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AbstractGlobal climate change is expected to further raise the frequency and severity of extreme events, such as droughts. The effects of extreme droughts on trees are difficult to disentangle given the inherent complexity of drought events (frequency, severity, duration, and timing during the growing season). Besides, drought effects might be modulated by trees’ phenotypic variability, which is, in turn, affected by long‐term local selective pressures and management legacies. Here we investigated the magnitude and the temporal changes of tree‐level resilience (i.e., resistance, recovery, and resilience) to extreme droughts. Moreover, we assessed the tree‐, site‐, and drought‐related factors and their interactions driving the tree‐level resilience to extreme droughts. We used a tree‐ring network of the widely distributed Scots pine (Pinus sylvestris) along a 2,800 km latitudinal gradient from southern Spain to northern Germany. We found that the resilience to extreme drought decreased in mid‐elevation and low productivity sites from 1980–1999 to 2000–2011 likely due to more frequent and severe droughts in the later period. Our study showed that the impact of drought on tree‐level resilience was not dependent on its latitudinal location, but rather on the type of sites trees were growing at and on their growth performances (i.e., magnitude and variability of growth) during the predrought period. We found significant interactive effects between drought duration and tree growth prior to drought, suggesting that Scots pine trees with higher magnitude and variability of growth in the long term are more vulnerable to long and severe droughts. Moreover, our results indicate that Scots pine trees that experienced more frequent droughts over the long‐term were less resistant to extreme droughts. We, therefore, conclude that the physiological resilience to extreme droughts might be constrained by their growth prior to drought, and that more frequent and longer drought periods may overstrain their potential for acclimation.

AbstractClimate warming should result in hotter droughts of unprecedented severity in this century. Such droughts have been linked with massive tree mortality, and data suggest that warming interacts with drought to aggravate plant performance. Yet how forests will respond to hotter droughts remains unclear, as does the suite of mechanisms trees use to deal with hot droughts. We used an ecosystem‐scale manipulation of precipitation and temperature on piñon pine (Pinus edulis) and juniper (Juniperus monosperma) trees to investigate nitrogen (N) cycling‐induced mitigation processes related to hotter droughts. We found that while negative impacts on plant carbon and water balance are manifest after prolonged drought, performance reductions were not amplified by warmer temperatures. Rather, increased temperatures for 5 years stimulated soil N cycling under piñon trees and modified tree N allocation for both species, resulting in mitigation of hotter drought impacts on tree water and carbon functions. These findings suggest that adjustments in N cycling are likely after multi‐year warming conditions and that such changes may buffer reductions in tree performance during hotter droughts. The results highlight our incomplete understanding of trees' ability to acclimate to climate change, raising fundamental questions about the resistance potential of forests to long‐term, compound climatic stresses.

Summary Over the last decades, spring leaf‐out of temperate and boreal trees has substantially advanced in response to global warming, affecting terrestrial biogeochemical fluxes and the Earth's climate system. However, it remains unclear whether leaf‐out will continue to advance with further warming because species’ effective chilling temperatures, as well as the amount of chilling time required to break dormancy, are still largely unknown for most forest tree species. Here, we assessed the progress of winter dormancy and quantified the efficiency of different chilling temperatures in six dominant temperate European tree species by exposing 1170 twig cuttings to a range of temperatures from −2°C to 10°C for 1, 3, 6 or 12 wk. We found that freezing temperatures were most effective for half of the species or as effective as chilling temperatures up to 10°C, that is, leading to minimum thermal time to and maximum success of budburst. Interestingly, chilling duration had a much larger effect on dormancy release than absolute chilling temperature. Our experimental results challenge the common assumption that optimal chilling temperatures range c. 4–6°C, instead revealing strong sensitivity to a large range of temperatures. These findings are valuable for improving phenological models and predicting future spring phenology in a warming world.

Abstract Key message Understory plant communities are essential for the recruitment of trees making up future forests. Independent of plant diversity, the understory across different forest ecosystems shows considerable physiological acclimation and structural stability towards drought events, which are expected to occur more frequently in future. Context Understory plant communities are essential for the recruitment of trees making up the future forest. It is so far poorly understood how climate change will affect understory in beech and conifer forests managed at different intensity levels. Aims We hypothesized that drought would affect transpiration and carbon isotope discrimination but not species richness and diversity. Moreover, we assumed that forest management intensity will modify the responses to drought of the understory community. Methods We set up roofs in forests with a gradient of management intensities (unmanaged beech—managed beech—intensively managed conifer forests) in three regions across Germany. A drought event close to the 2003 drought was imposed in two consecutive years. Results After 2 years, the realized precipitation reduction was between 27% and 34%. The averaged water content in the top 20 cm of the soil under the roof was reduced by 2% to 8% compared with the control. In the 1st year, leaf level transpiration was reduced for different functional groups, which scaled to community transpiration modified by additional effects of drought on functional group leaf area. Acclimation effects in most functional groups were observed in the 2nd year. Conclusion Forest understory shows high plasticity at the leaf and community level, and high structural stability to changing climate conditions with drought events.

AbstractUnprecedented tree dieback across Central Europe caused by recent global change‐type drought events highlights the need for a better mechanistic understanding of drought‐induced tree mortality. Although numerous physiological risk factors have been identified, the importance of two principal mechanisms, hydraulic failure and carbon starvation, is still debated. It further remains largely unresolved how the local neighborhood composition affects individual mortality risk. We studied 9435 young trees of 12 temperate species planted in a diversity experiment in 2013 to assess how hydraulic traits, carbon dynamics, pest infestation, tree height and neighborhood competition influence individual mortality risk. Following the most extreme global change‐type drought since record in 2018, one third of these trees died. Across species, hydraulic safety margins (HSMs) were negatively and a shift towards a higher sugar fraction in the non‐structural carbohydrate (NSC) pool positively associated with mortality risk. Moreover, trees infested by bark beetles had a higher mortality risk, and taller trees a lower mortality risk. Most neighborhood interactions were beneficial, although neighborhood effects were highly species‐specific. Species that suffered more from drought, especially Larix spp. and Betula spp., tended to increase the survival probability of their neighbors and vice versa. While severe tissue dehydration marks the final stage of drought‐induced tree mortality, we show that hydraulic failure is interrelated with a series of other, mutually inclusive processes. These include shifts in NSC pools driven by osmotic adjustment and/or starch depletion as well as pest infestation and are modulated by the size and species identity of a tree and its neighbors. A more holistic view that accounts for multiple causes of drought‐induced tree mortality is required to improve predictions of trends in global forest dynamics and to identify mutually beneficial species combinations.

AbstractLong generation times have been suggested to hamper rapid genetic adaptation of organisms to changing environmental conditions. We examined if environmental memory of the parental Scots pines (Pinus sylvestris L.) drive offspring survival and growth. We used seeds from trees growing under naturally dry conditions (control), irrigated trees (irrigated from 2003 to 2016), and formerly irrigated trees (“irrigation stop”; irrigated from 2003–2013; control condition since 2014). We performed two experiments, one under controlled greenhouse conditions and one at the experimental field site. In the greenhouse, the offspring from control trees exposed regularly to drought were more tolerant to hot–drought conditions than the offspring from irrigated trees and showed lower mortality even though there was no genetic difference. However, under optimal conditions (high water supply and full sunlight), these offspring showed lower growth and were outperformed by the offspring of the irrigated trees. This different offspring growth, with the offspring of the “irrigation‐stop” trees showing intermediate responses, points to the important role of transgenerational memory for the long‐term acclimation of trees. Such memory effects, however, may be overridden by climatic extremes during germination and early growth stages such as the European 2018 mega‐drought that impacted our field experiment.