Triggers of tree mortality under drought

Water availability is a major factor influencing the distribution and abundance of plant species in several communities (Allen and Breshears 1998; Engelbrecht et al. 2007). Most terrestrial ecosystems are subjected to seasonal droughts and global climate change models predict an increase in frequency and intensity of drought events in several regions of the world (Allen et al. 2010). These extreme drought events will likely induce widespread tree mortality and cause major shifts in plant community composition and ecosystem functioning (Nepstad et al. 2007). Understanding the diversity of plant drought responses and the mechanisms responsible for tree death is therefore of paramount importance to forecasting climate change impacts on the Earth’s biodiversity and ecosystem’s resilience (Anderegg et al. 2012; McDowell et al. 2008). These questions have recently become top priorities for global change research, because widespread and rapid drought-induced tree mortality is already an emerging global phenomenon, threatening the integrity of numerous ecosystems (Breshears et al. 2009; Phillips et al. 2010). Mediterranean-type ecosystems are especially vulnerable to climate change as global climate models predict rainfall reductions with a high degree of certainty in the near future for these regions (IPCC 2007). These wet-winter, dry-summer ecosystems harbor an enormous plant biodiversity. Of particular interest is south-western Australia, one of the world’s biodiversity hotspots and recently reported as one of the most vulnerable ecosystems to tipping points in Australia (Laurance et al. 2011). Can we predict the responses of plants from this species-rich flora to future drier climates? How resistant are different tree species to predicted increases in drought intensity and frequency? Can we predict with confidence changes in species distribution in response to climate change? Providing definite answers to these questions is neither simple nor straightforward, because of the numerous biotic and abiotic drivers that affect the performance of individual plants and the non-linear responses of ecosystems to climatic forcing. An ideal approach to address these questions would require a substantial multi-scale and multidisciplinary research effort. However, recent efforts have suggested that compiling functional trait variation along natural environmental gradients can substantially improve the mechanistic models used to Plant Soil DOI 10.1007/s11104-012-1508-9

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

Drought-induced tree mortality, which rapidly alters forest ecosystem composition, structure, and function, as well as the feedbacks between the biosphere and climate, has occurred worldwide over the past few decades, and is expected to increase pervasively as climate change progresses. The objectives of this review are to (1) highlight the likely ecological consequences of drought-induced tree mortality, (2) synthesize the hypotheses related to drought-induced tree mortality, (3) discuss the implications of current knowledge for modeling tree mortality processes under climate change, and (4) highlight future research needs. First, we emphasize the likely ecological consequences of tree mortality from ecosystem to biome to continental scales. We then document and criticize multiple non-exclusive tree mortality hypotheses (e.g., carbon starvation — carbon supply is less than carbon demand; and hydraulic failure — desiccation from failed water transport) from a more comprehensive ecological perspective. Next, we extend a forest decline concept model, Manion’s framework, by considering new emerging environmental conditions, for a more thorough understanding of the effects of climate change on forest decline. We find that an increase in drought frequency and (or) climate-change-type droughts may trigger increased background tree mortality rates and severe forest dieback events, accelerating species turnover and ecological regime shifts. The contribution of CO2 fertilization, rising temperature within the optimal growth range, and increased nitrogen deposition may defer or reduce this trend in tree mortality, but such contributions will vary between locations, species, and tree sizes. Multiple hypotheses proposed for drought-induced tree mortality are discussed, but coupling carbon and water cycles could help resolve the debate. The absence of a physiological understanding of tree mortality mechanisms limits the predictive ability of current models from stand-level process-based models to dynamic global vegetation models. We thus suggest that long-term observations, experiments, and models should be tightly interwoven during the research process to better forecast future climate changes and evaluate their impacts on forests.

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.

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere-atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.

Hydraulic determinants of drought-induced tree mortality and changes in tree abundance between two tropical forests with different water availability

Climate change is expected to increase the intensity of extreme precipitation events in Amazonia that in turn might produce more forest blowdowns associated with convective storms. Yet quantitative tree mortality associated with convective storms has never been reported across Amazonia, representing an important additional source of carbon to the atmosphere. Here we demonstrate that a single squall line (aligned cluster of convective storm cells) propagating across Amazonia in January, 2005, caused widespread forest tree mortality and may have contributed to the elevated mortality observed that year. Forest plot data demonstrated that the same year represented the second highest mortality rate over a 15‐year annual monitoring interval. Over the Manaus region, disturbed forest patches generated by the squall followed a power‐law distribution (scaling exponent α = 1.48) and produced a mortality of 0.3–0.5 million trees, equivalent to 30% of the observed annual deforestation reported in 2005 over the same area. Basin‐wide, potential tree mortality from this one event was estimated at 542 ± 121 million trees, equivalent to 23% of the mean annual biomass accumulation estimated for these forests. Our results highlight the vulnerability of Amazon trees to wind‐driven mortality associated with convective storms. Storm intensity is expected to increase with a warming climate, which would result in additional tree mortality and carbon release to the atmosphere, with the potential to further warm the climate system.

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.

TRIPLEX-Mortality model for simulating drought-induced tree mortality in boreal forests: Model development and evaluation

For tree seedlings in boreal forests, ectomycorrhizal (EM) fungal networks may promote, while root competition may impede establishment. Thus, disruption to EM fungal networks may decrease seedling establishment owing to the loss of positive interactions among neighbors. Widespread tree mortality can disrupt EM networks, but it is not clear whether seedling establishment will be limited by the loss of positive interactions or increased by the loss of negative interactions with surrounding roots. Depending upon the relative influence of these mechanisms, widespread tree mortality may have complicated consequences on seedling establishment, and in turn, the composition of future forests. To discern between these possible outcomes and the drivers of seedling establishment, we determined the relative importance of EM fungal networks, root presence, and the bulk soil on the establishment of lodgepole pine and white spruce seedlings along a gradient of beetle-induced tree mortality. We manipulated seedling contact with EM fungal networks and roots through the use of mesh-fabric cylinders installed in soils of lodgepole pine forests experiencing a range of overstorey tree mortality caused by mountain pine beetle. Lodgepole pine seedling survival was higher with access to EM fungal networks in undisturbed pine forests in comparison with that in beetle-killed stands. That is, overstorey tree mortality shifted fungal networks from being a benefit to a cost on seedling survival. In contrast, overstorey tree mortality did not change the relative strength of EM fungal networks, root presence and the bulk soil on survival and biomass of white spruce seedlings. Furthermore, the relative influence of EM fungal networks, root presence, and bulk soils on foliar N and P concentrations was highly contingent on seedling species and overstorey tree mortality. Our results highlight that following large-scale insect outbreak, soil-mediated processes can enable differential population growth of two common conifer species, which may result in species replacement in the future.

The effect of drought on wood-boring in trees and saplings in tropical rainforests

Natural disturbances maintain biological diversity and landscape heterogeneity and initiate ecosystem renewal and reorganization. However, the severity, frequency, and extent of many disturbances have increased substantially in recent decades as the result of anthropogenic climate change. Disturbances can be discrete, short-duration events, such as wildfires or hurricanes, or can exert persistent, cumulative stresses on an ecosystem (for example, ongoing warming of ocean or land surface temperatures). Landscape and ecosystem impacts can occur from a single disturbance, from several disturbances acting independently, or from the interactions of multiple, linked disturbances. Key, climate-related disturbances affecting global biomes and ecosystems include shifting temperature and hydrologic regimes (for example, warming surface temperatures and increasing aridity), increased frequency and magnitude of extreme events such as heatwaves, severe droughts, storms, and hurricanes, warming-induced permafrost thaw, and heightened wildfire activity and insect-caused tree mortality. For ecosystems and landscapes, the consequences of climate-amplified disturbance include forced poleward and upward movement of plant and animal species, widespread tree mortality and reduced forest productivity, changes in plant community structure and species distributions, reduced biodiversity, increased erosion, debris flows, wetland dynamism, declining sea ice extent, more frequent storm-driven tides and saltwater intrusion, and increased landscape flammability.

Drought stress causes widespread forest mortality globally, for instance recently in temperate forests of Central Europe during and after the extremely dry summers in 2018, 2019 and 2022. Trees may die in consequence of hydraulic damage from xylem embolism, but also due to long-term effects caused by allocation into repair and adaptation that may deplete carbon (C) reserves, reduce competition strength and lower resistance to subsequent insect and pathogen infestations. We review implementations of drought-induced tree and forest mortality in ecosystem models and test different implementations in LandscapeDNDC, a terrestrial ecosystem model designed for simulations of the C and nitrogen cycles at site and regional scales. Based on tree hydraulic processes recently integrated into the model, we simulate tree mortality either a) when a threshold in xylem hydraulic conductivity loss is exceeded, or b) when tree water storage is depleted. In addition, we consider c) tree mortality as a result of depleted C reserves and low growth efficiency caused by drought legacy effects. Direct and legacy effects of drought stress on tree mortality rates are parameterized for common European temperate tree species (Fagus sylvatica, Picea abies, Pinus sylvestris, Quercus robur). We evaluate our simulations of drought-related tree mortality rates by comparing them to estimates from forest inventory and remote sensing approaches covering recent drought events. An improved modelling of direct and lagged drought-induced forest mortality is essential to understand the response of the vegetation C cycle to climate change and the options of forest management to increase the resistance of European temperate forests to drought.

Tree characteristics and drought severity modulate the growth resilience of natural Mongolian pine to extreme drought episodes

Forest restoration treatments seek to increase resilience to wildfire and a changing climate while avoiding negative impacts to the ecosystem. The extent and intensity of treatments are often constrained by operational considerations and concerns over uncertainty in the trade-offs of addressing different management goals. The recent (2012–15) extreme drought in California, USA, resulted in widespread tree mortality, particularly in the southern Sierra Nevada, and provided an opportunity to assess the effects of restoration treatments on forest resilience to drought. We assessed changes in mixed-conifer forest structure following thinning and understorey burning at the Kings River Experimental Watersheds in the southern Sierra Nevada, and how treatments, topography and forest structure related to tree mortality in the recent drought. Treatments had negligible effect on basal area, tree density and canopy cover. Following the recent drought, average basal area mortality within the watersheds ranged from 5 to 26% across riparian areas and 12 to 44% across upland areas, with a range of 0 to 95% across all plots. Tree mortality was not significantly influenced by restoration treatments or topography. Our results suggest that the constraints common to many restoration treatments may limit their ability to mitigate the impacts of severe drought.