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Nanopore raw read accuracy has improved to over 99%, making it a potential tool for metabarcoding. For broad adoption, guidelines on quality filtering are needed to ensure reliable taxonomic unit recovery. This study aims to provide those guidelines for a fungal metabarcoding context and to apply them to a case study of ectomycorrhizae in the decaying bark of Fagus sylvatica . We introduce the eNano pipeline to test two standard metabarcoding approaches: (1) Reference-based mapping leveraging UNITE's species hypothesis system (SH approach); (2) Constructing 98% OTUs (OTU approach). Our results demonstrate that both approaches are effective with Nanopore data. When using a reference database, we recommend strict mapping criteria rather than Phred-based filtering. Leveraging the SH-system further enhances reproducibility and facilitates cross-study communication. For the 98% OTUs, filtering reads at ≥Q25 is recommended. Our case study reveals that the decay gradient is a primary determinant of community composition and that specific mycorrhizal fungi colonize decaying bark. Complementing our metabarcoding results with root tip morphotypification, we identify Laccaria amethystina and Tomentella sublilacina as key ectomycorrhizae of saplings on decaying logs. These findings demonstrate that Nanopore sequencing can provide valuable ecological insights and support its broader use in fungal metabarcoding as read quality continues to improve.

With ongoing global warming, increasing water deficits promote physiological stress on forest ecosystems with negative impacts on tree growth, vitality, and survival. How individual tree species will react to increased drought stress is therefore a key research question to address for carbon accounting and the development of climate change mitigation strategies. Recent tree-ring studies have shown that trees at higher latitudes will benefit from warmer temperatures, yet this is likely highly species-dependent and less well-known for more temperate tree species. Using a unique pan-European tree-ring network of 26,430 European beech (Fagus sylvatica L.) trees from 2118 sites, we applied a linear mixed-effects modeling framework to (i) explain variation in climate-dependent growth and (ii) project growth for the near future (2021-2050) across the entire distribution of beech. We modeled the spatial pattern of radial growth responses to annually varying climate as a function of mean climate conditions (mean annual temperature, mean annual climatic water balance, and continentality). Over the calibration period (1952-2011), the model yielded high regional explanatory power (R 2  = 0.38-0.72). Considering a moderate climate change scenario (CMIP6 SSP2-4.5), beech growth is projected to decrease in the future across most of its distribution range. In particular, projected growth decreases by 12%-18% (interquartile range) in northwestern Central Europe and by 11%-21% in the Mediterranean region. In contrast, climate-driven growth increases are limited to around 13% of the current occurrence, where the historical mean annual temperature was below ~6°C. More specifically, the model predicts a 3%-24% growth increase in the high-elevation clusters of the Alps and Carpathian Arc. Notably, we find little potential for future growth increases (-10 to +2%) at the poleward leading edge in southern Scandinavia. Because in this region beech growth is found to be primarily water-limited, a northward shift in its distributional range will be constrained by water availability., (© 2024 The Author(s). Global Change Biology published by John Wiley & Sons Ltd.)

Riparian corridors often act as low-land climate refugia for temperate tree species in their southern distribution range. A plausible mechanism is the buffering of regional climate extremes by local physiographic and biotic factors. We tested this idea using a 3-year-long microclimate dataset collected along the Ciron river, a refugia for European beech (Fagus sylvatica) in southwestern France. Across the whole network, canopy gap fraction was the main predictor for spatial microclimatic variations, together with two other landscape features (elevation above the river and woodland fraction within a 300m radius). However, within the riparian forest only (canopy gap fraction < 25%, distance to the river < 150m), variations of up to -4°C and + 15% in summertime daily maximum air temperature and minimum relative humidity, respectively, were still found from the plateau to the cooler, moister river banks, only ~ 5-10m below. Elevation above the river was then identified as the main predictor, and explained the marked variations from the plateau to the banks much better than canopy gap fraction. The microclimate measured near the river is as cool but moister than the macroclimate encountered at 700-1000m asl further east in F. sylvatica's main distribution range. Indeed, at all locations, we found that air relative humidity was higher than expected from a temperature-only effect, suggesting that extra moisture is brought by the river. Our results explain well why beech trees in this climate refugium are restricted to the river gorges where microtopographic variations are the strongest and canopy gaps are rare., (© 2024. The Author(s) under exclusive licence to International Society of Biometeorology.)

Ectomycorrhizal (ECM) fungi distribute tree-derived carbon (C) via belowground hyphal networks in forest ecosystems. Here, we asked the following: (1) Is C transferred belowground to a neighboring tree retained in fungal structures or transported within the recipient tree? (2) Is the overlap of ectomycorrhizal fungi in mycorrhizal networks related to the amount of belowground C transfer? We used potted sapling pairs of European beech (Fagus sylvatica) and North-American Douglas-fir (Pseudotsuga menziesii) for 13 CO 2 pulse-labeling. We compared 13 C transfer from beech (donor) to either beech or Douglas-fir (recipient) and identified the ECM species. We measured the 13 C enrichment in soil, plant tissues, and ECM fractions of fungal-containing parts and plant transport tissues. In recipients, only fungal-containing tissue of ectomycorrhizas was significantly enriched in 13 C and not the plant tissue. Douglas-fir recipients shared on average one ECM species with donors and had a lower 13 C enrichment than beech recipients, which shared on average three species with donors. Our results support that recently assimilated C transferred belowground is shared among fungi colonizing tree roots but not among trees. In mixed forests with beech and Douglas-fir, the links for C movement might be hampered due to low mycorrhizal overlap with consequences for soil C cycling., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

Marginal populations usually have low densities and are considered to be particularly vulnerable to environmental stochasticity. Using data collected in nest boxes, we analyzed the breeding performance of the edible dormouse ( Glis glis ), an obligate hibernating rodent and a seed predator in deciduous forests, in two populations at the distribution range's edge. Despite being only 20 km apart from each other, Montseny is a large patch of mixed deciduous forests (oaks and beech), whereas Montnegre would be the harshest habitat, that is, a small, isolated patch with only oaks. First, we studied the differences in climate and tree cover change in the two populations. Second, we analyzed the direct and indirect roles of local climate conditions and seed availability on breeding performance over 10 years in each population. Finally, we explored the influence of tree cover change on the occupancy dynamics in the two populations. Our results showed contrasting responses between populations: in Montseny, asynchronous seed production between oaks and beech precluded skip breeding, and breeding performance increased with seed availability. Furthermore, dormice in Montseny may use pollen production to anticipate the amount of beech nut resources and adjust their breeding effort. Boxes showed higher occupancy and colonization and fewer extinctions in Montseny than in Montnegre, where seed availability did not drive breeding performance. Results from Montnegre suggest that skip breeding was an adaptive response to a more pulsed, harsher environment. Here, females produced a similar number of pups than at Montseny. Long-term studies dealing with population responses in marginal habitats can lead to a deeper understanding of the capacities of organisms to adapt to harsh environments. Although local adaptation is frequently documented across various taxa, studies at the distribution edge may shed light on our still limited comprehension of the underlying mechanisms responsible for its occurrence., Competing Interests: None of the authors have a conflict of interest., (© 2024 The Author(s). Ecology and Evolution published by John Wiley & Sons Ltd.)

Increasingly frequent and severe droughts pose significant threats to forest ecosystems, particularly affecting photosynthesis, a crucial physiological process for plant growth and biomass production. This study investigates the impact of phosphorus fertilization on the photosynthesis of common beech ( Fagus sylvatica L.) and sessile oak ( Quercus petraea (Matt.) Liebl.). In a common garden experiment, saplings originating from two provenances (wetter KA and drier SB provenances) were exposed to regular watering and drought in interaction with moderate and high phosphorus concentrations in the growing substrate. Results indicated that drought significantly reduced pre-dawn leaf water potential (Ψ PD ), net photosynthesis (A net ), stomatal conductance (g s ) and photosynthetic performance index (PI abs ) in both species. Phosphorus fertilization had a negative impact on A net and PI abs , thus exacerbating the negative impact of drought on photosynthetic efficiency, potentially due to excessive phosphorus absorption by saplings. Provenance differences were notable, with the KA provenance showing better drought resilience. This research highlights the complexity of nutrient-drought interactions and underscores the need for cautious application of fertilization strategies in reforestation efforts under changing climatic conditions.

The future performance of the widely abundant European beech (Fagus sylvatica L.) across its ecological amplitude is uncertain. Although beech is considered drought-sensitive and thus negatively affected by drought events, scientific evidence indicating increasing drought vulnerability under climate change on a cross-regional scale remains elusive. While evaluating changes in climate sensitivity of secondary growth offers a promising avenue, studies from productive, closed-canopy forests suffer from knowledge gaps, especially regarding the natural variability of climate sensitivity and how it relates to radial growth as an indicator of tree vitality. Since beech is sensitive to drought, we in this study use a drought index as a climate variable to account for the combined effects of temperature and water availability and explore how the drought sensitivity of secondary growth varies temporally in dependence on growth variability, growth trends, and climatic water availability across the species' ecological amplitude. Our results show that drought sensitivity is highly variable and non-stationary, though consistently higher at dry sites compared to moist sites. Increasing drought sensitivity can largely be explained by increasing climatic aridity, especially as it is exacerbated by climate change and trees' rank progression within forest communities, as (co-)dominant trees are more sensitive to extra-canopy climatic conditions than trees embedded in understories. However, during the driest periods of the 20th century, growth showed clear signs of being decoupled from climate. This may indicate fundamental changes in system behavior and be early-warning signals of decreasing drought tolerance. The multiple significant interaction terms in our model elucidate the complexity of European beech's drought sensitivity, which needs to be taken into consideration when assessing this species' response to climate change., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Author(s). Published by Elsevier B.V. All rights reserved.)

Mesophyll resistance for CO 2 diffusion (r m ) is one of the main limitations for photosynthesis and plant growth. Breeding new varieties with lower r m requires knowledge of its distinct components. We tested new method for estimating the relative drawdowns of CO 2 concentration (c) across hypostomatous leaves of Fagus sylvatica. This technique yields values of the ratio of the internal CO 2 concentrations at the adaxial and abaxial leaf side, c d /c b , the drawdown in the intercellular air space (IAS), and intracellular drawdown between IAS and chloroplast stroma, c c /c bd . The method is based on carbon isotope composition of leaf dry matter and epicuticular wax isolated from upper and lower leaf sides. We investigated leaves from tree-canopy profile to analyse the effects of light and leaf anatomy on the drawdowns and partitioning of r m into its inter- (r IAS ) and intracellular (r liq ) components. Validity of the new method was tested by independent measurements of r m using conventional isotopic and gas exchange techniques. 73% of investigated leaves had adaxial epicuticular wax enriched in 13 C compared to abaxial wax (by 0.50‰ on average), yielding 0.98 and 0.70 for average of c d /c b and c c /c bd , respectively. The r IAS to r liq proportion were 5.5:94.5% in sun-exposed and 14.8:85.2% in shaded leaves. c c dropped to less than half of the atmospheric value in the sunlit and to about two-thirds of it in shaded leaves. This method shows that r IAS is minor but not negligible part of r m and reflects leaf anatomy traits, i.e. leaf mass per area and thickness., (© 2024 The Authors. Plant Biology published by John Wiley & Sons Ltd on behalf of German Society for Plant Sciences, Royal Botanical Society of the Netherlands.)

The persistence of plant populations depends crucially on successful regeneration. Yet, little is known about the effects of consecutive winter and spring frost events on the regeneration stage of trees from different seed sources, although this will partly determine the success of climate warming-driven poleward range shifts. In a common garden experiment with European beech ( Fagus sylvatica ) seedlings from winter 2015/2016 to autumn 2017, we studied how simulated successive spring and winter frost events affect leaf-out dates, growth performance, and survival rates of 1- to 2-year-old seedlings from provenances differing in climate at origin. We further investigated the combined effects of successive frost events. The first spring frost after germination led to a mortality rate up to 75%, resulting in reduced seedling numbers but better frost tolerance of the survivors, as reflected in a weaker impact of the following winter frost event in the survivors compared to the non-acclimated control. Final plant height was most strongly reduced by the spring frost in the second year. The winter frost event delayed leaf-out by up to 40 days, leading to severe growth impairment in 2017. Our results indicate partly successful frost acclimation and/or the selection of frost-hardier individuals, because the negative growth effects of consecutive frost events did not add up after exposure to more than one event. Both mechanisms may help to increase the frost tolerance of beech offspring. Nevertheless, mortality after the first spring frost was high, and frost exposure generally caused growth reductions. Thus, achieving higher frost tolerance may not be sufficient for beech seedlings to overcome frost-induced reductions in competitive strength caused by winter frost damage and delayed leaf enfolding., Competing Interests: All authors have no conflicts of interest to declare., (© 2024 The Author(s). Ecology and Evolution published by John Wiley & Sons Ltd.)

Similar to soils, tree stems emit and consume nitrous oxide (N 2 O) from the atmosphere. Although tree leaves dominate tree surface area, they have been completely excluded from field N 2 O flux measurements and therefore their role in forest N 2 O exchange remains unknown. We explored the contribution of leaf fluxes to forest N 2 O exchange. We determined the N 2 O exchange of mature European beech (Fagus sylvatica) stems and shoots (i.e., terminal branches) and of adjacent forest floor, in a typical temperate upland forest in Germany. The beech stems, and particularly the shoots, acted as net N 2 O sinks (-0.254 ± 0.827 μg N 2 O m -2 stem area h -1 and -4.54 ± 1.53 μg N 2 O m -2 leaf area h -1 , respectively), while the forest floor was a net source (2.41 ± 1.08 μg N 2 O m -2 soil area h -1 ). The unstudied tree shoots were identified as a significant contributor to the net ecosystem N 2 O exchange. Moreover, we revealed for the first time that tree leaves act as substantial N 2 O sinks. Although this is the first study of its kind, it is of global importance for the proper design of future flux studies in forest ecosystems worldwide. Our results demonstrate that excluding tree leaves from forest N 2 O flux measurements can lead to misinterpretation of tree and forest N 2 O exchange, and thus global forest greenhouse gas flux inventories., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

The hyperdiverse wood-inhabiting fungi play a crucial role in the global carbon cycle, but often are threatened by deadwood removal, particularly in temperate forests dominated by European beech ( Fagus sylvatica ) and Oriental beech ( Fagus orientalis ). To study the impact of abiotic drivers, deadwood factors, forest management and biogeographical patterns in forests of both beech species on fungal composition and diversity, we collected 215 deadwood-drilling samples in 18 forests from France to Armenia and identified fungi by meta-barcoding. In our analyses, we distinguished the patterns driven by rare, common, and dominant species using Hill numbers. Despite a broad overlap in species, the fungal composition with focus on rare species was determined by Fagus species, deadwood type, deadwood diameter, precipitation, temperature, and management status in decreasing order. Shifting the focus on common and dominant species, only Fagus species, both climate variables and deadwood type remained. The richness of species within the deadwood objects increased significantly only with decay stage. Gamma diversity in European beech forests was higher than in Oriental beech forests. We revealed the highest gamma diversity for old-growth forests of European beech when focusing on dominant species. Our results implicate that deadwood retention efforts, focusing on dominant fungi species, critical for the decay process, should be distributed across precipitation and temperature gradients and both Fagus species. Strategies focusing on rare species should additionally focus on different diameters and on the conservation of old-growth forests., Competing Interests: The authors have no conflicts of interest., (© 2024 The Author(s). Ecology and Evolution published by John Wiley & Sons Ltd.)

It is well established that solar irradiance greatly influences tree metabolism and growth through photosynthesis, but its effects acting through individual climate metrics have not yet been well quantified. Understanding these effects is crucial for assessing the impacts of climate change on forest ecosystems. To describe the effects of solar irradiance on tree growth, we installed 110 automatic dendrometers in two old-growth mountain forest reserves in Central Europe, performed detailed terrestrial and aerial laser scanning to obtain precise tree profiles, and used these to simulate the sum of solar irradiance received by each tree on a daily basis. Generalized linear mixed-effect models were applied to simulate the probability of growth and the growth intensity over seven growing seasons. Our results demonstrated various contrasting effects of solar irradiance on the growth of canopy trees. On the one hand, the highest daily growth rates corresponded with the highest solar irradiance potentials (i.e. the longest photoperiod). Intense solar irradiance significantly decreased tree growth, through an increase in the vapor pressure deficit. These effects were consistent for all species but had different magnitude. Tree growth is the most effective on long rainy/cloudy days with low solar irradiance., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

Forest trees adopt effective strategies to optimize nitrogen (N) use through internal N recycling. In the context of more recurrent environmental stresses due to climate change, the question remains of whether increased frequency of drought or defoliation threatens this internal N recycling strategy. We submitted 8-year-old beech trees to 2 years of either severe drought (Dro) or manual defoliation (Def) to create a state of N starvation. At the end of the second year before leaf senescence, we labeled the foliage of the Dro and Def trees, as well as that of control (Co) trees, with 15N-urea. Leaf N resorption, winter tree N storage (total N, 15N, amino acids, soluble proteins) and N remobilization in spring were evaluated for the three treatments. Defoliation and drought did not significantly impact foliar N resorption or N concentrations in organs in winter. Total N amounts in Def tree remained close to those in Co tree, but winter N was stored more in the branches than in the trunk and roots. Total N amount in Dro trees was drastically reduced (-55%), especially at the trunk level, but soluble protein concentrations increased in the trunk and fine roots compared with Co trees. During spring, 15N was mobilized from the trunk, branches and twigs of both Co and Def trees to support leaf growth. It was only provided through twig 15N remobilization in the Dro trees, thus resulting in extremely reduced Dro leaf N amounts. Our results suggest that stress-induced changes occur in N metabolism but with varying severity depending on the constraints: within-tree 15N transport and storage strategy changed in response to defoliation, whereas a soil water deficit induced a drastic reduction of the N amounts in all the tree organs. Consequently, N dysfunction could be involved in drought-induced beech tree mortality under the future climate., (© The Author(s) 2024. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Extreme droughts can have long-lasting effects on forest community dynamics and species interactions. Yet, our understanding of how drought legacy modulates ecological relationships is just unfolding. We tested the hypothesis that leaf chemistry and herbivory show long-term responses to premature defoliation caused by an extreme drought event in European beech (Fagus sylvatica L.). For two consecutive years after the extreme European summer drought in 2018, we collected leaves from the upper and lower canopy of adjacently growing drought-stressed and unstressed trees. Leaf chemistry was analyzed and leaf damage by different herbivore-feeding guilds was quantified. We found that drought had lasting impacts on leaf nutrients and on specialized metabolomic profiles. However, drought did not affect the primary metabolome. Drought-related phytochemical changes affected damage of leaf-chewing herbivores whereas damage caused by other herbivore-feeding guilds was largely unaffected. Drought legacy effects on phytochemistry and herbivory were often weaker than between-year or between-canopy strata variability. Our findings suggest that a single extreme drought event bears the potential to long-lastingly affect tree-herbivore interactions. Drought legacy effects likely become more important in modulating tree-herbivore interactions since drought frequency and severity are projected to globally increase in the coming decades., (© 2024 The Authors New Phytologist © 2024 New Phytologist Foundation.)

Background: Carbon (C) sink and stock are among the most important ecosystem services provided by forests in climate change mitigation policies. In this context, old-growth forests constitute an essential reference point for the development of close-to-nature silviculture, including C management techniques. Despite their small extent in Europe, temperate old-growth forests are assumed to be among the most prominent in terms of biomass and C stored. However, monitoring and reporting of C stocks is still poorly understood. To better understand the C stock amount and distribution in temperate old-growth forests, we estimated the C stock of two old-growth stands in the Dinaric Alps applying different assessment methods, including direct and indirect approaches (e.g., field measurements and allometric equations vs. IPCC standard methods). This paper presents the quantification and the distribution of C across the five main forest C pools (i.e., aboveground, belowground, deadwood, litter and soil) in the study areas and the differences between the applied methods., Results: We report a very prominent C stock in both study areas (507 Mg C ha - 1 ), concentrated in a few large trees (36% of C in 5% of trees). Moreover, we found significant differences in C stock estimation between direct and indirect methods. Indeed, the latter tended to underestimate or overestimate depending on the pool considered., Conclusions: Comparison of our results with previous studies and data collected in European forests highlights the prominence of temperate forests, among which the Dinaric Alps old-growth forests are the largest. These findings provide an important benchmark for the development of future approaches to the management of the European temperate forests. However, further and deeper research on C stock and fluxes in old-growth stands is of prime importance to understand the potential and limits of the climate mitigation role of forests., (© 2024. The Author(s).)

Background: European beech (Fagus sylvatica L.) trees produce seeds irregularly; therefore, it is necessary to store beech seeds for forestation. Despite the acquisition of desiccation tolerance during development, beech seeds are classified as intermediate because they lose viability during long-term storage faster than typical orthodox seeds. In this study, beech seeds stored for short (3 years) or long (20 years) periods under optimal conditions and displaying 92 and 30% germination capacity, respectively, were compared., Results: Aged seeds displayed increased membrane damage, manifested as electrolyte leakage and lipid peroxidation levels. Analyses have been based on embryonic axes, which contained higher levels of reactive oxygen species (ROS) and higher levels of protein-bound methionine sulfoxide (MetO) in aged seeds. Using label-free quantitative proteomics, 3,949 proteins were identified, of which 2,442 were reliably quantified pointing to 24 more abundant proteins and 35 less abundant proteins in beech seeds under long-term storage conditions. Functional analyses based on gene ontology annotations revealed that nucleic acid binding activity (molecular function), ribosome organization or biogenesis and transmembrane transport (cellular processes), translational proteins (protein class) and membranous anatomical entities (cellular compartment) were affected in aged seeds. To verify whether MetO, the oxidative posttranslational modification of proteins that can be reversed via the action of methionine sulfoxide reductase (Msr) enzymes, is involved in the aging of beech seeds, we identified and quantified 226 MetO-containing proteins, among which 9 and 19 exhibited significantly up- and downregulated MetO levels, respectively, in beech seeds under long-term storage conditions. Several Msr isoforms were identified and recognized as MsrA1-like, MsrA4, MsrB5 and MsrB5-like in beech seeds. Only MsrA1-like displayed decreased abundance in aged seeds., Conclusions: We demonstrated that the loss of membrane integrity reflected in the elevated abundance of membrane proteins had a higher impact on seed aging progress than the MetO/Msr system. Proteome analyses enabled us to propose protein Sec61 and glyceraldehyde-3-phosphate dehydrogenase as potential longevity modulators in beech seeds., (© 2024. The Author(s).)

Climate change effects on tree reproduction are poorly understood, even though the resilience of populations relies on sufficient regeneration to balance increasing rates of mortality. Forest-forming tree species often mast, i.e. reproduce through synchronised year-to-year variation in seed production, which improves pollination and reduces seed predation. Recent observations in European beech show, however, that current climate change can dampen interannual variation and synchrony of seed production and that this masting breakdown drastically reduces the viability of seed crops. Importantly, it is unclear under which conditions masting breakdown occurs and how widespread breakdown is in this pan-European species. Here, we analysed 50 long-term datasets of population-level seed production, sampled across the distribution of European beech, and identified increasing summer temperatures as the general driver of masting breakdown. Specifically, increases in site-specific mean maximum temperatures during June and July were observed across most of the species range, while the interannual variability of population-level seed production (CVp) decreased. The declines in CVp were greatest, where temperatures increased most rapidly. Additionally, the occurrence of crop failures and low seed years has decreased during the last four decades, signalling altered starvation effects of masting on seed predators. Notably, CVp did not vary among sites according to site mean summer temperature. Instead, masting breakdown occurs in response to warming local temperatures (i.e. increasing relative temperatures), such that the risk is not restricted to populations growing in warm average conditions. As lowered CVp can reduce viable seed production despite the overall increase in seed count, our results warn that a covert mechanism is underway that may hinder the regeneration potential of European beech under climate change, with great potential to alter forest functioning and community dynamics., (© 2024 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Fine roots are multifunctional organs that may change function with ageing or root branching events from primarily absorptive to resource transport and storage functions. It is not well understood, how fine root branching patterns and related root functional differentiation along the longitudinal root axis change with soil chemical and physical conditions. We examined the variation in fine root branching patterns (the relative frequency of 1st to 4th root orders) and root morphological and chemical traits of European beech trees with soil depth (topsoil vs. subsoil) and soil chemistry (five sites with acid to neutral/alkaline bedrock). Bedrock type and related soil chemistry had an only minor influence on branching patterns: base-poor, infertile sites showed no higher fine root branching than base-rich sites. The contribution of 1st-order root segments to total fine root length decreased at all sites from about 60% in the topsoil (including organic layer) to 45% in the lower subsoil. This change was associated with a decrease in specific root area and root N content and an increase in mean root diameter with soil depth, while root tissue density did not change consistently. We conclude that soil depth (which acts through soil physical and chemical drivers) influences the fine root branching patterns of beech much more than soil chemical variation across soil types. To examine whether changes in root function are indeed triggered by branching events or result from root ageing and diameter growth, spatially explicit root physiological and anatomical studies across root orders are needed., (© 2024. The Author(s).)

There are several highly damaging Phytophthora species pathogenic to forest trees, many of which have been spread beyond their native range by the international trade of live plants and infested materials. Such introductions can be reduced through the development of better tools capable of the early, rapid, and high-throughput detection of contaminated plants. This study utilized a volatilomics approach (solid-phase microextraction coupled to gas chromatography-mass spectrometry) to differentiate between several Phytophthora species in culture and discriminate between healthy and Phytophthora -inoculated European beech and pedunculate oak trees. We tentatively identified 14 compounds that could differentiate eight Phytophthora species from each other in vitro. All of the Phytophthora species examined, except Phytophthora cambivora , uniquely produced at least one compound not observed in the other species; however, most detected compounds were shared between multiple species. Phytophthora polonica had the most unique compounds and was the least similar of all the species examined. The inoculated seedlings had qualitatively different volatile profiles and could be distinguished from the healthy controls by the presence of isokaurene, anisole, and a mix of three unknown compounds. This study supports the notion that volatiles are suitable for screening plant material, detecting tree pathogens, and differentiating between healthy and diseased material.

The record-breaking drought in 2018 caused premature leaf discoloration and shedding (early browning) in many beech (Fagus sylvatica L.) dominated forests in Central Europe. However, a high degree of variability in drought response among individual beech trees was observed. While some trees were severely impacted by the prolonged water deficits and high temperatures, others remained vital with no or only minor signs of crown vitality loss. Why some beech trees were more susceptible to drought-induced crown damage than others and whether growth recovery is possible are poorly understood. Here, we aimed to identify growth characteristics associated with the variability in drought response between individual beech trees based on a sample of 470 trees in northern Switzerland. By combining tree growth measurements and crown condition assessments, we also investigated the possible link between crown dieback and growth recovery after drought. Beech trees with early browning exhibited an overall lower growth vigor before the 2018 drought than co-occurring vital beech trees. This lower vigor is mainly indicated by lower overall growth rates, stronger growth declines in the past decades, and higher growth-climate sensitivity. Particularly, warm previous year summer conditions negatively affected current growth of the early-browning trees. These findings suggest that the affected trees had less access to critical resources and were physiologically limited in their growth predisposing them to early browning. Following the 2018 drought, observed growth recovery potential corresponded to the amount of crown dieback and the local climatic water balance. Overall, our findings emphasize that beech-dominated forests in Central Europe are under increasing pressure from severe droughts, ultimately reducing the competitive ability of this species, especially on lowland sites with shallow soils and low water holding capacity., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier B.V. All rights reserved.)

Temperate forests are undergoing significant transformations due to the influence of climate change, including varying responses of different tree species to increasing temperature and drought severity. To comprehensively understand the full range of growth responses, representative datasets spanning extensive site and climatic gradients are essential. This study utilizes tree-ring data from 550 sites from the temperate forests of Czechia to assess growth trends of six dominant Central European tree species (European beech, Norway spruce, Scots pine, silver fir, sessile and pedunculate oak) over 1990-2014. By modeling mean growth series for each species and site, and employing principal component analysis, we identified the predominant growth trends. Over the study period, linear growth trends were evident across most sites (56% increasing, 32% decreasing, and 10% neutral). The proportion of sites with stationary positive trends increased from low toward high elevations, whereas the opposite was true for the stationary negative trends. Notably, within the middle range of their distribution (between 500 and 700 m a.s.l.), Norway spruce and European beech exhibited a mix of positive and negative growth trends. While Scots pine growth trends showed no clear elevation-based pattern, silver fir and oaks displayed consistent positive growth trends regardless of site elevation, indicating resilience to the ongoing warming. We demonstrate divergent growth trajectories across space and among species. These findings are particularly important as recent warming has triggered a gradual shift in the elevation range of optimal growth conditions for most tree species and has also led to a decoupling of growth trends between lowlands and mountain areas. As a result, further future shifts in the elevation range and changes in species diversity of European temperate forests can be expected., (© 2024 John Wiley & Sons Ltd.)

Common beech (Fagus sylvatica) is one of the most important deciduous tree species in European forests. However, climate-change-induced drought may threaten its dominant position. The Sonian Forest close to Brussels (Belgium) is home to some of the largest beech trees in the world. This UNESCO world heritage site is famous for its high density of very large beech trees as a result of its climatic suitability, fertile soil conditions, and past management. Here we utilized tree-ring data from increment cores to investigate the growth of these old and monumental beech trees, evaluating their growth trends, response to past climate, and the effect of mast years on 39 living and 16 recently wind-thrown trees. Our analysis reveals that the sampled trees were generally sensitive to spring and summer droughts but recovered quickly after such an extreme climatic event. The growth trend of living trees has remained high and only shows a slight, statistically insignificant, decline over the past 50 years. Although the overall growth rate remains strong (BAI 50 cm 2 /year), the past five decades have shown strong inter-annual growth variations due to frequent and more intense droughts combined with an increased frequency of mast years. We also found notable differences in growth patterns between the living trees and those that had recently been wind-thrown. While there were no significant differences between living and wind-thrown trees in response to droughts, heatwaves, or mast years when examining year-to-year growth changes, the wind-thrown trees did exhibit considerably lower overall growth rates and a significant downward trend in growth (BAI -0.57 cm 2 /year). This difference in growth trends has been apparent since at least the 1980s. Overall, the findings of this study can provide valuable insights for understanding the long-term dynamics of lowland beech forests and their responses to climate change., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Climate warming advances the onset of tree growth in spring, but above- and belowground phenology are not always synchronized. These differences in growth responses may result from differences in root and bud dormancy dynamics, but root dormancy is largely unexplored. We measured dormancy in roots and leaf buds of Fagus sylvatica and Populus nigra by quantifying the warming sum required to initiate above- and belowground growth in October, January and February. We furthermore carried out seven experiments, manipulating only the soil and not air temperature before or during tree leaf-out to evaluate the potential of warmer roots to influence budburst timing using seedlings and adult trees of F. sylvatica and seedlings of Betula pendula. Root dormancy was virtually absent in comparison with the much deeper winter bud dormancy. Roots were able to start growing immediately as soils were warmed during the winter. Interestingly, higher soil temperature advanced budburst across all experiments, with soil temperature possibly accounting for c. 44% of the effect of air temperature in advancing aboveground spring phenology per growing degree hour. Therefore, differences in root and bud dormancy dynamics, together with their interaction, likely explain the nonsynchronized above- and belowground plant growth responses to climate warming., (© 2023 The Authors New Phytologist © 2023 New Phytologist Foundation.)

Local survival of forest tree populations under climate change depends on existing genetic variation and their adaptability to changing environments. Responses to selection were studied in European beech (Fagus sylvatica) under field conditions. A total of 1087 adult trees, seeds, 1-year-old seedlings and established multiyear saplings were genotyped with 16 nuSSRs. Adult trees were assessed for phenotypic traits related to growth, phenology and reproduction. Parentage and paternity analyses were used to estimate effective female and male fecundity as a proxy of fitness and showed that few parents contributed to successful regeneration. Selection gradients were estimated from the relationship between traits and fecundity, while heritability and evolvability were estimated using mixed models and the breeder's equation. Larger trees bearing more fruit and early male flowering had higher total fecundity, while trees with longer growth season had lower total fecundity (directional selection). Stabilizing selection on spring phenology was found for female fecundity, highlighting the role of late frosts as a selection driver. Selection gradients for other traits varied between measurement years and the offspring cohort used to estimate parental fecundity. Compared to other studies in natural populations, we found low to moderate heritability and evolvability for most traits. Response to selection was higher for growth than for budburst, leaf senescence or reproduction traits, reflecting more consistent selection gradients across years and sex functions, and higher phenotypic variability in the population. Our study provides empirical evidence suggesting that populations of long-lived organisms such as forest trees can adapt locally, even at short-time scales., (© 2023 The Authors. Molecular Ecology published by John Wiley & Sons Ltd.)

In forest communities, light competition is a key process for community assembly. Species' differences in seedling and sapling tolerance to shade cast by overstory trees is thought to determine species composition at late-successional stages. Most forests are distant from these late-successional equilibria, impeding a formal evaluation of their potential species composition. To extrapolate competitive equilibria from short-term data, we therefore introduce the JAB model, a parsimonious dynamic model with interacting size-structured populations, which focuses on sapling demography including the tolerance to overstory competition. We apply the JAB model to a two-"species" system from temperate European forests, that is, the shade-tolerant species Fagus sylvatica L. and the group of all other competing species. Using Bayesian calibration with prior information from external Slovakian national forest inventory (NFI) data, we fit the JAB model to short time series from the German NFI. We use the posterior estimates of demographic rates to extrapolate that F. sylvatica will be the predominant species in 94% of the competitive equilibria, despite only predominating in 24% of the initial states. We further simulate counterfactual equilibria with parameters switched between species to assess the role of different demographic processes for competitive equilibria. These simulations confirm the hypothesis that the higher shade tolerance of F. sylvatica saplings is key for its long-term predominance. Our results highlight the importance of demographic differences in early life stages for tree species assembly in forest communities., Competing Interests: No competing interests., (© 2023 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Roots of forest trees are colonized by a diverse spectrum of ectomycorrhizal (EM) fungal species differing in their nitrogen (N) acquisition abilities. Here, we hypothesized that root N gain is the result of EM fungal diversity or related to taxon-specific traits for N uptake. To test our hypotheses, we traced 15 N enrichment in fine roots, coarse roots and taxon-specific ectomycorrhizas in temperate beech forests in two regions and three seasons, feeding 1 mM NH 4 NO 3 labelled with either 15 NH 4 + or 15 NO 3 - . We morphotyped > 45 000 vital root tips and identified 51 of 53 detected EM species by sequencing. EM root tips exhibited strong, fungal taxon-specific variation in 15 N enrichment with higher NH 4 + than NO 3 - enrichment. The translocation of N into the upper parts of the root system increased with increasing EM fungal diversity. Across the growth season, influential EM species predicting root N gain were not identified, probably due to high temporal dynamics of the species composition of EM assemblages. Our results support that root N acquisition is related to EM fungal community-level traits and highlight the importance of EM diversity for tree N nutrition., (© 2023 The Authors New Phytologist © 2023 New Phytologist Foundation.)

Prolonged drought and susceptibility to biotic stressors induced an extensive calamity in Norway spruce (Picea abies (L.) Karst.) and widespread crown defoliation in European beech (Fagus sylvatica L.) in Central Europe. For future management decisions, it is crucial to link changes in canopy cover to site conditions. However, current knowledge on the role of soil properties for drought-induced forest disturbance is limited due to the scarcity and low spatial resolution of soil information. We present a fine-scale assessment on the role of soil properties for forest disturbance in Norway spruce and European beech derived from optical remote sensing. A forest disturbance modeling framework based on Sentinel-2 time series was applied on 340 km 2 in low mountain ranges of Central Germany. Spatio-temporal information on forest disturbance was calculated at 10 m spatial resolution in the period 2019-2021 and intersected with high-resolution soil information (1:10,000) based on roughly 2850 soil profiles. We found distinct differences in disturbed area, depending on soil type, texture, stoniness, effective rooting depth and available water capacity (AWC). For spruce, we found a polynomial relationship between AWC (R 2  = 0.7) and disturbance, with highest disturbed area (65 %) for AWC between 90 and 160 mm. Interestingly, we found no evidence for generally higher disturbance on shallow soils, although stands on the deepest soils were significantly less affected. Noteworthy, sites affected first did not necessarily exhibit highest proportions of disturbed area post-drought, indicating recovery or adaptation. We conclude that site- and species-specific understanding of drought impacts benefits from a combination of remote sensing and fine-scale soil information. Since our approach revealed which sites were affected first and most, it qualifies for prioritizing in situ monitoring activities to most vulnerable stands in acute drought conditions as well as for developing long-term strategies for reforestation and site-specific risk assessment for precision forestry., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Mast seeding causes strong fluctuations in populations of forest animals. Thus, this phenomenon can be used as a natural experiment to examine how variation in host abundance affects parasite loads. We investigated fleas infesting yellow-necked mice in beech forest after 2 mast and 2 non-mast years. We tested 2 mutually exclusive scenarios: (1) as predicted by classical models of density-dependent transmission, an increase in host density will cause an increase in ectoparasite abundance (defined as the number of parasites per host), versus (2) an increase in host density will cause a decline in flea abundance ("dilution," which is thought to occur when parasite population growth is slower than that of the host). In addition, we assessed whether masting alters the relationship between host traits (sex and body mass) and flea abundance. We found a hump-shaped relationship between host and flea abundance. Thus, the most basic predictions are too simple to describe ectoparasite dynamics in this system. In addition, masting modified seasonal dynamics of flea abundance, but did not affect the relationship between host traits and flea abundance (individuals with the highest body mass hosted the most fleas; after controlling for body mass, parasite abundance did not vary between sexes). Our results demonstrate that pulses of tree reproduction can indirectly, through changes in host densities, drive patterns of ectoparasite infestation., (© 2022 The Authors. Integrative Zoology published by International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

Against the backdrop of global change, the intensity, duration, and frequency of droughts are projected to increase and threaten forest ecosystems worldwide. Tree responses to drought are complex and likely to vary among species, drought characteristics, and site conditions. Here, we examined the drought response patterns of three major temperate tree species, s. fir (Abies alba), E. beech (Fagus sylvatica), and N. spruce (Picea abies), along an ecological gradient in the South - Central - East part of Germany that included a total of 37 sites with varying climatic and soil conditions. We relied on annual tree-ring data to assess the influence of different drought characteristics and (micro-) site conditions on components of tree resilience and to detect associated temporal changes. Our study revealed that nutrient regime, drought frequency, and hydraulic conditions in the previous and subsequent years were the main determinants of drought responses, with pronounced differences among species. Specifically, we found that (a) higher drought frequency was associated with higher resistance and resilience for N. spruce and E. beech; (b) more favorable climatic conditions in the two preceding and following years increased drought resilience and determined recovery potential of E. beech after extreme drought; (c) a site's nutrient regime, rather than micro-site differences in water availability, determined drought responses, with trees growing on sites with a balanced nutrient regime having a higher capacity to withstand extreme drought stress; (d) E. beech and N. spruce experienced a long-term decline in resilience. Our results indicate that trees under extreme drought stress benefit from a balanced nutrient supply and highlight the relevance of water availability immediately after droughts. Observed long-term trends confirm that N. spruce is suffering from persistent climatic changes, while s. fir is coping better. These findings might be especially relevant for monitoring, scenario analyses, and forest ecosystem management., Competing Interests: Declaration of competing interest None declared., (Copyright © 2023 The Authors. Published by Elsevier B.V. All rights reserved.)

In forests, mycorrhizal fungi regulate carbon (C) and nitrogen (N) dynamics. We evaluated the interplay among ectomycorrhizas (ECM), ecosystem C fluxes, tree productivity, C and N exchange and isotopic fractionation along the soil-ECM-plant continuum in a Mediterranean beech forest. From bud break to leaf shedding, we monitored: net ecosystem exchange (NEE, a measure of the net exchange of C between an ecosystem and the atmosphere), leaf area index, stem growth, N concentration, δ 13 C and δ 15 N in rhizosphere soil, ectomycorrhizal fine root tips (ERT), ECM-free fine root portions (NCR) and leaves. Seasonal changes in ERT relative biomass were strictly related to NEE and mimicked those detected in the radial growth. The analysis of δ 13 C in ERT, leaves and NCR highlighted the impact of canopy photosynthesis on ERT development and an asynchronous seasonal C allocation strategy between ERT and NCR at the root tips level. Concerning N, δ 15 N of leaves was negatively related to that of ERT and dependent on seasonal 15 N differences between ERT and NCR. Our results unravel a synchronous C allocation towards ERT and tree stem driven by the increasing NEE in spring-early summer. Moreover, they highlighted a phenology-dependent 15 N fractionation during N transfer from ECM to their hosts. This evidence, obtained in mature beech trees under natural conditions, may improve the knowledge of Mediterranean forests functionality., (© 2022 John Wiley & Sons Ltd.)

Extreme climate events such as late spring frosts (LSFs) negatively affect productivity and tree growth in temperate beech forests. However, detailed information on how these forests recover after such events are still missing. We investigated how LSFs affected forest cover and radial growth in European beech (Fagus sylvatica L.) populations located at different elevations at four sites in the Italian Apennines, where LSFs have been recorded. We combined tree-ring and remote-sensing data to analyse the sensitivity and recovery capacity of beech populations to LSFs. Using daily temperature records, we reconstructed LSF events and assessed legacy effects on growth. We also evaluated the role played by elevation and stand structure as modulators of LSFs impacts. Finally, using satellite images we computed Normalized Difference Vegetation Index (NDVI), Enhanced Vegetation Index (EVI) and LAI (Leaf Area Index) to evaluate the post-LSF canopy recovery. The growth reduction in LSF-affected trees ranged from 36 % to 84 %. We detected a negative impact of LSF on growth only during the LSF year, with growth recovery occurring within 1-2 years after the event. LSF-affected stands featured low vegetation indices until late June, i.e. on average 75 days after the frost events. We did not find a clear relationship between beech forest elevation and occurrence of LSFs defoliations. Our results indicate a high recovery capacity of common beech and no legacy effects of LSFs., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Leaf cuticular waxes play an important role in reducing evapotranspiration via diffusion. However, the ability of mature trees to regulate the biosynthesis of waxes to changing conditions (e.g., drought, light exposition) remain an open question, especially during the late growing season. This holds also true for one of the most widely distributed trees in Central Europe, the European beech tree ( Fagus sylvatica L. ). In order to investigate the ongoing formation of wax constituents like alkanes and fatty acids, we conducted a 13 CO 2 pulse-chase labelling experiment on sun-exposed and shaded branches of a mature beech tree during the late summer 2018. The 13 C-label was traced via compound-specific δ 13 C isotope analysis of n -alkanes and fatty acids to determine the de-novo biosynthesis within these compound classes. We did not observe a significant change in lipid concentrations during the late growing season, but we found higher n -alkane concentrations in sun-exposed compared to shaded leaves in August and September. The n- alkane and fatty acid composition showed ongoing modifications during the late growing season. Together with the uptake and following subsequent decrease of the 13 C-label, this suggests ongoing de-novo biosynthesis, especially of fatty acids in European beech leaves. Moreover, there is a high variability in the 13 C-label among individual branches and between sun-exposed and shaded leaves. At the same time, sun-exposed leaves invest more of the assimilated C into secondary metabolites such as lipids than shaded leaves. This indicates that the investigated mature beech tree could adjust its lipid production and composition in order to acclimate to changes in microclimates within the tree crown and during the investigated period., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2023 Speckert, Petibon and Wiesenberg.)

Genetic diversity influences the evolutionary potential of forest trees under changing environmental conditions, thus indirectly the ecosystem services that forests provide. European beech ( Fagus sylvatica L.) is a dominant European forest tree species that increasingly suffers from climate change-related die-back. Here, we conducted a systematic literature review of neutral genetic diversity in European beech and created a meta-data set of expected heterozygosity ( He ) from all past studies providing nuclear microsatellite data. We propose a novel approach, based on population genetic theory and a min-max scaling to make past studies comparable. Using a new microsatellite data set with unprecedented geographic coverage and various re-sampling schemes to mimic common sampling biases, we show the potential and limitations of the scaling approach. The scaled meta-dataset reveals the expected trend of decreasing genetic diversity from glacial refugia across the species range and also supports the hypothesis that different lineages met and admixed north of the European mountain ranges. As a result, we present a map of genetic diversity across the range of European beech which could help to identify seed source populations harboring greater diversity and guide sampling strategies for future genome-wide and functional investigations of genetic variation. Our approach illustrates how to combine information from several nuclear microsatellite data sets to describe patterns of genetic diversity extending beyond the geographic scale or mean number of loci used in each individual study, and thus is a proof-of-concept for synthesizing knowledge from existing studies also in other species., Supplementary Information: The online version contains supplementary material available at 10.1007/s11295-022-01577-4., Competing Interests: Conflict of interestThe authors declare no competing interests., (© The Author(s) 2022.)

During the particularly severe hot summer drought in 2018, widespread premature leaf senescence was observed in several broadleaved tree species in Central Europe, particularly in European beech (Fagus sylvatica L.). For beech, it is yet unknown whether the drought evoked a decline towards tree mortality or whether trees can recover in the longer term. In this study, we monitored crown dieback, tree mortality and secondary drought damage symptoms in 963 initially live beech trees that exhibited either premature or normal leaf senescence in 2018 in three regions in northern Switzerland from 2018 to 2021. We related the observed damage to multiple climate- and stand-related parameters. Cumulative tree mortality continuously increased up to 7.2% and 1.3% in 2021 for trees with premature and normal leaf senescence in 2018, respectively. Mean crown dieback in surviving trees peaked at 29.2% in 2020 and 8.1% in 2019 for trees with premature and normal leaf senescence, respectively. Thereafter, trees showed first signs of recovery. Crown damage was more pronounced and recovery was slower for trees that showed premature leaf senescence in 2018, for trees growing on drier sites, and for larger trees. The presence of bleeding cankers peaked at 24.6% in 2019 and 10.7% in 2020 for trees with premature and normal leaf senescence, respectively. The presence of bark beetle holes peaked at 22.8% and 14.8% in 2021 for trees with premature and normal leaf senescence, respectively. Both secondary damage symptoms occurred more frequently in trees that had higher proportions of crown dieback and/or showed premature senescence in 2018. Our findings demonstrate context-specific differences in beech mortality and recovery reflecting the importance of regional and local climate and soil conditions. Adapting management to increase forest resilience is gaining importance, given the expected further beech decline on dry sites in northern Switzerland., (© 2022 The Authors. Plant Biology published by John Wiley & Sons Ltd on behalf of German Society for Plant Sciences, Royal Botanical Society of the Netherlands.)

Traditionally focussed on maximising productivity, forest management increasingly has to consider other functions performed by the forest stands, such as biodiversity conservation. Terrestrial plant communities typically possess a hump-back relationship between biomass productivity and plant species richness. However, there is evidence of a reverse relationship in forests dominated by beech, one of the most competitive and widespread tree species in temperate Europe. To fully explore the tree productivity-species richness relationship, we investigated above- and below-ground drivers of understorey plant species richness. We focussed on managed beech forests growing along an elevation gradient in Central Europe. We found that the lowest understorey plant diversity was under conditions optimal for beech. Tree fine root mass, canopy openness, soil C/N ratio, the interaction between tree fine root mass and stoniness, and stand structural diversity explain the variation of understorey species richness. We show that the competition for soil resources is the main driver of plant species diversity in managed forests; maximising beech growth in optimal conditions may thus come at the expense of understorey plant richness., Competing Interests: Declaration of competing interest We have no conflict of interest to declare., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Temperate trees could cope with climate change through phenotypic plasticity of phenological key events or adaptation in situ via selection on genetic variation. However, the relative contribution of local adaptation and phenotypic plasticity to phenological change is unclear for many ecologically important tree species. Here, we analyzed the leaf-out data of European beech (Fagus sylvatica L.) from 50 provenances planted in 7 trial sites. We first constructed a function between chilling accumulation (CA) and photoperiod-associated heat requirement (PHR) of leaf-out date for each provenance and quantified the relationship between parameters of the CA-PHR function and climatic variables at provenance origins by using the random forest model. Furthermore, we used the provenance-specific CA-PHR function to simulate future leaf-out dates under two climate change scenarios (RCP 4.5 and 8.5) and two assumptions (no adaptation and adaptation). The results showed that both CA, provenance, and their interactions affected the PHR of leaf-out. The provenances from southeastern Europe exhibited a stronger response of PHR to CA and thus flushed earlier than northwestern provenances. The parameters of the CA-PHR function were connected with climatic variables (e.g., mean diurnal temperature range, temperature seasonality) at the originating sites of each provenance. If only considering the phenotypic plasticity, the leaf-out date of European beech in 2070-2099 will advance by 6.8 and 9.0 days on average relative to 1951-2020 under RCP 4.5 and RCP 8.5, respectively. However, if F. sylvatica adapts to future climate change by adopting the current strategy, the advance of the leaf-out date will weaken by 1.4 and 3.4 days under RCP 4.5 and RCP 8.5, respectively. Our results suggest that the European beech could slow down its spring phenological advances and reduce its spring frost risk if it adopts the current strategy to adapt to future climate change., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Temperature (T) and vapour pressure deficit (VPD) are important drivers of plant hydraulic conductivity, growth, mortality, and ecosystem productivity, independently of soil water availability. Our goal was to disentangle the effects of T and VPD on plant hydraulic responses. Young trees of Fagus sylvatica L., Quercus pubescens Willd. and Quercus ilex L. were exposed to a cross-combination of a T and VPD manipulation under unlimited soil water availability. Stem hydraulic conductivity and leaf-level hydraulic traits (e.g., gas exchange and osmotic adjustment) were tracked over a full growing season. Significant loss of xylem conductive area (PLA) was found in F. sylvatica and Q. pubescens due to rising VPD and T, but not in Q. ilex. Increasing T aggravated the effects of high VPD in F. sylvatica only. PLA was driven by maximum hydraulic conductivity and minimum leaf conductance, suggesting that high transpiration and water loss after stomatal closure contributed to plant hydraulic stress. This study shows for the first time that rising VPD and T lead to losses of stem conductivity even when soil water is not limiting, highlighting their rising importance in plant mortality mechanisms in the future., (© 2022 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

In the future, climate change is expected to affect the spatial distribution of most tree species in Europe. The European beech ( Fagus sylvatica ), a drought-sensitive tree species, is currently distributed throughout Europe, where it is an ecologically and economically important species. In Slovenia, the European beech represents 33% of the growing stock, but such a proportion greatly varies across Europe. Whether such a variation is related to the climate environmental gradients or because of historical or management decisions is an as-yet unexplored question. For this study, we employed the Slovenian Forests Service inventory, where the proportion of beech in the forest stock has been monitored in 341,341 forest stands across the country. Modeled climate data from the SLOCLIM database, calculated for each of the stands, was also used to test the hypothesis that although beech forests have always been influenced by human activity, the dominance of beech trees in forest stands is at least partially dictated by the climate. The results showed the distribution of the main climate variables (annual precipitation, the share of summer and spring precipitation, and annual maximum and minimum temperatures) and how they affect the current dominance of beech trees at the stand level. Due to the large number and variability of forest stands studied, the results should be transferable to better understand and manage the climatic suitability and risks of Fagus sylvatica . The modeled data is publicly available in the web repository Zenodo.

The objective of this study was to investigate the role of the oomycete Phytophthora × cambivora in the decline affecting European beech ( Fagus sylvatica ) in the Nebrodi Regional Park (Sicily, southern Italy). In a survey of a beech forest stand in the heart of the park, Phytophthora × cambivora was the sole Phytophthora species recovered from the rhizosphere soil and fine roots of trees. Both A1 and A2 mating type isolates were found. Direct isolation from the stem bark of trees showing severe decline symptoms and bleeding stem cankers yielded exclusively P. gonapodyides , usually considered as an opportunistic pathogen. The mean inoculum density of P. × cambivora in the rhizosphere soil, as determined using the soil dilution plating method and expressed in terms of colony forming units (cfus) per gm of soil, the isolation frequency using leaf baiting, and the percentage of infected fibrous roots from 20 randomly selected beech trees with severe decline symptoms (50 to 100 foliage transparency classes) were 31.7 cfus, 80%, and 48.6%, respectively. These were significantly higher than the corresponding mean values of 20 asymptomatic or slightly declining trees, suggesting P. × cambivora is a major factor responsible for the decline in the surveyed stand.

Two deciduous forest ecosystems, one dominated by Fagus sylvatica and a mixed one with Quercus cerris and Quercus frainetto , were monitored from an ecophysiological perspective during a five-year period, in order to assess seasonal fluctuations, establish links between phenology and ecophysiology, and reveal climatic controls. Field measurements of leaf area index (LAI), chlorophyll content, leaf specific mass (LSM), water potential (Ψ) and leaf photosynthesis (Aleaf) were performed approximately on a monthly basis. LAI, chlorophylls and LSM fluctuations followed a recurrent pattern yearly, with increasing values during spring leaf burst and expansion, relatively stable values during summer and decreasing values during autumn senescence. However, pre-senescence leaf fall and chlorophyll reductions were evident in the driest year. The dynamically responsive Aleaf and Ψ presented considerable inter-annual variation. Both oak species showed more pronounced depressions of Aleaf and Ψ compared to beech, yet the time-point of their appearance coincided and was the same for all species each year. Spring temperature had a positive role in the increasing phase of all ecophysiological processes while rising autumn temperature resulted in retarded senescence. Precipitation showed asymmetric effects on the measured ecophysiological parameters. The between-species differences in responses, climate sensitivity and climate memory are identified and discussed.

Sunflecks are transient patches of direct radiation that provide a substantial proportion of the daily irradiance to leaves in the lower canopy. In this position, faster photosynthetic induction would allow for higher sunfleck-use efficiency, as is commonly reported in the literature. Yet, when sunflecks are too few and far between, it may be more beneficial for shade leaves to prioritize efficient photosynthesis under shade. We investigated the temporal dynamics of photosynthetic induction, recovery under shade, and stomatal movement during a sunfleck, in sun and shade leaves of Fagus sylvatica from three provenances of contrasting origin. We found that shade leaves complete full induction in a shorter time than sun leaves, but that sun leaves respond faster than shade leaves due to their much larger amplitude of induction. The core-range provenance achieved faster stomatal opening in shade leaves, which may allow for better sunfleck-use efficiency in denser canopies and lower canopy positions. Our findings represent a paradigm shift for future research into light fluctuations in canopies, drawing attention to the ubiquitous importance of sunflecks for photosynthesis, not only in lower-canopy leaves where shade is prevalent, but particularly in the upper canopy where longer sunflecks are more common due to canopy openness., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

European beech (Fagus sylvatica) was among the most affected tree species during the severe 2018 European drought. It not only suffered from instant physiological stress but also showed severe symptoms of defoliation and canopy decline in the following year. To explore the underlying mechanisms, we used the Swiss-Canopy-Crane II site and studied in branches of healthy and symptomatic trees the repair of hydraulic function and concentration of carbohydrates during the 2018 drought and in 2019. We found loss of hydraulic conductance in 2018, which did not recover in 2019 in trees that developed defoliation symptoms in the year after drought. Reduced branch foliation in symptomatic trees was associated with a gradual decline in wood starch concentration throughout summer 2019. Visualization of water transport in healthy and symptomatic branches in the year after the drought confirmed the close relationship between xylem functionality and supported branch leaf area. Our findings showed that embolized xylem does not regain function in the season following a drought and that sustained branch hydraulic dysfunction is counterbalanced by the reduction in supported leaf area. It suggests acclimation of leaf development after drought to mitigate disturbances in canopy hydraulic function., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

The onset of the growing season in temperate forests is relevant for forest ecology and biogeochemistry and is known to occur earlier with climate change. Variation in tree phenology among individual trees of the same stand and species, however, is not well understood. Yet, natural selection acts on this inter-individual variation, which consequently affects the adaptive potential to ongoing environmental changes. Budburst dates of 146 mature individuals of Fagus sylvatica , the dominant natural forest tree of central Europe, were recorded over 12 years in one forest stand of 1 ha in the Müritz National Park, Germany. The tree-specific location, topographical differences, as well as social status, were measured to explain the inter-individual variation in budburst. Furthermore, inter-individual differences in bud dormancy were quantified. Additional phenology and weather data across Germany from 405 sites over a 25-year period was used to put the insights from the single stand into perspective. Consistent phenological ranking over the years with respect to early and late flushing trees was observed within the single forest stand, with 23 trees consistently flushing 3-6 days earlier and 22 trees consistently flushing 3-10 days later than the median. Trees flushing consistently early varied most in their spring budburst dates and were less dormant than late-flushing trees already in mid-winter. The higher variation in earlier flushing trees was best explained by a slower warming rate during their budburst period in the observed stand as well as across Germany. Likewise, years with a lower warming rate during the budburst period were more variable in budburst dates. The rate of warming during spring time is crucial to accurately project future within-species variation and the resulting adaptive potential in spring phenology of dominant forest tree species., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Malyshev, van der Maaten, Garthen, Maß, Schwabe and Kreyling.)

Environmental conditions affect tree-ring width (TRW), wood structure, and, consequently, wood density, which is one of the main wood quality indicators. Although studies on inter- and intra-annual variability in tree-ring features or density exist, studies demonstrating a clear link between wood structure on a cellular level and its effect on wood density on a macroscopic level are rare. Norway spruce with its simple coniferous structure and European beech, a diffuse-porous angiosperm species were selected to analyze these relationships. Increment cores were collected from both species at four sites in Slovenia. In total, 24 European beech and 17 Norway spruce trees were sampled. In addition, resistance drilling measurements were performed just a few centimeters above the increment core sampling. TRW and quantitative wood anatomy measurements were performed on the collected cores. Resistance drilling density values, tree-ring (TRW, earlywood width-EWW, transition-TWW, and latewood width-LWW) and wood-anatomical features (vessel/tracheid area and diameter, cell density, relative conductive area, and cell wall thickness) were then averaged for the first 7 cm of measurements. We observed significant relationships between tree-ring and wood-anatomical features in both spruce and beech. In spruce, the highest correlation values were found between TRW and LWW. In beech, the highest correlations were observed between TRW and cell density. There were no significant relationships between wood-anatomical features and resistance drilling density in beech. However, in spruce, a significant negative correlation was found between resistance drilling density and tangential tracheid diameter, and a positive correlation between resistance drilling density and both TWW + LWW and LWW. Our findings suggest that resistance drilling measurements can be used to evaluate differences in density within and between species, but they should be improved in resolution to be able to detect changes in wood anatomy., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Arnič, Krajnc, Gričar and Prislan.)

The critical level of ozone flux for forest trees is based entirely on biomass data from fumigation experiments with saplings, mostly in open-top chambers. Extrapolation to mature forests asks, therefore, for validation, which may be performed by epidemiological data analysis. This requires a multivariable regression analysis with a number of covariates to account for potential confounding factors. The present paper analyses the ozone sensitivity of volume increments of mature European beech ( Fagus sylvatica ) and Norway spruce ( Picea abies ), with the addition, or removal, of covariates. The comparison of the epidemiological dose-response relationship with experimental data shows very good agreement in beech and a more sensitive relationship in the epidemiological analysis of Norway spruce compared to the experiments. In Norway spruce, there was also a strong interaction between the effects of ozone and temperature; at high July temperatures, the ozone effect was stronger. This interaction may explain the disagreement between the epidemiological study and the experiments, of which the majority were performed in Sweden.

Mineral nitrogen (N) is a major nutrient showing strong fluctuations in the environment due to anthropogenic activities. The acquisition and translocation of N to forest trees are achieved mainly by highly diverse ectomycorrhizal fungi (EMF) living in symbioses with their host roots. Here, we examined colonized root tips to characterize the entire root-associated fungal community by DNA metabarcoding-Illumina sequencing of the fungal internal transcribed spacer 2 (ITS2) molecular marker and used RNA sequencing to target metabolically active fungi and the plant transcriptome after N application. The study was conducted with beech (Fagus sylvatica L.), a dominant tree species in central Europe, grown in native forest soil. We demonstrate strong enrichment of 15 N from nitrate or ammonium in the ectomycorrhizal roots by stable-isotope labeling. The relative abundance of the EMF members in the fungal community was correlated with their transcriptional abundances. The fungal metatranscriptome covered Kyoto Encyclopedia of Genes and Genomes (KEGG) and Eukaryotic Orthologous Groups (KOG) categories similar to those of model fungi and did not reveal significant changes related to N metabolization but revealed species-specific transcription patterns, supporting trait stability. In contrast to the resistance of the fungal metatranscriptome, the transcriptome of the host exhibited dedicated nitrate- or ammonium-responsive changes with the upregulation of transporters and enzymes required for nitrate reduction and a drastic enhancement of glutamine synthetase transcript levels, indicating the channeling of ammonium into the pathway for plant protein biosynthesis. Our results support that naturally assembled fungal communities living in association with the tree roots buffer nutritional signals in their own metabolism but do not shield plants from high environmental N levels. IMPORTANCE Although EMF are well known for their role in supporting tree N nutrition, the molecular mechanisms underlying N flux from the soil solution into the host through the ectomycorrhizal pathway remain widely unknown. Furthermore, ammonium and nitrate availability in the soil solution is subject to frequent oscillations that create a dynamic environment for the tree roots and associated microbes during N acquisition. Therefore, it is important to understand how root-associated mycobiomes and the tree roots handle these fluctuations. We studied the responses of the symbiotic partners by screening their transcriptomes after a sudden environmental flux of nitrate or ammonium. We show that the fungi and the host respond asynchronously, with the fungi displaying resistance to increased nitrate or ammonium and the host dynamically metabolizing the supplied N sources. This study provides insights into the molecular mechanisms of the symbiotic partners operating under N enrichment in a multidimensional symbiotic system.

Legacies of past climate conditions and historical management govern forest productivity and tree growth. Understanding how these processes interact and the timescales over which they influence tree growth is critical to assess forest vulnerability to climate change. Yet, few studies address this issue, likely because integrated long-term records of both growth and forest management are uncommon. We applied the stochastic antecedent modelling (SAM) framework to annual tree-ring widths from mixed forests to recover the ecological memory of tree growth. We quantified the effects of antecedent temperature and precipitation up to 4 years preceding the year of ring formation and integrated management effects with records of harvesting intensity from historical forest management archives. The SAM approach uncovered important time periods most influential to growth, typically the warmer and drier months or seasons, but variation among species and sites emerged. Silver fir responded primarily to past climate conditions (25-50 months prior to the year of ring formation), while European beech and Scots pine responded mostly to climate conditions during the year of ring formation and the previous year, although these responses varied among sites. Past management and climate interacted in such a way that harvesting promoted growth in young silver fir under wet and warm conditions and in old European beech under drier and cooler conditions. Our study shows that the ecological memory associated with climate legacies and historical forest management is species-specific and context-dependent, suggesting that both aspects are needed to properly evaluate forest functioning under climate change., Supplementary Information: The online version contains supplementary material available at 10.1007/s10021-021-00650-8., (© The Author(s) 2021.)

European beech is an important component of European lowland forests in terms of ecology, and produces irregular seeds categorized as intermediate due to their limited longevity. Removal of the excess of reactive oxygen species is crucial for redox homeostasis in growing plant tissues. Hydrogen peroxide (H 2 O 2 ) is detoxified via the plant-specific ascorbate-glutathione cycle, and enzymatically, mainly by catalase (CAT). The reduced and oxidized (redox) forms of ascorbate (AsA, DHA) and glutathione (GSH, GSSG) decreased during maturation as the content of redox forms of nicotinamide adenine dinucleotide (NADH, NAD + ) phosphate (NADPH, NADP + ), cofactors of ascorbate-glutathione enzymes, declined and limited this cycle. The degree of oxidation of glutathione peaked at approximately 80%, at the exact time when the NADP content was the lowest and the NADPH/NADP + ratio reached the highest values. The glutathione pool was reflected in changes in the NADP pool, both in embryonic axes (R 2 = 0.61) and in cotyledons (R 2 = 0.98). A large excess of NADPH was reported in embryonic axes, whereas cotyledons displayed more unified levels of NADP redox forms. As a result, anabolic redox charge and reducing power were higher in embryonic axes. CAT was recognized as two proteins, and the abundance of the 55 kDa protein was correlated with all redox forms of ascorbate, glutathione, NAD, and NADP, whereas the 37 kDa protein was oppositely regulated in embryonic axes and cotyledons. Here, we discuss the role of NAD(P) in the regulation of the ascorbate-glutathione cycle, catalase, and seed longevity concerning a putative role of NAD(P)H as a redox biomarker involved in predefining seed quality, because NAD(P)H-derived redox homeostasis was found to be better controlled in embryonic axes than cotyledons.

Background: The tick Ixodes ricinus is an important vector of tick-borne diseases including Lyme borreliosis. In continental Europe, the nymphal stage of I. ricinus often has a bimodal phenology with a large spring peak and a smaller fall peak. There is consensus about the origin of the spring nymphal peak, but there are two alternative hypotheses for the fall nymphal peak. In the direct development hypothesis, larvae quest as nymphs in the fall of the same year that they obtained their larval blood meal. In the developmental diapause hypothesis, larvae overwinter in the engorged state and quest as nymphs one year after they obtained their larval blood meal. These two hypotheses make different predictions about the time lags that separate the larval blood meal and the density of questing nymphs (DON) in the spring and fall., Methods: Inter-annual variation in seed production (masting) by deciduous trees is a time-lagged index for the density of vertebrate hosts (e.g., rodents) which provide blood meals for larval ticks. We used a long-term data set on the masting of the European beech tree and a 15-year study on the DON at 4 different elevation sites in western Switzerland to differentiate between the two alternative hypotheses for the origin of the fall nymphal peak., Results: Questing I. ricinus nymphs had a bimodal phenology at the three lower elevation sites, but a unimodal phenology at the top elevation site. At the lower elevation sites, the DON in the fall was strongly correlated with the DON in the spring of the following year. The inter-annual variation in the densities of I. ricinus nymphs in the fall and spring was best explained by a 1-year versus a 2-year time lag with the beech tree masting index. Fall nymphs had higher fat content than spring nymphs indicating that they were younger. All these observations are consistent with the direct development hypothesis for the fall peak of I. ricinus nymphs at our study site. Our study provides new insight into the complex bimodal phenology of this important disease vector., Conclusions: Public health officials in Europe should be aware that following a strong mast year, the DON will increase 1 year later in the fall and 2 years later in the spring. Studies of I. ricinus populations with a bimodal phenology should consider that the spring and fall peak in the same calendar year represent different generations of ticks., (© 2021. The Author(s).)