatial analysis of embolism occurrence in field-grown trees suggests that embolism under natural drought probably propagates by air spreading from embolized into neighbouring conduits in a circumferential pattern. This pattern offers the possibility to understand the temporal aspects of embolism occurrence by examining stem cross-sections.

Eco-hydrological responses to recent droughts in tropical South America

This study assesses the ecohydrological effects of recent meteorological droughts in tropical South America based on multiple sources of data, and investigates the possible mechanisms underlying the drought response and recovery of different ecohydrological systems. Soil drought response and recovery lag behind the meteorological drought, with delays longer in the dry region (Nordeste) than in the wet region (Amazonia), and longer in deep soil than in shallow soil. Evapotranspiration (ET) and vegetation in Nordeste are limited by water under normal conditions and decrease promptly in response to the onset of shallow soil drought. In most of the Amazon where water is normally abundant, ET and vegetation indices follow an increase-then-decrease pattern, increase at the drought onset due to increased sunshine and decrease when the drought is severe enough to cause a shift from an energy-limited regime to a water-limited regime. After the demise of meteorological droughts, ET and vegetation rapidly recover in Nordeste with the replenishment of shallow soil moisture (SM), but take longer to recover in southern Amazon due to their dependence on deep SM storage. Following severe droughts, the negative anomalies of ET and vegetation indices in southern Amazon tend to persist well beyond the end of soil drought, indicating drought-induced forest mortality that is slow to recover from. Findings from this study may have implications on the possibility of a future forest dieback as drought is projected to become more frequent and more severe in a warmer climate.

Plant water content integrates hydraulics and carbon depletion to predict drought-induced seedling mortality.

Widespread drought-induced forest mortality (DIM) is expected to increase with climate change and drought, and is expected to have major impacts on carbon and water cycles. For large-scale assessment and management, it is critical to identify variables that integrate the physiological mechanisms of DIM and signal risk of DIM. We tested whether plant water content, a variable that can be remotely sensed at large scales, is a useful indicator of DIM risk at the population level. We subjected Pinus ponderosa Douglas ex C. Lawson seedlings to experimental drought using a point of no return experimental design. Periodically during the drought, independent sets of seedlings were sampled to measure physiological state (volumetric water content (VWC), percent loss of conductivity (PLC) and non-structural carbohydrates) and to estimate population-level probability of mortality through re-watering. We show that plant VWC is a good predictor of population-level DIM risk and exhibits a threshold-type response that distinguishes plants at no risk from those at increasing risk of mortality. We also show that plant VWC integrates the mechanisms involved in individual tree death: hydraulic failure (PLC), carbon depletion across organs and their interaction. Our results are promising for landscape-level monitoring of DIM risk.

Multi-temporal influence of vegetation on soil respiration in a drought-affected forest

Abstract: Aboveground plant activity influences fine roots and rhizosphere activity, which is reflected on soil respiration (SR). However, it is still unclear and poorly understood the nature of plant activity control over SR, especially under drought conditions. We studied the plant activity-SR relationship at different timescales in a water-limited mixed Mediterranean forest where Scots pines (Pinus sylvestris L.) are undergoing drought-induced die-off and are being replaced by the more drought-resistant Holm oak (Quercus ilex L.). Half-hourly sap flow (SF), as a proxy of photosynthesis, coupled with measures of SR using solid-state CO2 sensors, were monitored during nine months in four different trees, representative of the diversity and health condition of the forest. SF was strongly associated with SR at both daily and seasonal timescales. At daily timescales, almost no lags were found between SF and SR, indicating a fast control of photosynthesis on SR. However, the association between SF and SR weakened during the summer drought. These temporal patterns were not constant across the trees representing the die-off and replacement processes. SR beneath living pines was highly controlled by SF at daily scale, whereas Holm oak seemed to be more controlled by SF at seasonal scale. The relationship between SF and SR measured beneath dead pine and Holm oak at the daily and seasonal scales was consistent with the colonization of soil gaps by holm oak roots following Scots pine death and suggests that surviving Scots pines are unable to expand their root system in these gaps. Our results collectively show how drought modulates the link between canopy photosynthesis and soil respiration, and increase our understanding on how belowground processes may be affected by the successional dynamics following drought-induced forest mortality.

Teresa Rosas.

Teresa Rosas.

Remotely sensed predictors of conifer tree mortality during severe drought

Widespread, drought-induced forest mortality has been documented on every forested continent over the last two decades, yet early pre-mortality indicators of tree death remain poorly understood. Remotely sensed physiological-based measures offer a means for large-scale analysis to understand and predict drought-induced mortality. Here, we use laser-guided imaging spectroscopy from multiple years of aerial surveys to assess the impact of sustained canopy water loss on tree mortality. We analyze both gross canopy mortality in 2016 and the change in mortality between 2015 and 2016 in millions of sampled conifer forest locations throughout the Sierra Nevada mountains in California. On average, sustained water loss and gross mortality are strongly related, and year-to-year water loss within the drought indicates subsequent mortality. Both relationships are consistent after controlling for location and tree community composition, suggesting that these metrics may serve as indicators of mortality during a drought.

Tree mortality across biomes is promoted by drought intensity, lower wood density and higher specific leaf area.

Drought events are increasing globally, and reports of consequent forest mortality are widespread. However, due to a lack of a quantitative global synthesis, it is still not clear whether drought-induced mortality rates differ among global biomes and whether functional traits influence the risk of drought-induced mortality. To address these uncertainties, we performed a global meta-analysis of 58 studies of drought-induced forest mortality. Mortality rates were modelled as a function of drought, temperature, biomes, phylogenetic and functional groups and functional traits. We identified a consistent global-scale response, where mortality increased with drought severity [log mortality (trees trees-1 year-1 ) increased 0.46 (95% CI=0.2-0.7) with one SPEI unit drought intensity]. We found no significant differences in the magnitude of the response depending on forest biomes or between angiosperms and gymnosperms or evergreen and deciduous tree species. Functional traits explained some of the variation in drought responses between species (i.e. increased from 30 to 37% when wood density and specific leaf area were included). Tree species with denser wood and lower specific leaf area showed lower mortality responses. Our results illustrate the value of functional traits for understanding patterns of drought-induced tree mortality and suggest that mortality could become increasingly widespread in the future.

Carbohydrate dynamics and mortality in a piñon-juniper woodland under three future precipitation scenarios.

Drought-induced forest mortality is an increasing global problem with wide-ranging consequences, yet mortality mechanisms remain poorly understood. Depletion of non-structural carbohydrate (NSC) stores has been implicated as an important mechanism in drought-induced mortality, but experimental field tests are rare. We used an ecosystem-scale precipitation manipulation experiment to evaluate leaf and twig NSC dynamics of two co-occurring conifers that differ in patterns of stomatal regulation of water loss and recent mortality: the relatively desiccation-avoiding piñon pine (Pinus edulis) and the relatively desiccation-tolerant one-seed juniper (Juniperus monosperma). Piñon pine experienced 72% mortality after 13-25 months of experimental drought and juniper experienced 20% mortality after 32-47 months. Juniper maintained three times more NSC in the foliage than twigs, and converted NSC to glucose and fructose under drought, consistent with osmoregulation requirements to maintain higher stomatal conductance during drought than piñon. Despite these species differences, experimental drought caused decreased leaf starch content in dying trees of both species (P < 0.001). Average dry-season leaf starch content was also a good predictor of drought-survival time for both species (R(2) = 0.93). These results, along with observations of drought-induced reductions to photosynthesis and growth, support carbon limitation as an important process during mortality of these two conifer species.

Feature: Improving our knowledge of drought‐induced forest mortality through experiments, observations, and modeling

Feature: Improving our knowledge of drought‐induced forest mortality through experiments, observations, and modeling

Drought-induced tree species replacement is reflected in the spatial variability of soil respiration in a mixed Mediterranean forest

As episodes of drought-induced forest mortality are being increasingly reported worldwide and may become more frequent in the future as a result of climate change, it is essential to characterize their functional implications in terms of ecosystem carbon and water fluxes. We investigated the spatial variability of soil respiration in a mixed Mediterranean forest located on rugged terrain, where Scots pine (Pinus sylvestris) is affected by drought-induced dieback and appears to have been replaced by Holm oak (Quercus ilex) as the dominant tree species. Soil respiration was measured in spring 2010 on two plots (16.2×16.2m) using a static closed chamber method (soda lime technique) and a systematic sampling (1.8-m grid) including 100 points per plot. Biotic and abiotic variables, such as soil moisture, soil temperature, soil organic matter content, stoniness, pH, fine root C:N ratio and biomass, tree basal area and tree species and health condition of nearest neighbouring tree were also recorded. Our results showed that the spatial variability of soil respiration under optimal environmental conditions (spring) was high and showed no spatial autocorrelation on the scale studied (1–18m). A mixed-effects model applied to explain the spatial variability of soil respiration indicated that only the variables related to forest structure (i.e., health condition and basal area) explained any of the observed variability of soil respiration (R2=0.45). Our model revealed that soil respiration was highest in soils close to dead pines and under Holm oak trees, suggesting that tree mortality and species replacement of pine trees by Holm oak may lead to higher soil respiration fluxes. The direct effect of tree mortality on soil respiration may be a transitory response caused by fine root mortality. Furthermore, the fact that tree species replacement as a result of drought-induced die-off is accompanied by concomitant changes in soil respiration has important implications for soil and ecosystem carbon balance.

Drought characteristics' role in widespread aspen forest mortality across Colorado, USA

Globally documented widespread drought-induced forest mortality has important ramifications for plant community structure, ecosystem function, and the ecosystem services provided by forests. Yet the characteristics of drought seasonality, severity, and duration that trigger mortality events have received little attention despite evidence of changing precipitation regimes, shifting snow melt timing, and increasing temperature stress. This study draws upon stand level ecohydrology and statewide climate and spatial analysis to examine the drought characteristics implicated in the recent widespread mortality of trembling aspen (Populus tremuloides Michx.). We used isotopic observations of aspen xylem sap to determine water source use during natural and experimental drought in a region that experienced high tree mortality. We then drew upon multiple sources of climate data to characterize the drought that triggered aspen mortality. Finally, regression analysis was used to examine the drought characteristics most associated with the spatial patterns of aspen mortality across Colorado. Isotopic analysis indicated that aspens generally utilize shallow soil moisture with little plasticity during drought stress. Climate analysis showed that the mortality-inciting drought was unprecedented in the observational record, especially in 2002 growing season temperature and evaporative deficit, resulting in record low shallow soil moisture reserves. High 2002 summer temperature and low shallow soil moisture were most associated with the spatial patterns of aspen mortality. These results suggest that the 2002 drought subjected Colorado aspens to the most extreme growing season water stress of the past century by creating high atmospheric moisture demand and depleting the shallow soil moisture upon which aspens rely. Our findings highlight the important role of drought characteristics in mediating widespread aspen forest mortality, link this aspen die-off to regional climate change trends, and provide insight into future climate vulnerability of these forests.