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.

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.

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.

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.

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.

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.

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.

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.

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.

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.