Local carbon reserves are insufficient for phloem terpene induction during drought in Pinus edulis in response to bark beetle-associated fungi.

Stomatal closure during drought inhibits carbon uptake and may reduce a tree's defensive capacity. Limited carbon availability during drought may increase a tree's mortality risk, particularly if drought constrains trees' capacity to rapidly produce defenses during biotic attack. We parameterized a new model of conifer defense using physiological data on carbon reserves and chemical defenses before and after a simulated bark beetle attack in mature Pinus edulis under experimental drought. Attack was simulated using inoculations with a consistent bluestain fungus (Ophiostoma sp.) of Ips confusus, the main bark beetle colonizing this tree, to induce a defensive response. Trees with more carbon reserves produced more defenses but measured phloem carbon reserves only accounted for c. 23% of the induced defensive response. Our model predicted universal mortality if local reserves alone supported defense production, suggesting substantial remobilization and transport of stored resin or carbon reserves to the inoculation site. Our results show that de novo terpene synthesis represents only a fraction of the total measured phloem terpenes in P. edulis following fungal inoculation. Without direct attribution of phloem terpene concentrations to available carbon, many studies may be overestimating the scale and importance of de novo terpene synthesis in a tree's induced defense response.

Using climate envelopes and earth system model simulations for assessing climate change induced forest vulnerability.

Changing climatic conditions threaten forest ecosystems. Drought, disease and infestation, are leading to forest die-offs which cause substantial economic and ecological losses. In central Europe, this is especially relevant for commercially important coniferous tree species. This study uses climate envelope exceedance (CEE) to approximate species risk under different future climate scenarios. To achieve this, we used current species presence-absence and historical climate data, coupled with future climate scenarios from various Earth System Models. Climate scenarios tended towards drier and warmer conditions, causing strong CEEs especially for spruce. However, we show that annual averages of temperature and precipitation obscure climate extremes. Including climate extremes reveals a broader increase in CEEs across all tree species. Our study shows that the consideration of climate extremes, which cannot be adequately reflected in annual averages, leads to a different assessment of the risk of forests and thus the options for adapting to climate change.

Effects of canopy gaps on microclimate, soil biological activity and their relationship in a European mixed floodplain forest.

Forest canopy gaps can influence understorey microclimate and ecosystem functions such as decomposition. Gaps can arise from silviculture or tree mortality, increasingly influenced by climate change. However, to what degree canopy gaps affect the buffered microclimate in the understorey under macroclimatic changes is unclear. We, therefore, investigated the effect of forest gaps differing in structure and size (25 gaps: single tree gaps up to 0.67 ha cuttings) on microclimate and soil biological activity compared to closed forest in a European mixed floodplain forest. During the investigation period in the drought year 2022 between May and October, mean soil moisture and temperature as well as soil and air temperature fluctuations increased with increasing openness. In summer, the highest difference of monthly means between cuttings and closed forest in the topsoil was 3.98 ± 9.43 % volumetric moisture and 2.05 ± 0.89 °C temperature, and in the air at 30 cm height 0.61 ± 0.35 °C temperature. For buffering, both the over- and understorey tree layers appeared as relevant with a particularly strong influence of understorey density on soil temperature. Three experiments, investigating soil biological activity by quantifying decomposition rates of tea and wooden spatulas as well as mesofauna feeding activity with bait-lamina stripes, revealed no significant differences between gaps and closed forest. However, we found a positive significant effect of mean soil temperature on feeding activity throughout the season. Although soil moisture decreased during this period, it showed no counteracting effect on feeding activity. Generally, very few significant relationships were observed between microclimate and soil biological activity in single experiments. Despite the dry growing season, decomposition rates remained high, suggesting temperature had a stronger influence than soil moisture. We conclude that the microclimatic differences within the gap gradient of our experiment were not strong enough to affect soil biological activity considerably.

Meta-analysis of livestock effects on tree regeneration in oak agroforestry systems.

Livestock grazing occupies over a quarter of terrestrial land and is prevalent to agroforestry ecosystems, potentially affecting the survival, growth, and density of trees' early developmental stages, such as seeds, seedlings, and saplings. To address the effects of livestock on tree recruitment in the face of ongoing debates about their impacts, we conducted a 33-year meta-analysis in Quercus-dominated agroforestry systems. Our analysis revealed a consistently negative effect of livestock on oak acorns, seedlings, and saplings. Significantly, livestock body size influenced oak regeneration, with small-sized livestock, notably sheep and goats, having a more pronounced negative impact compared to mixed-size systems, mainly involving cattle and sheep. The effects of small-sized livestock were markedly detrimental on acorn survival and seedling/sapling density, although no studies eligible for meta-analysis examined large livestock impacts on acorns. Overall, mixed-size livestock systems, often involving cattle and sheep, lessen the negative effects. Our findings indicate that the body size and foraging behaviors of livestock should be considered for the ecological sustainability of the tree component in agroforestry systems. While protective measures have long been integral to well-managed agroforestry systems, our results underscore the importance of integrating diverse livestock sizes and applying specific protective strategies, particularly for acorns and saplings, to further refine these practices. Future research should expand to underrepresented regions and livestock types to refine global agroforestry management practices.

Hydraulic properties and drought response of a tropical bamboo (Cephalostachyum pergracile).

Bamboo plants are an essential component of tropical ecosystems, yet their vulnerability to climate extremes, such as drought, is poorly understood due to limited knowledge of their hydraulic properties. Cephalostachyum pergracile, a commonly used tropical bamboo species, exhibited a substantially higher mortality rate than other co-occurring bamboos during a severe drought event in 2019, but the underlying mechanisms remain unclear. This study investigated the leaf and stem hydraulic traits related to drought responses, including leaf-stem embolism resistance (P50leaf; P50stem) estimated using optical and X-ray microtomography methods, leaf pressure-volume and water-releasing curves. Additionally, we investigated the seasonal water potentials, native embolism level (PLC) and xylem water source using stable isotope. We found that C. pergracile exhibited strong resistance to embolism, showing low P50leaf, P50stem, and turgor loss point, despite its rapid leaf water loss. Interestingly, its leaves displayed greater resistance to embolism than its stem, suggesting a lack of effective hydraulic vulnerability segmentation (HVS) to protect the stem from excessive xylem tension. During the dry season, approximately 49% of the water was absorbed from the upper 20-cm-deep soil layer. Consequently, significant diurnal variation in leaf water potentials and an increase in midday PLC from 5.87 ± 2.33% in the wet season to 12.87 ± 4.09% in the dry season were observed. In summary, this study demonstrated that the rapid leaf water loss, high reliance on surface water, and a lack of effective HVS in C. pergracile accelerated water depletion and increased xylem embolism even in the typical dry season, which may explain its high mortality rate during extreme drought events in 2019.

Positive effects of tree species diversity on productivity switch to negative after severe drought mortality in a temperate forest experiment.

The synthesis of a large body of evidence from field experiments suggests more diverse plant communities are more productive as well as more resistant to the effects of climatic extremes like drought. However, this view is strongly based on data from grasslands due to the limited empirical evidence from tree diversity experiments. Here we report on the relationship between tree diversity and productivity over 10 years in a field experiment established in 2005 that was then affected by the 2018 mega-drought in central Europe. Across a number of years, tree species diversity and productivity were significantly positively related; however, the slope switched to negative in the year of the drought. Net diversity effects increased through time, with complementarity effects making greater contributions to the net diversity effect than selection effects. Complementarity effects were clearly positive in three- and five-species mixtures before the drought (2012-2016) but were found to decrease in the year of the drought. Selection effects were clearly positive in 2016 and remained positive in the drought year 2018 in two-, three-, and five-species mixtures. The survival of Norway spruce (Picea abies) plummeted in response to the drought, and a negative relationship between species diversity and spruce survival was found. Taken together, our findings suggest that tree diversity per se may not buffer communities against the impacts of extreme drought and that tree species composition and the drought tolerance of tree species (i.e., species identity) will be important determinants of community productivity as the prevalence of drought increases.

Unpacking the point of no return under drought in poplar: insight from stem diameter variation.

A specific, robust threshold for drought-induced tree mortality is needed to improve the prediction of forest dieback. Here, we tested the relevance of continuous measurements of stem diameter variations for identifying such a threshold, their relationship with hydraulic and cellular damage mechanisms, and the influence of growth conditions on these relationships. Poplar saplings were grown under well-watered, water-limited, or light-limited conditions and then submitted to a drought followed by rewatering. Stem diameter was continuously measured to investigate two parameters: the percentage loss of diameter (PLD) and the percentage of diameter recovery (DR) following rewatering. Water potentials, stomatal conductance, embolism, and electrolyte leakage were also measured, and light microscopy allowed investigating cell collapse induced by drought. The water release observed through loss of diameter occurred throughout the drought, regardless of growth conditions. Poplars did not recover from drought when PLD reached a threshold and this differed according to growth conditions but remained linked to cell resistance to damage and collapse. Our findings shed new light on the mechanisms of drought-induced tree mortality and indicate that PLD could be a relevant indicator of drought-induced tree mortality, regardless of the growth conditions.

Stem heating results in hydraulic dysfunction in Symplocos tinctoria: implications for post-fire tree death.

Fire-induced heating of stems can impair plant water transport by deforming xylem and increasing vulnerability to cavitation, but it is not clear whether these effects can result in tree death, or how quickly this may occur. In field experiments, we heated stems of Symplocos tinctoria (L.) L'Hér saplings to 90 °C using a thin-film resistive heater, and we monitored stomatal conductance, leaf water potential, sap flow and hydraulic conductivity until stem death. Sap flow and stomatal conductance declined quickly after heating, while whole-plant hydraulic conductance and leaf water potential remained high for the first week. In fact, leaf water potential increased during the first days after heating, indicating that stomatal closure was not initially caused by leaf water deficit induced by impaired water transport. After 1 week, leaf water potential and whole-plant conductance declined below unheated controls, while stomatal conductance and sap flow continued declining, approaching zero after 2 weeks. To better understand the cause of these declines, we directly measured hydraulic conductivity of heated stems. Stems underwent a progressive decline in conductivity after heating, and by the time that samples were severely wilted or desiccated, the heated portion of stems had little or no conductivity. Importantly, conductivity of heated stems was not recovered by flushing stems to remove embolisms, suggesting the existence of physical occlusions. Scanning electron micrographs did not reveal deformed cell walls, nor did it identify alternative causes of blockages. These results reveal that stem heating can result in xylem dysfunction and mortality, but neither response is immediate. Dysfunction was likely caused by wound responses rather than embolism, but improved understanding of the mechanisms of heat-induced hydraulic failure is needed.

Mechanistic links between physiology and spectral reflectance enable previsual detection of oak wilt and drought stress.

Tree mortality due to global change-including range expansion of invasive pests and pathogens-is a paramount threat to forest ecosystems. Oak forests are among the most prevalent and valuable ecosystems both ecologically and economically in the United States. There is increasing interest in monitoring oak decline and death due to both drought and the oak wilt pathogen (Bretziella fagacearum). We combined anatomical and ecophysiological measurements with spectroscopy at leaf, canopy, and airborne levels to enable differentiation of oak wilt and drought, and detection prior to visible symptom appearance. We performed an outdoor potted experiment with Quercus rubra saplings subjected to drought stress and/or artificially inoculated with the pathogen. Models developed from spectral reflectance accurately predicted ecophysiological indicators of oak wilt and drought decline in both potted and field experiments with naturally grown saplings. Both oak wilt and drought resulted in blocked water transport through xylem conduits. However, oak wilt impaired conduits in localized regions of the xylem due to formation of tyloses instead of emboli. The localized tylose formation resulted in more variable canopy photosynthesis and water content in diseased trees than drought-stressed ones. Reflectance signatures of plant photosynthesis, water content, and cellular damage detected oak wilt and drought 12 d before visual symptoms appeared. Our results show that leaf spectral reflectance models predict ecophysiological processes relevant to detection and differentiation of disease and drought. Coupling spectral models that detect physiological change with spatial information enhances capacity to differentiate plant stress types such as oak wilt and drought.

"Mortality, or not mortality, that is the question ": How to Treat Removals in Tree Survival Analysis of Central European Managed Forests.

Tree mortality is an objective forest health criterion and is particularly suitable for long-term and large-scale studies of forest condition. However, it is impossible to determine actual tree mortality in Central European managed forests where trees are removed for various reasons. In this case, the only way to approximate tree mortality is to define the range in which it occurs. This can be carried out by including in the mortality calculations either dead trees that remain in the stand at the end of the assessment period or additionally trees that have been removed from the stand. We used data from the annual forest monitoring surveys in Poland from 2009 to 2022 for pine, spruce, oak and birch to perform a survival analysis in which we included all removals or sanitary cuttings either as censored or complete observations. The differences between the calculated mortality rates were significant, indicating the importance of how removals are treated in the analysis. To assess which method used for mortality calculation was more appropriate, we compared values for last recorded defoliation and severity of damage from live, dead and thinned or salvaged trees. For all species studied, significant differences were found between dead trees or trees removed by sanitation cuts and living trees or trees removed by thinning, suggesting that not only dead trees remaining in the forest, but also trees removed by sanitation cuts, should be considered when calculating mortality in managed stands. We also recommend the use of survival analysis in forest monitoring as a routine method for assessing the health of stands.

Impacts of a severe storm on carbon accumulation in coarse woody debris within a secondary Atlantic Forest fragment in Brazil.

The alarming increase in extreme weather events, such as severe storms with torrential rain and strong winds, is a direct result of climate change. These events have led to discernible shifts in forest structure and the carbon cycle, primarily driven by a surge in tree mortality. However, the impacts caused by these severe storms on the production and carbon increment from coarse woody debris (CWD) are still poorly understood, especially in the Brazilian Atlantic Forest. Thus, the goal proposed by the study was to quantify the CWD volume, necromass, and carbon stock before and after the occurrence of a severe storm and to determine the importance of spatial, structural, and qualitative variables of trees in the CWD carbon increment. The increase in carbon by the storm was 2.01 MgC ha-1, with a higher concentration in the CWD less decomposed and smaller diameter class. The forest fragment plots showed distinct increments (0.05-0.35 MgC), being influenced by spatial (elevation, declivity, and slope angle) structural (basal area) and qualitative factors (trunk quality and tree health), intrinsic to the forest. Thus, it is concluded that severe storms cause a large increase in carbon in CWD, making it essential to understand the susceptibility of forests to the action of intense rains and strong winds to model and monitor the future impacts of these extreme weather events on Atlantic Forest and other tropical forests in the world.

Evaluating urban park ecosystem services and modeling improvement scenarios.

Three ecosystem services of the 25 public parks in Bangkok, including carbon sequestration, avoided runoff, four air pollutant removals (CO, NO2, PM10, and PM2.5), and the relevant monetary values, were determined using i-Tree Eco software. Two modeling scenarios (MS) including MS1 (no greening improvement) and MS2 (improvement by increasing either green area or tree planting, or both in the parks) with tree annual mortality rates (AMR) of 1 and 3% were developed to forecast the parks' ecosystem services for 50 years after 2020 (2021-2071). The results revealed the synergistic interactions of the different tree planting specifications (MS1 and MS2) and tree mortality rates on the parks' ecosystem services. For MC2 with the assigned 1% AMR, the parks' optimal ecosystem services were obtained and the average annual monetary value (0.55 million USD) of the total ecosystem services of the 25 parks over the 50-year forecast was 150% higher than that (0.22 million USD) in 2020. Based on MS1 and MS2, tree rotations should be conducted in the parks after 2057 and 2065, respectively, for the low tree AMR (≤1%) but not later than 2041 and 2043, respectively, for the higher tree AMR.

Different hydraulic and photosynthetic responses to summer drought between newly sprouted and established Moso bamboo culms.

The subtropical regions in China are prone to recurrent summer droughts induced by the Western Pacific Subtropical High-Pressure, which has induced the death of tens of millions of culms of Moso bamboo (Phyllostachys edulis (Carriere) J. Houzeau), a widely distributed giant bamboo with high economic and ecological values. In the future, the intensity and frequency of the summer drought are projected to increase in these areas due to global climate change, which may lead to significant age-specific mortality of Moso bamboo. So far, it is still unclear about the age-specific response mechanisms of hydraulic traits and carbon balance of Moso bamboo when it is suffering to an ongoing summer drought. This study aimed to investigate the hydraulic and photosynthetic responses of newly sprouted (1 year old) and established (2-5 years old) culms of Moso bamboo to summer drought, which was manipulated by throughfall reduction in Lin'an of Zhejiang. The results showed that both newly sprouted and established culms had a gradually weakening hydraulic conductivity and photosynthesis during the whole drought process. In the early stage of the manipulated drought, the established culms had more loss of hydraulic conductivity than the newly sprouted culms. However, the newly sprouted culms had significant more loss of hydraulic conductivity and lower photosynthetic rates and stomatal conductance in the middle and late stages of the manipulated drought. The results suggest that the newly sprouted culms were more susceptible to summer drought than established culms due to the combined effects of hydraulic damage and photosynthetic restriction, explaining why the newly sprouted culms have higher mortality than elder culms when subjected to extreme drought. These findings provided insights into the mechanisms of Moso bamboo's age-specific drought-induced mortality, which will help for the anti-drought management of bamboo.

Partitioning the biodiversity effects on productivity into density and size components.

Plant density and size - two factors that represent plant survival and growth - are key determinants of yield but have rarely been analysed explicitly in the context of biodiversity-productivity relationships. Here, we derive equations to partition the net, complementarity and selection effects of biodiversity into additive components that reflect diversity-induced changes in plant density and size. Applications of the new method to empirical datasets reveal contrasting ways in which plant density and size regulate yield in species mixtures. In an annual plant diversity experiment, overyielding is largely explained by selection effects associated with increased size of highly productive plant species. In a tree diversity experiment, the cause of overyielding shifts from enhanced growth in tree size to reduced mortality by complementary use of canopy space during stand development. These results highlight the capability of the new method to resolve crucial, yet understudied, demographic links between biodiversity and productivity.

Drought survival in conifer species is related to the time required to cross the stomatal safety margin.

The regulation of water loss and the spread of xylem embolism have mostly been considered separately. The development of an integrated approach taking into account the temporal dynamics and relative contributions of these mechanisms to plant drought responses is urgently needed. Do conifer species native to mesic and xeric environments display different hydraulic strategies and temporal sequences under drought? A dry-down experiment was performed on seedlings of four conifer species differing in embolism resistance, from drought-sensitive to extremely drought-resistant species. A set of traits related to drought survival was measured, including turgor loss point, stomatal closure, minimum leaf conductance, and xylem embolism resistance. All species reached full stomatal closure before the onset of embolism, with all but the most drought-sensitive species presenting large stomatal safety margins, demonstrating that highly drought-resistant species do not keep their stomata open under drought conditions. Plant dry-down time to death was significantly influenced by the xylem embolism threshold, stomatal safety margin, and minimum leaf conductance, and was best explained by the newly introduced stomatal margin retention index (SMRIΨ50) which reflects the time required to cross the stomatal safety margin. The SMRIΨ50 may become a key tool for the characterization of interspecific drought survival variability in trees.

Climate-driven tree growth and mortality in the Black Forest, Germany-Long-term observations.

Episodic tree mortality can be caused by various reasons. This study describes climate-driven tree mortality and tree growth in the Black Forest mountain range in Germany. It is based on a 68-year consistent data series describing the annual mortality of all trees growing in a forest area of almost 250 thousand ha. The study excludes mortality caused by storm, snow and ice, and fire. The sequence of the remaining mortality, the so-called "desiccated trees," is analyzed and compared with the sequence of the climatic water balance during the growing season and the annual radial growth of Norway spruce in the Black Forest. The annual radial growth series covers 121 years and the climatic water balance series 140 years. These unique time series enable a quantitative assessment of multidecadal drought and heat impacts on growth and mortality of forest trees on a regional spatial scale. Data compiled here suggest that the mortality of desiccated trees in the Black Forest during the last 68 years is driven by the climatic water balance. Decreasing climatic water balance coincided with an increase in tree mortality and growth decline. Consecutive hot and dry summers enhance mortality and growth decline as a consequence of drought legacies lasting several years. The sensitivity of tree growth and mortality to changes in the climatic water balance increases with the decreasing trend of the climatic water balance. The findings identify the climatic water balance as the main driver of mortality and growth variation during the 68-year observation period on a landscape-scale including a variety of different sites. They suggest that bark beetle population dynamics modify mortality rates. They as well provide evidence that the mortality during the last 140 years never was as high as in the most recent years.

The Database of European Forest Insect and Disease Disturbances: DEFID2.

Insect and disease outbreaks in forests are biotic disturbances that can profoundly alter ecosystem dynamics. In many parts of the world, these disturbance regimes are intensifying as the climate changes and shifts the distribution of species and biomes. As a result, key forest ecosystem services, such as carbon sequestration, regulation of water flows, wood production, protection of soils, and the conservation of biodiversity, could be increasingly compromised. Despite the relevance of these detrimental effects, there are currently no spatially detailed databases that record insect and disease disturbances on forests at the pan-European scale. Here, we present the new Database of European Forest Insect and Disease Disturbances (DEFID2). It comprises over 650,000 harmonized georeferenced records, mapped as polygons or points, of insects and disease disturbances that occurred between 1963 and 2021 in European forests. The records currently span eight different countries and were acquired through diverse methods (e.g., ground surveys, remote sensing techniques). The records in DEFID2 are described by a set of qualitative attributes, including severity and patterns of damage symptoms, agents, host tree species, climate-driven trigger factors, silvicultural practices, and eventual sanitary interventions. They are further complemented with a satellite-based quantitative characterization of the affected forest areas based on Landsat Normalized Burn Ratio time series, and damage metrics derived from them using the LandTrendr spectral-temporal segmentation algorithm (including onset, duration, magnitude, and rate of the disturbance), and possible interactions with windthrow and wildfire events. The DEFID2 database is a novel resource for many large-scale applications dealing with biotic disturbances. It offers a unique contribution to design networks of experiments, improve our understanding of ecological processes underlying biotic forest disturbances, monitor their dynamics, and enhance their representation in land-climate models. Further data sharing is encouraged to extend and improve the DEFID2 database continuously. The database is freely available at https://jeodpp.jrc.ec.europa.eu/ftp/jrc-opendata/FOREST/DISTURBANCES/DEFID2/.

Observed forest trait velocities have not kept pace with hydraulic stress from climate change.

The extent to which future climate change will increase forest stress and the amount to which species and forest ecosystems can acclimate or adapt to increased stress is a major unknown. We used high-resolution maps of hydraulic traits representing the diversity in tree drought tolerance across the United States, a hydraulically enabled tree model, and forest inventory observations of demographic shifts to quantify the ability for within-species acclimation and between-species range shifts to mediate climate stress. We found that forests are likely to experience increases in both acute and chronic hydraulic stress with climate change. Based on current species distributions, regional hydraulic trait diversity was sufficient to buffer against increased stress in 88% of forested areas. However, observed trait velocities in 81% of forested areas are not keeping up with the rate required to ameliorate projected future stress without leaf area acclimation.

Modeling the mechanisms of conifer mortality under seawater exposure.

Relative sea level rise (SLR) increasingly impacts coastal ecosystems through the formation of ghost forests. To predict the future of coastal ecosystems under SLR and changing climate, it is important to understand the physiological mechanisms underlying coastal tree mortality and to integrate this knowledge into dynamic vegetation models. We incorporate the physiological effect of salinity and hypoxia in a dynamic vegetation model in the Earth system land model, and used the model to investigate the mechanisms of mortality of conifer forests on the west and east coast sites of USA, where trees experience different form of sea water exposure. Simulations suggest similar physiological mechanisms can result in different mortality patterns. At the east coast site that experienced severe increases in seawater exposure, trees loose photosynthetic capacity and roots rapidly, and both storage carbon and hydraulic conductance decrease significantly within a year. Over time, further consumption of storage carbon that leads to carbon starvation dominates mortality. At the west coast site that gradually exposed to seawater through SLR, hydraulic failure dominates mortality because root loss impacts on conductance are greater than the degree of storage carbon depletion. Measurements and modeling focused on understanding the physiological mechanisms of mortality is critical to reducing predictive uncertainty.

The cumulative impacts of droughts and N deposition on Norway spruce (Picea abies) in Switzerland based on 37 years of forest monitoring.

Norway spruce is one of the most important tree species in Central Europe, however, it is facing major problems with recent droughts. In this study we present 37 years (1985-2022) of long-term forest observation data on 82 different forest sites across Switzerland including 134'348 tree observations. The sites consists of managed spruce or mixed forest stands with beech (Fagus sylvatica) and comprise large gradients in altitude (290-1870 m), precipitation (570-2448 mm a-1), temperature (3.6-10.9 °C) or total nitrogen deposition (8.5-81.2 kg N ha-1 a-1). Long-term tree mortality has increased more than fivefold due to the multiple drought years 2019, 2020 and 2022, which is more than double the increase following the 2003 drought. We used a Bayesian multilevel model including three years of lagged drought indicator to predict spruce mortality. Besides stand age, the most important factors were drought and N deposition. Especially under drought conditions spruce mortality was increased on sites with high N deposition. Moreover, N deposition increased the imbalance of foliar phosphorus concentrations, with negative impacts on tree mortality. Mortality was increased by a factor of 1.8 in spruce compared to mixed beech and spruce stands. Stands with high mortality rates showed previously an increased proportion of trees with damaged crowns, especially after the droughts of 2003 and 2018. Taken together, we found evidences of an increase in spruce mortality droughts amplified under high N depositions. The perennial drought of 2018-2020 resulted in a cumulative spruce mortality of 12.1 % (564 dead trees in 82 sites) in only three years. With a Bayesian change-point regression framework we estimated a critical empirical load for nitrogen of 10.9 ± 4.2 kg N ha-1 a-1, which is in line with current thresholds, above which future plantings of spruce in Switzerland may not be a sustainable option due to the observed interaction between drought and nitrogen deposition.