Stand age diversity (and more than climate change) affects forests' resilience and stability, although unevenly.

Stand age significantly influences the functioning of forest ecosystems by shaping structural and physiological plant traits, affecting water and carbon budgets. Forest age distribution is determined by the interplay of tree mortality and regeneration, influenced by both natural and anthropogenic disturbances. Unfortunately, human-driven alteration of tree age distribution presents an underexplored avenue for enhancing forest stability and resilience. In our study, we investigated how age impacts the stability and resilience of the forest carbon budget under both current and future climate conditions. We employed a state-of-the-science biogeochemical, biophysical, validated process-based model on historically managed forest stands, projecting their future as undisturbed systems, i.e., left at their natural evolution with no management interventions (i.e., forests are left to develop undisturbed). Such a model, forced by climate data from five Earth System Models under four representative climate scenarios and one baseline scenario to disentangle the effect of climate change, spanned several age classes as representative of the current European forests' context, for each stand. Our findings indicate that Net Primary Production (NPP) peaks in the young and middle-aged classes (16- to 50-year-old), aligning with longstanding ecological theories, regardless of the climate scenario. Under climate change, the beech forest exhibited an increase in NPP and maintained stability across all age classes, while resilience remained constant with rising atmospheric CO2 and temperatures. However, NPP declined under climate change scenarios for the Norway spruce and Scots pine sites. In these coniferous forests, stability and resilience were more influenced. These results underscore the necessity of accounting for age class diversity -lacking in most, if not all, the current Global Vegetation Models - for reliable and robust assessments of the impacts of climate change on future forests' stability and resilience capacity. We, therefore, advocate for customized management strategies that enhance the adaptability of forests to changing climatic conditions, taking into account the diverse responses of different species and age groups to climate.

Retrospective analysis of wood anatomical traits and tree-ring isotopes suggests site-specific mechanisms triggering Araucaria araucana drought-induced dieback.

In 2010-2018, Northern Patagonia featured the longest severe drought of the last millennium. This extreme dry spell triggered widespread growth decline and forest dieback. Nonetheless, the roles played by the two major mechanisms driving dieback, hydraulic failure and carbon starvation, are still not clear and understudied in this seasonally dry region. Here, for the 1800-2017 period, we apply a retrospective analysis of radial growth, wood anatomical traits (lumen area, cell-wall thickness) and δ13 C and δ18 O stable isotopes to assess dieback causes of the iconic conifer Araucaria araucana. We selected three stands where declining (defoliated) and nondeclining (not defoliated) trees coexisted along a precipitation gradient from the warm-dry Coastal Range to the cool-wet Andes. At all sites declining trees showed lower radial growth and lower theoretical hydraulic conductivity, suggesting a long-lasting process of hydraulic deterioration in their water transport system compared to nondeclining, coexisting trees. Wood anatomical traits evidenced that this divergence between declining and nondeclining trees started at least seven decades before canopy dieback. In the drier stands, declining trees showed higher water-use efficiency (WUE) throughout the whole period, which we attributed to early stomatal closure, suggesting a greater carbon starvation risk consistent with thinner cell walls. In the wettest stand, we found the opposite pattern. Here, a reduction in WUE coupled with thicker cell walls suggested increased carbon assimilation rates and exposure to drought-induced hydraulic failure. The δ18 O values indicated different strategies of gas exchange between sites, which are likely a consequence of microsite conditions and water sources. Multiproxy, retrospective quantifications of xylem anatomical traits and tree-ring isotopes provide a robust tool to identify and forecast, which stands or trees will show dieback or, on the contrary, which will likely withstand and be more resilient to future hotter droughts.

Wood anatomical traits in black spruce reveal latent water constraints on the boreal forest.

The effects of climate change on high-latitude forest ecosystems are complex, making forecasts of future scenarios uncertain. The predicted lengthening of the growing season under warming conditions is expected to increase tree growth rates. However, there is evidence of an increasing sensitivity of the boreal forest to drought stress. To assess the influence of temperature and precipitation on the growth of black spruce (Picea mariana), we investigated long-term series of wood anatomical traits on 20 trees from four sites along 600 km, the latitudinal range of the closed boreal forest in Quebec, Canada. We correlated the anatomical traits resolved at intraring level with daily temperature, vapor pressure deficit (VPD), and precipitation during the 1943-2010 period. Tree-ring width, number of cells per ring and cell wall thickness were positively affected by spring and summer daily mean and maximum temperature at the northern sites. These results agree with the well-known positive effect of high temperatures on tree ring formation at high latitudes. However, we captured, for the first time in this region, the latent impact of water availability on xylem traits. Indeed, in all the four sites, cell lumen area showed positive correlations with daily precipitation (mostly at low latitude), and/or negative correlations with daily mean and maximum temperature and VPD (mostly at high latitude). We inferred that drought, due to high temperatures, low precipitations, or both, negatively affects cell enlargement across the closed boreal forest, including the northernmost sites. The production of tracheids with narrower lumen, potentially more resistant to cavitation, could increase xylem hydraulic safety under a warmer and drier climate. However, this would result in lower xylem conductivity, with consequent long-term hydraulic deterioration, growth decline, and possibly lead to tree dieback, as observed in other forest ecosystems at lower latitudes.

Contrasting patterns of water use efficiency and annual radial growth among European beech forests along the Italian peninsula.

Tree mortality and forest dieback episodes are increasing due to drought and heat stress. Nevertheless, a comprehensive understanding of mechanisms enabling trees to withstand and survive droughts remains lacking. Our study investigated basal area increment (BAI), and δ13C-derived intrinsic water-use-efficiency (iWUE), to elucidate beech resilience across four healthy stands in Italy with varying climates and soil water availability. Additionally, fist-order autocorrelation (AR1) analysis was performed to detect early warning signals for potential tree dieback risks during extreme drought events. Results reveal a negative link between BAI and vapour pressure deficit (VPD), especially in southern latitudes. After the 2003 drought, BAI decreased at the northern site, with an increase in δ13C and iWUE, indicating conservative water-use. Conversely, the southern sites showed increased BAI and iWUE, likely influenced by rising CO2 and improved water availability. In contrast, the central site sustained higher transpiration rates due to higher soil water holding capacity (SWHC). Despite varied responses, most sites exhibited reduced resilience to future extreme events, indicated by increased AR1. Temperature significantly affected beech iWUE and BAI in northern Italy, while VPD strongly influenced the southern latitudes. The observed increase in BAI and iWUE in southern regions might be attributed to an acclimation response.