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Leaf morphological and physiological traits control the carbon and water relations of mature trees and are determinants of drought tolerance, but it is not well understood how they are modified in response to water deficits. We analysed five sun-canopy leaf traits (mean leaf size (LS), specific leaf area (SLA), Huber value (HV), water potential at turgor loss point (Ψtlp ) and foliar carbon isotope signature (δ13 C)) in European beech (Fagus sylvatica L.) across three precipitation gradients sampled in moist (2010), dry (2019) and very dry (2018) summers, and tested their response to short-term water deficits (climatic water balance (CWB) preceding sample collection) and long-term water availability (mean annual precipitation (MAP), plant-available soil water capacity (AWC) and neighbourhood competition). Across the 34 sites, LS varied seven-fold (3.9-27.0 cm2 ), SLA four-fold (77.1-306.9 cm²·g-1 ) and HV six-fold (1.0-6.65 cm2 ·m-2 ). In the 2018 dataset, LS showed a negative and HV a positive relationship to MAP, which contradicts relations found in multi-species samples. Average Ψtlp ranged from -1.90 to -2.62 MPa and decreased across the sites with decreasing CWB in the month prior to measurement, as well as with decreasing MAP and AWC in 2019. Studied leaf traits varied considerably between years, suggesting that mast fruiting and the severe 2018 drought caused the formation of smaller leaves. We conclude that sun-canopy leaf traits of European beech exhibit considerable plasticity in response to climatic and edaphic aridity, and that osmotic adjustment may be an important element in the drought response strategy of this anisohydric tree species.

Using hierarchical clustering with unweighted pair-group method with arithmetic mean (UPGMA) we arranged 602 phytosociological releves of beech forests on the present forest line, mainly from the Julian Alps and the Trnovo Forest Plateau (we also included the releves from the Karawanks and the Kamnik Alps) into 32 clusters. Based on their analysis and comparison with previously described similar (alti)montane-subalpine beech communities we classified most of the releves into the association Polysticho lonchitis-Fagetum and its new subassociations ericetosum carneae, cardaminetosum trifoliae, luzuletosum niveae, luzuletosum luzuloidis, calamagrostieteosum variae, allietosum victorialis, adoxetosum moschatellinae, stellarietosum nemorum and several new variants. The altitude of the studied stands is predominantly 1400 to 1550 m (the upper line is at 1660 m); they occur at all aspects, frequently on steep and very steep slopes, mainly on limestone and dolomite limestone, the predominant soil type is rendzina. These stands are species rich (on average 61 species per releve, altogether more than 500 vascular plants) and have many species in common with the stands of associations Rhododendro hirsuti-Fagetum and Rhodothamno-Laricetum. They belong to the habitat type of Community interest 91K0 Illyrian Fagus sylvatica forests and are some of the best preserved forest communities in the Southeastern Alps and northwestern Dinaric Alps, comprising small areas of virgin forest remnants and performing an important protective role as a biotope.

High genetic variation and extensive gene flow may help forest trees with adapting to ongoing climate change, yet the genetic bases underlying their adaptive potential remain largely unknown. We investigated range-wide patterns of potentially adaptive genetic variation in 64 populations of European beech (Fagus sylvatica L.) using 270 SNPs from 139 candidate genes involved either in phenology or in stress responses. We inferred neutral genetic structure and processes (drift and gene flow) and performed differentiation outlier analyses and gene-environment association (GEA) analyses to detect signatures of divergent selection.Beech range-wide genetic structure was consistent with the species’ previously identified postglacial expansion scenario and recolonization routes. Populations showed high diversity and low differentiation along the major expansion routes. A total of 52 loci were found to be putatively under selection and 15 of them turned up in multiple GEA analyses. Temperature and precipitation related variables were equally represented in significant genotype-climate associations. Signatures of divergent selection were detected in the same proportion for stress response and phenology-related genes. The range-wide adaptive genetic structure of beech appears highly integrated, suggesting a balanced contribution of phenology and stress-related genes to local adaptation, and of temperature and precipitation regimes to genetic clines. Our results imply a best-case scenario for the maintenance of high genetic diversity during range shifts in beech (and putatively other forest trees) with a combination of gene flow maintaining within-population neutral diversity and selection maintaining between-population adaptive differentiation.

Atmospheric deposition of nitrogen (N) has exceeded its demand for plant increment in forest ecosystems in Germany. High N inputs increased plant growth, the internal N cycling within the ecosystem, the retention of N in soil and plant compartments, and the N output by seepage water. But the processes involved are not fully understood, notably the effect of fructification in European beech (Fagus sylvatica L.) on N fluxes. The frequency of fructification has increased together with air temperature and N deposition, but its impact on N fluxes and the sequestration of carbon (C) and N in soils have been hardly studied. A field experiment using 15 N-labeled leaf litter exchange was carried out over a 5.5-year period at seven long-term European beech (Fagus sylvatica L.) monitoring sites to study the impact of current mast frequency on N cycling. Mean annual leaf litterfall contained 35 kg N ha −1 , but about one-half of that was recovered in the soil 5.5 years after the establishment of the leaf litter 15 N exchange experiment. In these forests, fructification occurred commonly at intervals of 5 to 10 years, which has now changed to every 2 years as observed during this study period. Seed cupules contributed 51 % to the additional litterfall in mast years, which creates a high nutrient demand during their decomposition due to the very high ratios of C to N and C to phosphorus (P). Retention of leaf litter 15 N in the soil was more closely related to the production of total litterfall than to the leaf litterfall, indicating the role of seed cupules in the amount of leaf N retained in the soil. Higher mast frequency increased the mass of mean annual litterfall by about 0.5 Mg ha −1 and of litterfall N by 8.7 kg ha −1 . Mean net primary production (NPP) increased by about 4 %. Mean total N retention in soils calculated by input and output fluxes was unrelated to total litterfall, indicating that mast events were not the primary factor controlling total N retention in soils. Despite reduced N deposition since the 1990s, about 5.7 out of 20.7 kg N ha −1 deposited annually between 1994 and 2008 was retained in soils, notably at acid sites with high N / P and C / P ratios in the organic layers and mineral soils, indicating P limitation for litter decomposition. Trees retained twice as much N compared to soils by biomass increment, particularly in less acidic stands where the mineral soils had low C / N ratios. These results have major implications for our understanding of the C and N cycling and N retention in forest ecosystems. In particular the role of mast products in N retention needs more research in the future.

The current tendency towards the silvicultural promotion of mixed tree species has increased the variability in the crown structure within stands. This study shows how neighbouring trees can influence both the external crown features and internal wood properties of trees. Using terrestrial laser scanning, the crown features of 100 European beech trees (Fagus sylvatica L.) from pure beech stands and mixed stands of beech with Douglas fir (Pseudotsuga menziesii (Mirb.) Franco), Norway spruce (Picea abies (L.) Karst.), sessile oak (Quercus petraea (Matt.) Liebl.), and Scots pine (Pinus sylvestris L.) were recorded. After felling and sawing, the dynamic modulus of elasticity was determined on 1623 boards from the two lowest 4.1 m logs. Significant differences were found between beech trees from pure stands and those from beech–pine mixed stands in terms of crown volume (415 vs 766 m3), crown ratio (50.0% vs 71.5%), crown projection ratio (0.182 vs 0.253 m·cm−1), and branch angle (30.7° vs 54.1°). Multiple regression mixed models revealed significant relationships between timber stiffness and crown volume (–1.7 N·mm−2·m−3), crown ratio (–28.4 N·mm−2·%−1), and crown projection ratio (–9835 N·mm−2·m−1·cm). Thus, the crown morphology of broad-leaved species reflects the tree’s long-term competitive status and suggests indicators for the assessment of mechanical–physical wood properties.

Background and Aims Many recent studies emphasize that mixed species is a promising silvicultural option for sustainable ecosystem management under uncertain and risky future environmental conditions. However, compared with monocultures, knowledge of mixed stands is still rather fragmentary. This comprehensive study analysed the most common Central European tree species combinations to determine the extent to which mono-layered species mixing (1) can increase stand productivity and stem diameter growth, (2) increase stand density or growth efficiency, and (3) reduce competition and attenuate the relationship between stand density and stem diameter growth compared with mono-specific stands. Methods The study was based on 63 long-term experimental plots in Germany with repeated spatially explicit stand inventories. They covered mono-specific and mixed species stands of Norway spruce (Picea abies), silver fir (Abies alba), Scots pine (Pinus sylvestris), European beech (Fagus sylvatica), sessile oak (Quercus petraea), European ash (Fraxinus excelsior) and sycamore maple (Acer pseudoplatanus). Based on spatially explicit measurement, we quantified for each tree the intra- or inter-specific neighbourhood, local stand density and growth. We applied mixed models to analyse how inter-specific neighbourhoods modify stand productivity, stand density, growth efficiency, individual tree growth and the trade-off between individual tree growth and stand productivity. Key Results We found stand productivity gains of 7–53 % of mixed versus mono-specific stands continuing over the entire rotation. All mixtures achieved a 3–36 % higher leaf area index until advanced stand age. Stem diameter growth increased by up to 31 % in mixed stands. The growth efficiency of the leaf area was up to 31 % higher, except in mixtures of sessile oak and European beech. The trade-off between stem diameter growth and stand productivity was attenuated by the mixture. Conclusions The increased productivity was mainly based on a density increase in the case of Norway spruce/silver fir/European beech and sessile oak/European beech and it was based on a more efficient resource use given the same stand density in the case of Scots pine/European beech and European ash/sycamore maple. In the other species assemblages the increased productivity was based on a combination of density and efficiency increase. We hypothesize that the density effect may be site-invariant and mainly depends on the structural species complementarity. The efficiency increase of growth may depend on the growth-limiting factor that is remedied by mixture and thus be co-determined by the site conditions. For forest management, the results indicate increased stand and tree size growth by species mixing. For the common mixtures examined in this study the results show that thinning for the acceleration of stem growth requires less density reduction and causes less stand growth losses than in monocultures. We discuss the consequences of our findings for silvicultural prescriptions for mixed-species stands.

As of 2019, the total area of forest stands with the varying proportion of Scots pine (Pinus sylvestris L.) and beech (Fagus sylvatica L.) in composition on the Ukrainian Roztochchia was 11,251.3 ha and the area of pine stands involving European beech in composition was 6879.5 ha. Pine stands involving beech in composition are formed on the fresh, moist and boggy relatively pure and relatively rich soil condition. Pine forest stands (average share of pine exceeds 56%), which include beech in composition occupy more than 80% of the total area of interest. Mature and overmature forest stands dominate most of the landscape. More than 85% of stands are characterized by an average relative completeness. Undergrowth of deciduous tree species, which is mainly formed by European beech or hornbeam (Carpinus betulus L.) and rarely by common oak (Quercus robur L.) or sycamore maple (Acer pseudoplatanus), is available under a canopy of middle-aged, ripening and mature stands. The study points to the need to rethink forest management approaches and eventually to use such methods of forest cuttings that ensure natural regeneration not only of a European beech and common oak, but primarily of a pine. The practice of creating pine plantations in such conditions favourable for its growth and development should be used only in those cases when it is impossible to achieve natural regeneration of pine.

Mycorrhizal fungi play an important role for the nitrogen (N) supply of trees. The influence of different mycorrhizal types on N acquisition in tree–tree interactions is, however, not well understood, particularly with regard to the competition for growth-limiting N. We studied the effect of competition between temperate forest tree species on their inorganic and organic N acquisition in relation to their mycorrhizal type (i.e., arbuscular mycorrhiza or ectomycorrhiza). In a field experiment, we quantified net N uptake capacity from inorganic and organic N sources using 15N/13C stable isotopes for arbuscular mycorrhizal tree species (i.e., Acer pseudoplatanus L., Fraxinus excelsior L., and Prunus avium L.) as well as ectomycorrhizal tree species (i.e., Carpinus betulus L., Fagus sylvatica L., and Tilia platyphyllos Scop.). All species were grown in intra- and interspecific competition (i.e., monoculture or mixture). Our results showed that N sources were not used complementarily depending on a species’ mycorrhizal association, but their uptake rather depended on the competitor, indicating species-specific effects. Generally, ammonium was preferred over glutamine and glutamine over nitrate. In conclusion, our findings suggest that the inorganic and organic N acquisition of the studied temperate tree species is less regulated by mycorrhizal association but rather by the availability of specific N sources in the soil as well as the competitive environment of different tree species.

The potential utilisation of natural regeneration of European beech (Fagus sylvatica L.) for forest conversion has received little attention to date. Ecological knowledge is necessary to understand and predict successful natural regeneration of beech. The objective of this study was to improve understanding of what drives the occurrence of beech regeneration and, once regeneration is present, what drives its density. In the study, we utilised a forest inventory dataset provided by Sachsenforst, the state forestry service of Saxony, Germany. The dataset was derived from 8725 permanent plots. Zero-altered negative binomial models (ZANB) with spatial random effects were used to analyse factors influencing occurrence and density simultaneously. The results provided by the spatial ZANB models revealed that the probability of the occurrence of beech regeneration is highly dependent on seed availability, i.e. dependent on source trees in close proximity to a plot. The probability of beech regeneration rises with the increasing diameter of a potential seed tree and decreases with increasing distance to the nearest potential seed source. The occurrence of regeneration is affected by overstorey composition and competition exerted by spruce regeneration. Where sites are affected by groundwater or temporary waterlogging, the impact on the occurrence of regeneration is negative. Although distance to the nearest potential seed source has an influence on occurrence, this variable exerts no influence on density. A high regeneration density arises in conjunction with a high beech basal area in the overstorey. Beech regeneration density, but not occurrence, is negatively affected by browsing intensity. These variables can be used to predict the occurrence and density of beech regeneration in space to a high level of precision. The established statistical tool can be used for decision-making when planning forest conversion using natural regeneration.

Outcomes after an abrupt, storm-caused loss of spruce shelterwood (Picea abies L. Karst.) overstories on advanced planted beech (Fagus sylvatica L.) were investigated, including effects of developmental stage and beech stand density on growth after release. Six years after overstorey loss by storm Kyrill in January 2007, heights and root collar diameters were measured, along with annual length of the main shoot and ring widths of the stem and strongest branch. No significant difference in the total height of released and sheltered beeches was found at six years after shelter loss, but annual growth of the main shoot of released beeches increased from the second year after release. Overall growth patterns of released and sheltered beech differed significantly. Diameter growth and that of the strongest branch increased strongly after shelter loss. Height-to-diameter ratio (H/RCD) indicated that sheltered beeches with higher densities were slenderer. However, no differences were found in growth response between young stands and beech in the thicket live stage. Overall, advanced planted beech beneath a spruce shelterwood of medium canopy closure showed vigorous height growth and a qualitatively desired form. Sudden release of the planted beech enhanced diameter growth of stem and branches, which is undesirable for timber quality. Findings suggest that advanced planted beech should not be released by abruptly cutting the shelterwood. Instead, with stepwise canopy opening beech should gradually adapt to the open conditions.

1. Forest regrowth following farmland (agriculture and pasture) abandonment has been positively associated with a number of processes including the regulation of hydrological cycling, the enhancement of soil functioning and an increase in forest productivity and carbon (C) sequestration. Although these changes in eco-system functioning post-farmland abandonment have been observed in multiple locations and studies, the ecophysiological basis underpinning these patterns re-mains unclear. Here, we examine whether increased forest expansion following pastureland abandonment is associated with greater water-use efficiency (WUE) and legacies from previous land use in terms of nitrogen (N) availability., 2. We thus explored differences in leaf traits and N availability between recently es-tablished (post-1950) beech Fagus sylvatica (L.) forests on former pastureland and long-established beech forests (pre-1950). The investigated leaf traits were SLA, leaf N concentration (%N) and intrinsic WUE (iWUE, i.e. the ratio between photo-synthesis and stomatal conductance); as well, leaf and soil stable N isotope compo-sition (δ15N) and total %N were used to assess changes in N availability. Finally, we compared the correlation strength between the above-mentioned parameters and those associated with tree productivity (wood density and basal area increment, BAI) and the richness of ectomycorrhizal fungi (ECM) in these two forest types., 3. Recent forests had greater iWUE than long-established forests, which was associated more with lower SLA than leaf %N. Leaf and soil δ15N were more robust proxies than %N for detecting differences in N availability. Less negative leaf and soil δ15N values in recent versus long-established forests suggest, on the one hand, greater N avail-ability, probably due to higher historical N input originating from animal excreta on these former pasturelands, and, on the other hand, an increase in N loss pathways., 4. Our results point to greater correlations between leaf δ15N, tree iWUE and pro-ductivity in recent forests than in long-established forests, thereby suggesting a close link between C and N cycles. Our findings also highlight different N dynam-ics between the two forest types, with recent forests showing ‘leaky’ N cycling wherever lower N retention by trees and associated ECM fungi occurs as a legacy of previous land use., This study was supported by the projects: SPONFOREST (APCIN_ 2016_0174), NEWFORLAND (RTI2018-099397-B-C22 MCIU/AEI/ERDF, EU) and BEEMED (SGR913 Generalitat de Catalunya). Etal.was supported by the postdoctoral grant Juan de la Cierva-Formación from the Spanish Ministry of Economy, Industry and Competitiveness (grant no. FJCI-2015-26848). T

Trees have a strong influence on the chemical properties of the soil in which they grow. Establishing plantations with different tree species combinations thus potentially leads to divergence in soil chemistry. To study the degree to which differentiation already occurs during the first years after establishment, we made use of a biodiversity–ecosystem functioning experiment in Belgium, FORBIO. The multi-site experiment replicates tree species richness (1–4 species) and composition in three sites. The sites represent contrasting site contexts, mostly land-use history, soil, and climate. Soil samples (0–10 cm) were taken at the time of planting and approximately 8 years later. We measured the total C and N concentration, the Olsen P, the pH-H2O, and the concentration of base and Al cations. The change in chemical composition was strongly dependent on site conditions including former land use. Afforestation on former cropland had a positive impact on total C and Olsen P and a negative effect on base cations and pH. On sites reforested after clear-cut, soil texture and particular site preparation played an important role. On top of that, we found several significant effects of species composition. Strongest composition effects were detected in the reforested site on loamy soil with little soil disturbance. This study highlights that species choice can already affect soil chemistry in early stages of forest development, but that the nature of the effects may strongly depend on the context in which the plantations are established. Further research is needed to identify the most important contextual factors.

As the demand for dried timber has been increasing worldwide, drying of timbers is required to supply the demand in the manufacturing industries of furniture and other interior woodwork. During the drying process, some condensation will occur, resulting in liquids' emission, which requires disposal. in the liquid condensate, there are many substances at low concentrations that make their recovery uneconomic. After revealing the potential of the substances available in these condensates, biotechnological processes can be employed to produce high value-added products. This condensate contains tree sap, including proteins, enzymes, and many natural compounds such as terpenoids, alkaloids, phenolic compounds. Thus, it is considered that these natural compounds might be beneficial in pharmaceutical applications. Natural compounds such as phenolic compounds are bioactive substances that have antioxidant, antimicrobial, and cytotoxic activities. This study was aimed to characterize the condensate from the Rf-vacuum drying process of beech (Fagus sylvatica L.), walnut (Juglans regia L.), and restharrow (Onosis arvensis L.) in terms of antioxidant capacity, total phenol content, antimicrobial activity, and cytotoxicity. Obtained data revealed that condensates from the Rf-vacuum drying process inhibited the growth of both S. epidermidis and E. coli significantly.

Recently, many studies worldwide tapped tree ring pattern for detection of growth events and trends caused by weather extremes and climate change. As long-term experiments with permanent survey of all trees are rare, growth trend analyses are mostly based on retrospective measurements of growth via increment coring or stem analyses of the remaining individual trees in older forest stands. However, the growth of the survivor trees in older stands may only unsufficiently represent the course of growth of the dominant trees throughout the stand development. Here, the more than 100 years survey data of the European beech (Fagus sylvatica L.) thinning experiment Fabrikschleichach in South Germany are used to show the long-term changes in social ranking of trees and their consequences for growth trend detection by retrospective tree ring analyses, for stand dynamics and silvicultural management. Firstly, a significant social upwards drift of initially medium-sized trees till 2010 is shown based on the trees' percentiles in the stem diameter distribution in 1904 versus 2010. The social climbing is stronger on the thinned compared to the unthinned plots. Secondly, we show that 40–60% of the 100 tallest trees in 1904 were replaced by social climbers and down-ranked below the 100 tallest trees till 2010. Linear mixed model analyses reveal that the long-term trend of the diameter growth of the 100 dominant survivors in 2010 was on average by 23% steeper than the trend of the 100 tallest starters in 1904. This indicates that the survivors had a steeper and longer lasting growth than the originally dominant trees. Thirdly, the diameter growth trend in the last 20 years, from 1990 to 2010, is analyzed in dependency on the current and past social position. A linear model shows that early subdominance or suppression can significantly steepen the growth trend a century later and vice versa.Finally, we discuss the implications of the social drift for the survivor-based growth trend analyses, for the stand dynamics, and silvicultural management.

More frequent and severe disturbances increasingly open the forest canopy and initiate tree regeneration. Simultaneously, increasing weather extremes, such as drought and heat, are threatening species adapted to cool and moist climate. The magnitude of the microclimatic buffering capacity of forest canopies to mitigate hot and dry weather conditions and its disturbance-induced reduction remains poorly quantified. Also, the influence of disturbance legacies (e.g., deadwood) on forest microclimate is unresolved. In a unique manipulation experiment we investigated (i) the microclimatic buffering capacity of forest canopies in years with different climatic conditions; (ii) the impacts of spatial disturbance patterns on surface light and microclimate; and (iii) the effect of deadwood presence and type on microclimate. Treatments included two disturbance patterns (i.e., aggregated and distributed), four deadwood types (i.e., standing, downed, standing and downed, removed), and one untreated control (i.e., nine treatments in total), replicated at five sites dominated by European beech (Fagus sylvatica L.) in southeastern Germany. We measured forest floor light conditions and derived diurnal extremes and variation in temperature (T) and vapor pressure deficit (VPD) during four consecutive summer seasons (2016 – 2019). The buffering capacity of intact forest canopies was higher in warm and dry years. Surface light was significantly higher in spatially aggregated disturbance gaps compared to distributed disturbances of similar severity. An increase in surface light by 10 % relative to closed canopies elevated Tmax and VPDmax by 0.42°C and 0.04 kPa, respectively. Deadwood presence and type did not affect the forest microclimate significantly. Microclimatic buffering under forest canopies can dampen the effects of climate change. However, increasing canopy disturbances result in more light penetrating the canopy, reducing the microclimatic buffering capacity of forests. We conclude that forest management should foster microclimatic buffering in forests as one element of a multi-pronged strategy to counter climate change.

[Motivation] Trait variation within species can reveal plastic and/or genetic responses to environmental gradients, and may indicate where local adaptation has occurred. Here, we present a dataset of rangewide variation in leaf traits from seven of the most ecologically and economically important tree species in Europe. Sample collection and trait assessment are embedded in the GenTree project (EU-Horizon 2020), which aims at characterizing the genetic and phenotypic variability of forest tree species to optimize the management and sustainable use of forest genetic resources. Our dataset captures substantial intra- and interspecific leaf phenotypic variability, and provides valuable information for studying the relationship between ecosystem functioning and trait variability of individuals, and the response and resilience of species to environmental changes., [Main types of variable contained] We chose morphological and chemical characters linked to trade-offs between acquisition and conservation of resources and water use, namely specific leaf area, leaf size, carbon and nitrogen content and their ratio, and the isotopic signature of stable isotope 13C and 15N in leaves., [Spatial location and grain] We surveyed between 18 and 22 populations per species, 141 in total, across Europe., [Time period] Leaf sampling took place between 2016 and 2017., [Major taxa and level of measurement] We sampled at least 25 individuals in each population, 3,569 trees in total, and measured traits in 35,755 leaves from seven European tree species, i.e. the conifers Picea abies, Pinus pinaster and Pinus sylvestris, and the broadleaves Betula pendula, Fagus sylvatica, Populus nigra and Quercus petraea., [Software format] The data files are in ASCII text, tab delimited, not compressed., All authors acknowledge support from the European Union’s Horizon 2020 Research and Innovation Programme under grant agreement no. 676876 (Project GenTree). Laboratory work was also supported by other projects: COMEDIAS (CGL2017- 83170-R, Spanish Ministry of Science, Innovation and Universities), REMEDINAL IV (TE-CM S2018/EMT-4338, Project of the Autonomous Community of Madrid, Spain), SPONFOREST (BIODIVERSA-3, PCIN-2016-055), and International Laboratory of Global Change (LINCGlobal 4540-143AP). The Swiss contribution was supported by the Swiss Secretariat for Education, Research and Innovation (SERI) under contract no. 6.0032. BC was funded by a Scholarship from Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES (Brazil).

Is it possible to untangle the historical mosaic of human interference with the origin of forest plantations in Europe? This topic is relevant at the northern frontier of species expansion such as western Lithuania, where European beech was introduced by German foresters in the eighteenth century. Our aim was to identify the origin and assess the genetic diversity of the introduced European beech populations in Lithuania with the aid of DNA markers. We used 10 nuclear microsatellite loci to genotype 1260 European beech trees from the introduced populations in Lithuania and 18 natural reference populations from Germany, Sweden and Poland as the most likely sources of the introduction. We calculated common genetic diversity indexes and used several clustering approaches to investigate the genetic associations. There was a significant differentiation among the populations at all the loci (Dest = 0.07 to 0.24). The Bayesian clustering returned high likelihood for the Lithuanian European beech stands to originate from three distinct sources: (a) the Bavarian Alps, (b) north-eastern Poland, and (c) south-eastern Poland and the Carpathian Mountains. Despite the high allelic diversity, the Lithuanian European beech populations possessed a markedly lower observed heterozygosity than the natural populations. We assume that the seeds for the Lithuanian stands were collected over a geographically wide range (resulting in high allelic diversity), but at each spot, a few adjacent and likely related trees were sampled. A DNA-based tracking system could efficiently reveal historic transfers of forest reproduce material Europe and help to improve the adaptability of future forests.

The mass loss kinetics of thermally modified wood species was analyzed as a time–temperature function. European beech (Fagus sylvatica L.), English oak (Quercus robur L.), Norwegian spruce (Picea abies L. Karst.) and Scots pine (Pinus sylvestris L.) wood specimens of dimensions 20 × 20 × 10 mm3 were thermally modified at 140 °C, 160 °C, 180 °C, 200 °C and 220 °C for 1–6 h using atmospheric pressure and superheated steam environment. The process intensity was determined by mass loss (ML), based on oven-dry mass before and after the thermal modification. Furthermore, the equilibrium moisture content (EMC) was determined before and after thermal modification to analyze the effect of mass loss on the sorption properties. Measured mass loss data were compared with the three-dimensional analytical function and its applicability to mass loss prediction was verified. For the studied wood species, the ML was found to be less than 1–1.5% when temperature of 140 °C and 160 °C was applied. Differences between studied species were more significant at temperatures higher than 160 °C. With the highest tested temperature (220 °C), mass loss reached 13.5% (beech), 18.8% (oak), 6.7% (spruce) and 13.5% (pine). According to the results, hardwoods have been shown to be more sensitive to the thermal degradation than softwoods as demonstrated by the higher mass loss recorded for the same modification time and temperature. The three-dimensional analytical function was confirmed as valid for all the species studied (R2 = 0.89–0.99) and relevant for the mass loss prediction using fitted parameters. The EMC was reduced after thermal modification within the range of 4–48%, 0.4–47%, 1–32% and 0.7–40% for beech, oak, spruce and pine, respectively. Further, the EMC correlates exponentially (R2 = 0.91–0.95) with the decrease in the specimens’ mass depending on the wood species used and modification temperature applied. However, the EMC seems to be almost stabilized beyond a limit value of approximately 10–12% of mass loss. The results provide a better insight into the mass loss and EMC kinetics of thermally modified wood species and can be used as a tool for prediction of mass loss values and required material properties (EMC) for designed wooden products.

Temperate forest ecosystems are exposed to a higher frequency, duration and severity of drought. To promote forest longevity in a changing climate, we require a better understanding of the long-term impacts of repetitive drought events on fine-root dynamics in mature forests. Using minirhizotron methods, we investigated the effect of seasonal drought on fine-root dynamics in single-species and mixed-species arrangements of Fagus sylvatica (European beech) and Picea abies (Norway spruce) by means of a 4-yr-long throughfall-exclusion experiment. Fine-root production of both species decreased under drought. However, this reduction was not evident for P. abies when grown intermixed with F. sylvatica. Throughfall-exclusion prolonged the lifespan of P. abies roots but did not change the lifespan of F. sylvatica roots, except in 2016. Fagus sylvatica responded to drought by reducing fine-root production at specific depths and during roof closure. This is the first study to examine long-term trends in mature forest fine-root dynamics under repetitive drought events. Species-specific fine-root responses to drought have implications for the rate and depth of root-derived organic matter supply to soil. From a root dynamics perspective, intermixing tree species is not beneficial to all species but dampens drought impacts on the belowground productivity of P. abies.

Fagus sylvatica (beech) dominates the montane forests of the Apennines and builds old-growth high-conservation value stands. However, recent severe drought-induced diebacks raise concern on the future persistence of these forests and of Southern European mesophilous woodlands overall, growing at their dry edge. To explore the history of Apennine beech-dominated forests, we draw on the multiproxy paleoecological record from Lago Verdarolo, which includes a robust vegetation-independent temperature reconstruction. Numerical techniques are used to investigate the drivers of long-term Mediterranean mountain forest dynamics. Specifically, we focus on disentangling the ecological factors that caused the shift from high-diversity mixed forests to beech-dominated stands and on assessing the occurrence of legacy effects on present-day forests. Abrupt climate change largely drove vegetation dynamics during the Late Glacial and Early Holocene. Species-rich mixed Abies alba (silver fir) forests dominated about 10,500—5500 years ago, under rather dry and warmer-than-today conditions (+ 1—2 °C) and limited fire occurrence. Cooler and moister summers and increasing fire activity caused declines in several fire-sensitive temperate deciduous trees (for example, Ulmus, Tilia, Fraxinus) and favored the establishment of fir-beech forests around 5500 years ago. Further enhancement of fire activity and farming around 2000 years ago led to local Abies alba extinction and forest impoverishment. We conclude that the currently widespread monospecific Apennine beech forests are the result of multi-millennial land-use intensification superimposed on Late Holocene cooling and moistening. Given their higher drought-tolerance compared to beech stands, reviving ancient species-rich mixed fir forests represents a feasible and ‘tested’ possibility to adapt forests to climate change. Electronic supplementary material The online version of this article (10.1007/s10021-020-00587-4) contains supplementary material, which is available to authorized users.

Treating wood with water-soluble resins is a well-known and effective method to improve the durability of wood. However, there has been no systematic work to date related to the influence of average molecular size of phenol-formaldehyde (PF) resin on the decay resistance of wood, especially of hardwoods. Therefore, the goal of this study was to investigate the effect of average molecular size of PF resin treatment on the resistance of beech wood against brown- and white-rot fungi. Four different average molecular weights (M w) of resol type resin oligomers (297, 421, 655 and 854 g/mol) were examined. Different weight percent gains (WPGs) in European beech (Fagus sylvatica) wood blocks (15 × 20 × 50 mm3) were attained through vacuum impregnation using various concentrations of aqueous-PF solutions. Afterwards treated wood blocks passed the leaching and were exposed to brown-rot fungi (Gloeophyllum trabeum; Coniophora puteana) and white-rot fungi (Trametes versicolor) for 16 weeks. No effect of oligomer size on the resistance against G. trabeum decay of wood blocks was observed, resulting in resin loadings of 7–8%. The required WPG for resistance to brown-rot decay by C. puteana increased slightly with increasing oligomer molecular size: 6, 7, 10 and 11% for wood treated with 297, 421, 655 and 854 g/mol, respectively. The extent of white-rot fungal decay resistance of treated wood was affected by the molecular size of oligomers. Resin loadings of 8% and of 17% against T. versicolor were required to attain similar durability levels for beech wood treated with M w = 297 and 854 g/mol, respectively.

In this study, we analysed patterns of spatial variation in the basal area of live and dead trees and structural complexity in close-to-primeval forests in the Dinaric Mts. The results were compared with an analogous study conducted in the Western Carpathians. The research was carried out in the Janj, Lom and Perucića forest reserves (Bosnia and Herzegovina) in mixed-species stands of silver fir Abies alba Mill., European beech Fagus sylvatica L. and Norway spruce Picea abies (L.) H. Karst. In the core zones of the reserves, concentric sample plots (154 and 708 m2) were set in a regular 20 × 20 m grid covering approximately 10 ha. The analyses revealed varying distribution patterns of live canopy trees, suggesting that these characteristics may fluctuate to some extent at the regional level. At the spatial scale of 708 m2, attractive associations between dead canopy trees were found, but this tendency disappeared with increasing area. Although stands in the Dinaric Mts. are characterized by an almost twofold greater biomass accumulation compared to those from the Western Carpathians, the study revealed analogous bell-shaped distributions of stand basal areas of live trees and a very similar trend of decreasing variation in stand basal area and structural heterogeneity with increasing spatial scale. Nonetheless, the higher growing stocks, lower ratios of dead to live tree basal area and lower proportion of homogeneous structure types found in the Dinaric Mts. may suggest a less severe disturbance history over recent decades in this region compared to the Western Carpathians.

The aim of the present study was to generate a consistent data set for biorefinery intermediates from novel energy crop Silphium perfoliatum compared with Fagus sylvatica, including process streams from liquid hot water pretreatment, enzymatic hydrolysis, and fermentation with Bacillus licheniformis. The consistency of the data set was further supported by the application of a consistent analytical method based on high performance anion exchange chromatography with pulsed amperometric detection and validated for process intermediates, which renders the technique a versatile analytical tool also for alcoholic compounds. For the first time, Silphium perfoliatum was used for liquid hot water pretreatment which resulted in a maximal absolute glucose recovery after enzymatic hydrolysis for feedstock pretreated with 200 °C, 20 min, 20% solid loading. Under these conditions, 68% glucose were recovered for Silphium perfoliatum and 62% for Fagus sylvatica. Further, enzymatic hydrolyzates of both feedstocks were successfully used as single carbon sources for 2,3-butanediol fermentation with Bacillus licheniformis resulting in a 2,3-butanediol yield of 39% and 31% of the theoretical yield for Silphium perfoliatum and Fagus sylvatica, respectively. Thus, the technical suitability of Silphium perfoliatum as feedstock for a liquid hot water–based biorefinery process was comprehensively demonstrated and successfully bore the comparison with Fagus sylvatica.

Soils play a significant role in climate regulation, especially due to soil organic carbon (SOC). The SOC pool is therefore modeled for various environments, and forest floor and topsoil thicknesses are important parameters for most of these models as they store most of the SOC. However, the forest floor and topsoil thicknesses show high spatial variability which is a result of multiple factors which are not agreed upon among scientists. Out of these factors, we choose topography parameters (elevation, slope, and topography wetness index) and forest stand characteristics (stand age, dominant tree species, and forest floor cover), and soil moisture, and we analyzed their relationship to the forest floor and topsoil thicknesses. The study was performed in a managed submontaneous forest in Central Europe dominated by Picea abies (L.) Karsten with small patches of Fagus sylvatica L. or other species. The thicknesses of the O horizons (Oi, Oe, Oa) and topsoil were measured at 221 sampling pits. Geographically weighted regression showed that the spatial variability of the overall forest floor plus topsoil thickness (OA) is responsible for 8% of its variability. The thickness of the OA is the most strongly controlled by forest floor cover explaining approximately 6% of its variability and soil moisture explaining 2–6% of the variability. The Oi + Oe horizon thickness is controlled only by forest floor cover explaining 10.7% of its variability, and the thickness of Oa + A horizon can be explained mainly by soil moisture in mineral horizon explaining 9% of the variability.

Tree species differ in their functioning at the scale of an individual tree which will result in differences in ecosystem service provision. Replacement trees for diseased trees should take account of functional differences. Globally tree species composition is changing due to species loss from pests and pathogens. The impact of this change on ecological functioning is rarely tested. Using six sites across the UK, with multiple tree species at each site, we test for functional differences between three species threatened by disease in the UK: Quercus petraea, Q. robur and Fraxinus excelsior and six other species: Acer pseudoplatanus, Castanea sativa, Fagus sylvatica, Quercus cerris, Quercus rubra, and Tilia x europaea, which have previously been suggested as ecological replacements. Differences between species were detected for all the variables measured: nitrogen mineralization, decomposition rate, total soil carbon and nitrogen, soil pH, soil temperature, and bark water holding capacity. Non-native Quercus species were only suitable replacements for native Quercus for some of the functions measured but replicating native Quercus functioning using a mixture of other species may be possible. The functioning of F. excelsior was different from most other tree species, suggesting that replicating its functioning with replacement tree species is difficult. The work highlighted that which species replaces diseased trees, even at the scale of single trees, will impact on the functions and ecosystem services provided.

In this study, the relatively new picture frame method applied to wood is compared with three established shear test methods, namely the experimental modal analysis, the square plate twist method and the torsion test. For the investigations, the wood species European beech (Fagus sylvatica L.) and spruce (Picea abies L. Karst.) were used and the shear tests were conducted in LR and RL direction. The results show comparable shear moduli for beech and spruce in the range of 931–1289 Nmm−2 and 495–842 Nmm−2, respectively. In contrast to the theory of linear elastic orthotropic materials, significant differences in the results of the picture frame method between LR and RL direction were observed for spruce.

Juveniles and canopy trees may not exhibit similar nitrogen acquisition responses to soil temperature change caused by variation in snow cover over winter. The use of 15N tracer is a powerful tool for tracking the effects of variation in soil frost on plant nitrogen acquisition. While the responses of juvenile trees to environmental change are often used to infer the responses of canopy trees, the 15N enrichment responses of juveniles and mature canopy trees may not be comparable. We conducted a winter soil temperature manipulation study (snow exclusion, ambient snow or soil insulation) in a lowland Fagus sylvatica forest. 15N tracer was applied the following spring and the 15N enrichments of soil, juvenile and mature canopy trees were examined in late fall. Within canopy trees and juveniles, the relative treatment effects on 15N enrichment were consistent among all sampled tissues (roots, stem cores, leaves, buds and the current year’s shoot growth). For juveniles, 15N enrichment was highest under snow exclusion (coldest soil) and lowest under soil insulation (warmest soil), and lower 15N enrichment occurred under ambient conditions than under snow exclusion. For canopy trees, 15N enrichment also was highest under snow exclusion and lowest under soil insulation, but there was no difference in 15N enrichment between ambient conditions and the snow exclusion treatment. Therefore, our results indicate that sampling of juveniles may overestimate the nitrogen acquisition responses of mature trees to winter temperature variation.

This paper presents the results of extensive investigations on the lamination strength grading, the production and the mechanical properties of European beech (Fagus sylvatica L.) glued laminated timber (GLT). Based on the analysis of potential influencing parameters on strength and stiffness as well as subsequent tension tests parallel to the grain on single boards, a combined visual/machine approach for grading the raw material into tensile strength classes T50, T42, T33 and T22 was developed. Boards strength graded with the developed procedure were then finger-jointed by a Swiss GLT producer and the strength of the finger joints was investigated by means of tension and bending tests. The strength and durability of the bonding was investigated and verified by means of tensile-shear and delamination tests. It could be shown that the required finger-joint and bondline strengths for GLT of strength classes GL40 and GL48 can be achieved, but that the process parameters for finger jointing (in particular the geometrical properties of the finger joint profile) have to be optimized in order to be able to produce GLT of strength class GL55. Finally, an extensive experimental testing campaign was performed to investigate the mechanical properties of European beech GLT produced based on the strength grading rules and production techniques developed before. Bending, tensile and compressive parallel to the grain, as well as shear tests were carried out on GLT specimens of strength classes GL40, GL48 and GL55 in different sizes in terms of cross-section and length. Based on these investigations and complementing numerical simulations, characteristic strength and stiffness values and formulae for consideration of size effects in bending, tension and shear were determined.

The introduction of non-native species with various ecological and functional traits to European forests may be a potential tool for mitigating climate risks. We analyzed the growth sensitivity to climate of seven alien (Acer rubrum, Betula maximowicziana, Castanea sativa, Cryptomeria japonica, Metasequoia glyptostroboides, Thuja plicata and Tsuga heterophylla) and two native (Fagus sylvatica and Picea abies) tree species on a productive forest site in western Germany in order to assess their potential suitability in light of climate change. Retrospective basal area increment at breast height (BAI) and total stem volume increment (VI) were statistically related to a total of 11 multiscale climatic variables showing minor discrepancies between BAI and VI in climatic sensitivity. Across the tree species, the strongest climate signals were found for the standardized precipitation evapotranspiration index (spei), vapor pressure deficit (vpd) and the number of frost days (nfr) representing water availability, heat and coldness, respectively. Growth of most species was positively affected by mild winters and growing season spei. Importance of the investigated variables differed among tree species with, e.g., C. sativa and T. plicata being mainly driven by winter vpd and A. rubrum, C. japonica and P. abies by water availability in the growing seasons. We conclude that particularly more anisohydric species (e.g., C. sativa) may profit from their low sensitivity to drier conditions. Highly sensitive and isohydric species (e.g., A. rubrum) will undergo significant growth reductions when climate becomes more variable.

UK trees require increased conservation efforts due to sparse and fragmented populations. Ex situ conservation, including seed banking, can be used to better manage these issues. We conducted accelerated ageing tests on seeds of 22 UK native woody species, in order to assess their likely longevity and optimize their conservation in a seed bank. Germination at four ageing time points was determined to construct survival curves, and it was concluded that multiple samples within a species showed comparable responses for most species tested, except for Fraxinus excelsior. Of all species studied, one could be classified as very short-lived, four as short-lived and 17 as medium, with none exceeding the medium category. The most important finding of this manuscript is that although some taxonomic trends were observed, the results indicate the need for caution when making broad conclusions on potential seed storage life at a species, genus or family level. Longevity predictions were compared to actual performance of older collections held in long-term storage at the Millennium Seed Bank, Kew. Although most collections remain high in viability in storage after more than 20 years, for short-lived species at least, there is some indication that accelerated ageing predicts longevity in seed bank conditions. For species with reduced potential longevity, such as Fagus sylvatica and Ulmus glabra, additional storage options are recommended for long-term gene banking.

As currently framed, the forest cycle model that underlies close-to-nature management in temperate beech forests throughout the globe specifies an orderly sequence of temporal development within even-aged patches comprising the forest mosaic. Although this model has been widely applied to European beech (Fagus sylvatica L.) forests, the underlying assumptions of disturbance-induced even-agedness (i.e., within-patch age homogeneity) and competition-induced size differentiation (i.e., within-patch size heterogeneity) have not been tested in natural beech forests due to prohibitions on tree coring in primeval forest reserves. In a rare and unprecedented test dataset of spatially explicit tree ages in an old-growth European beech forest, we employed triangulated irregular networks of Delaunay triangles to objectively identify natural tree neighborhoods to determine if neighboring (i.e., within-patch) trees were even- or, at most, two-aged. Age differences among neighboring trees (summarized in 25-yr age classes) were rarely 50 yrs, while the few "even-aged" patches were very small (100 m2) and relatively young (

European beech is a temperate species with peripheral populations in the Mediterranean mountains. These Mediterranean beech forests have often been interpreted as climatic relicts. However, the use of this term does not always seem appropriate because their recent post-glacial expansion in the Iberian Peninsula makes it difficult to disentangle the effects of climate from those associated with land use. Moreover, the observed recovery of Mediterranean beech forests after land-use abandonment is not what might be expected of a climatic relict. Our objective is to clarify this issue by studying one of these Mediterranean “relict” populations located at the south-western margin of this species (Iberian Central Range), discriminating between subpopulations with dissimilar land-use legacies. Differences between the climatic range of this population and the rest of the European beech forests were explored. Historic and recent shifts in the species distribution area were studied by reviewing paleorecords from the literature and contrasting historical and recent cartography. Field data on population structure and demography were also examined. The climatic analyses do not point to climatic marginality of the peripheral population studied. Furthermore, the results revealed no evidence of reduction in area or confinement over recent decades. Contrary to what is expected for climatic relict species under current climate, beech is expanding in the studied mountain following the abandonment of traditional land uses, particularly since the middle of the twentieth century CE. Our results do not support the concept of a climatic relict but rather, they point to the importance of anthropogenic legacies to explain the distribution area, population dynamics and current conservation status of Mediterranean beech forests., This work was funded by the Spanish Ministry of Science, Innovation and University through Projects: AGL2013-46028-R (SCALyFOR), AGL2016-76769-C2-1-R (FORESTCHANGE) and AGL2014-61175-(MEDFORMEC). Acknowledgements

Beech (Fagus sylvatica) and silver fir (Abies alba) are often cultivated in mixed stands and, hence, compete for water and nutrients. Besides nitrogen (N), also phosphorus (P) is an important nutrient for growth and development. Beech trees in Central Europe grow on both P-poor and P-rich soils, thereby showing similar growth and low variation in foliar P. The central aim of the present study was to test the hypothesis that variations in foliar P contents of beech are driven by seasonal changes rather than by the competition with silver fir. It was further hypothesized that P contents in silver fir needles depend on needle age and forest site. To test these hypotheses, P contents and P fractions, i.e. organic-bound P (Porg) and inorganic phosphate P (Pi), were measured in the foliage of beech trees from pure beech and mixed beech/silver fir plots as well as in needles of silver fir of the mixed plots. The forest sites investigated are located in Central Europe in the Black Forest, Germany, and in Croatia near the south-eastern distribution limit of beech and are all poor in plant-available soil P. The analyses showed that the main driver of P contents and P fractions in beech leaves at all forest sites is the season and that competition with silver fir had no effect. Hence, the present results demonstrate the high plasticity of beech trees to adapt to both poor plant-available soil P and competition with silver fir. Total P contents of silver fir needles were higher at the Croatian site compared to the Black Forest sites and originated from higher foliar Pi contents. One third of the P present in current-year needles in late summer was remobilized and exported until the needles reached the age of 1 year. The difference in P contents between current-year and 1-year-old needles can be seen as the amount of P resorbed from 1-year-old needles in summer during the generation of new needles to support the P demand of current-year needles for growth and development.

Cultivation of seedlings in trays requires the use of specifically developed substrates. This study presents the results of the analyses of selected physical and mechanical parameters of a peat–perlite substrate, in which seedlings of the Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies (L.) H. Karst), European beech (Fagus sylvatica L.), and pedunculate oak (Quercus robur L.) were grown during a production season. For each species, the substrate parameters changed throughout the production season, substrate dry weight decreased, whereas its compactness increased with time. Independent of the species, the bulk density and total porosity of the substrate changed or deviated from the optimum range, although the values of porosity were close to maximum and those of density were close to the minimum. In certain periods in the V265 trays with beech and oak seedlings, the substrate was characterized by very low water capacity and excessive air capacity. Compactness measured with a cone penetrometer showed, that this parameter might be used for monitoring the substrate properties.

Foliar water uptake (FWU), the direct uptake of water into leaves, is a global phenomenon, having been observed in an increasing number of plant species. Despite the growing recognition of its functional relevance, our understanding of how FWU occurs and which foliar surface structures are implicated, is limited. In the present study, fluorescent and ionic tracers, as well as microcomputed tomography, were used to assess potential pathways for water entry in leaves of beech, a widely distributed tree species from European temperate regions. Although none of the tracers entered the leaf through the stomatal pores, small amounts of silver precipitation were observed in some epidermal cells, indicating moderate cuticular uptake. Trichomes, however, were shown to absorb and redistribute considerable amounts of ionic and fluorescent tracers. Moreover, microcomputed tomography indicated that 72% of empty trichomes refilled during leaf surface wetting and microscopic investigations revealed that trichomes do not have a cuticle but are covered with a pectin-rich cell wall layer. Taken together, our findings demonstrate that foliar trichomes, which exhibit strong hygroscopic properties as a result of their structural and chemical design, constitute a major FWU pathway in beech.

The performance of adhesive-hardwood bonds can often be sensitive to humidity and temperature variation. Therefore, it is frequently challenging to achieve standard requirements for structural applications. To gain a better understanding of the wood-adhesive bond, the properties of the individual constituents as well as the local interface of European beech (Fagus sylvatica L.) wood cell walls in contact with structural adhesives were analyzed by means of nanoindentation. These results are compared to classical lap-shear strength. As adhesives two different one-component polyurethane adhesives (1C PUR) and a phenol resorcinol formaldehyde adhesive (PRF) were used. In one case, the beech wood was additionally pre-treated with an adhesion-promoting agent (primer) prior to bonding with 1C PUR. Beech wood joints were analyzed subsequent to several treatments, namely standard climate, after wet storage and in re-dried conditions. In addition, the influence of the primer on the hydroxyl accessibility of beech wood was investigated with dynamic vapor sorption (DVS). The lap-shear strength revealed good performance in dry and re-dried conditions for all adhesives on beech. Both polyurethane adhesives obtained deficits when tested in wet conditions. The use of a primer significantly improved the PUR performance in wet condition. DVS experiment demonstrated a decrease in hydroxyl group accessibility when using a high primer concentration. As novelty, nanoindentation was used for the first time to characterize the local wood–adhesive-interface properties in wet conditions. Nanoindentation showed that all tested 1C PUR perform quite similar in room climate, while PRF achieves considerable higher values for reduced E-modulus and hardness. Wet storage led to a considerable reduction in mechanical properties for all adhesives, while the highest relative change was observed for PRF. After re-drying, the adhesives re-gained a large part of their original mechanical properties in room climate. No distinct effect of the primer on the local micromechanical properties could be detected with nanoindentation in terms of specific work of indentation.

Common gardens are experimental plantations for comparing the performance of tree species while eliminating many of the variables that prevail in natural tree stands. The aim of this study was to evaluate the biodiversity of corticolous lichens on Danish tree species (Acer pseudoplatanus, Alnus glutinosa, Betula pendula, Fagus sylvatica, Fraxinus excelsior, Quercus robur and Tilia cordata) under common garden conditions and to examine the height distribution of particular lichen species. Observations were recorded through regular sampling of at least 36 lichen species on the main stems (from the base of the stem to the treetops) of 44-year-old trees at four common garden sites. Acer pseudoplatanus and Fraxinus excelsior had the greatest lichen species richness and Shannon diversity values while these measures were significantly lower for Betula pendula and Fagus sylvatica. The distribution of lichen species appeared biased among tree species. The general lichen distribution and relative sample height were weakly related (nonmetric multidimensional scaling). However, single lichen species showed a clear differential distribution along the tree stem (P Lepraria incana, Pseudosagedia aenea and Arthonia atra were mainly found at the stem base while Lecanora carpinea, L. chlarotera, Lecidella elaeochroma, Physcia tenella and Xanthoria parietina, were most abundant at around 70% of the total tree height. The differential distribution of single lichen species presumably reflects different specific requirements during spore germination and thallus growth. By isolating the unique effect of key variables (tree species and height), this study contributes to the knowledge base of corticolous lichen ecology.

Forest ecosystems nowadays provide multiple ecosystem goods and services at a time and throughout all development phases. Species mixing is considered an effective measure to gain benefits beyond purely additive effects. However, the complex structural processes behind interspecific competition and temporal and spatial facilitative effects through mixing are still far from being understood and predictable. In particular the mixture of European beech (Fagus sylvatica L.) and sessile oak (Quercus petraea [Matt.] Liebl.) is gaining even more importance due to the fact that forests from these species are considered more tolerant to climatic effects and are expected to expand their natural range to the north due to global warming. The 30 long-term experimental plots analysed in this study reveal the structural processes in mixed beech-oak stands based on data at the tree and stand level. Using spatial and non-spatial structural indices, we can show an increasing dominance of beech over oak in unmanaged stands and the effectiveness of thinning operations to support oak. Those processes are representative for other light-demanding tree species in mixtures with shade-tolerant species. Improving the knowledge on the structural processes in mixed-species stands is particularly relevant when trying to modify forest structure in order to adapt forest management to shifting environmental conditions and the increasing demand for ecosystem services.

Promoting tree species diversity is commonly advocated in the anticipation of predicted increases in drought frequency and severity. However, mixing effects on drought responses vary substantially with site conditions and species identity. We combined annually resolved tree-ring data and repeated forest inventory data spanning the last 90 years to examine the effect of species-specific neighbourhood competition on the drought response (resistance, recovery and resilience) of European beech (Fagus sylvatica), silver fir (Abies alba) and Norway spruce (Picea abies) for six drought events that occurred since the 1970s at three sites in Switzerland. We found predominantly weak neighbourhood competition and tree species diversity effects, with significant interspecific influences only for resistance and recovery of beech. These minor neighbourhood effects were outweighed by tree age and size effects. Although age effects depended on species identity and components of resilience, tree size consistently negatively affected all species. Our results emphasize that diversity effects may vary for each given species combination which makes broader conclusions challenging. This is because species interact through their specific set of traits and interactions vary in space and time. Adaptive management strategies are likely to be more effective when they promote more drought-tolerant species and reductions in stand density. Despite the absence of an unequivocal advantage of tree diversity on drought resilience, striving towards species-rich forests nonetheless allows for a risk spreading among multiple species and the reinforced provision of numerous ecosystem services.

Many theoretical models predict when genetic evolution and phenotypic plasticity allow adaptation to changing environmental conditions. These models generally assume stabilizing selection around some optimal phenotype. We however often ignore how optimal phenotypes change with the environment, which limit our understanding of the adaptive value of phenotypic plasticity. Here, we propose an approach based on our knowledge of the causal relationships between climate, adaptive traits, and fitness to further these questions. This approach relies on a sensitivity analysis of the process-based model Phenofit, which mathematically formalizes these causal relationships, to predict fitness landscapes and optimal budburst dates along elevation gradients in three major European tree species. Variation in the overall shape of the fitness landscape and resulting directional selection gradients were found to be mainly driven by temperature variation. The optimal budburst date was delayed with elevation, while the range of dates allowing high fitness narrowed and the maximal fitness at the optimum decreased. We also found that the plasticity of the budburst date should allow tracking the spatial variation in the optimal date, but with variable mismatch depending on the species, ranging from negligible mismatch in fir, moderate in beech, to large in oak. Phenotypic plasticity would therefore be more adaptive in fir and beech than in oak. In all species, we predicted stronger directional selection for earlier budburst date at higher elevation. The weak selection on budburst date in fir should result in the evolution of negligible genetic divergence, while beech and oak would evolve counter-gradient variation, where genetic and environmental effects are in opposite directions. Our study suggests that theoretical models should consider how whole fitness landscapes change with the environment. The approach introduced here has the potential to be developed for other traits and species to explore how populations will adapt to climate change.

Beech wood, due to its properties, is one of the most versatile and successfully used construction materials. The wood properties could even be improved, and among different wood modification processes, the thermal modification approach is usually considered as an environment-friendly technology based only on the heat and water application during wood treatment. Changes in material properties resulting from the thermal treatment of wood increase applicability of this material, but on the other hand, detailed knowledge of the modified properties is definitely necessary for the proper application of such materials to construction engineering. Unfortunately, credible data on thermal characteristics of thermally modified wood are usually provided in a very limited way, and there is no information on specific heat in particular. An original calorimetric method was used to determine the specific heat of untreated and thermally modified European beech wood (Fagus sylvatica L.). The inverse modeling was implemented to estimate the anisotropic thermal conductivity, and significant differences were found for the radial and tangential directions. The thermal modification highly influenced the increase in the thermal conductivity in the longitudinal direction. The validation procedure showed credibility of the applied methods, and it is clear that modeling of heat transfer in thermally modified wood leads to erroneous results when using thermal properties determined merely for untreated wood.

AIM: Trees need to avoid frost damage to their young leaves by leafing out after the occurrence of the last frost, yet they also need to start photosynthesis early in the season to achieve sufficient growth. This trade‐off leads to the hypothesis that ‘safety margins’ against spring frost should become shorter, the longer the winter duration, perhaps reaching an asymptotic limit where frost damage would occur in most years. Physiologically, shorter safety margins in high‐latitude ecotypes might be achieved by lower degree‐day requirements for leaf‐out, compared to low‐latitude ecotypes. LOCATION: Europe. TIME PERIOD: 1902–2009. MAJOR TAXA STUDIED: Temperate trees. METHODS: Using herbarium collections of Acer platanoides, Carpinus betulus, Fagus sylvatica and Prunus spinosa made over 108 years at 40° to 60° N latitude, we related historic leaf‐out dates to winter and spring temperatures (chilling and degree‐days), winter duration, and date of last frost occurrence in the relevant years and locations. RESULTS: In all species, frost safety margins decreased towards high‐latitude regions with long winters, with each day increase in winter duration reducing frost safety margins by 0.48 days in Fagus and 0.32–0.21 days in Prunus, Acer and Carpinus. These latitudinal differences correlate with northern ecotypes’ shorter degree‐day requirements for leaf‐out. MAIN CONCLUSIONS: The decline in spring frost safety margins in regions with long winters supports the new hypothesis that species may reach their geographic range limit where they ‘bump up’ against experiencing regular frost injury to their young leaves. Larger datasets are necessary to further corroborate our hypothesis and future efforts should thus be directed toward increasing the latitudinal range of existing phenological databases.

Fagus sylvatica in Europe is expected to be severely affected by the ongoing climate change. In this article, seed adaptive traits, in terms of morphology and germination, of F. sylvatica populations of different postglacial lineage and intrapopulation genetic diversity were evaluated. Eight plots from two geographical provenances, Evros and Drama, were selected. Provenance shaped both morphology and germination patterns, but the effect was more pronounced on germination. Seeds from Drama were larger and heavier than those from Evros but exhibited a higher degree of dormancy and slower germination. High among-plots variability on morphology and germination was also observed, especially in Evros. This higher variability was consistent with the higher level of genetic diversity observed at genomic and chloroplast DNA markers at small or larger spatial scales from previous published studies on the same plots. Results suggested the existence of different seed adaptation strategies, mainly between provenances, as a result of possible adaptation to different environmental conditions, whereas a possible influence of a generally complex pattern of admixture between different beech subspecies and postglacial lineages could not be excluded.

This paper explores the effects of three types of admixture treatments: (1) monospecific (M: European beech surrounded by six European beech seedlings), (2) single-admixture (S: European beech surrounded by six Norway spruce seedlings), and (3) group-admixture (G: European beech surrounded by three Norway spruce and three European beech seedlings), and two different CO2 concentrations ([CO2]): ambient [AMBCO2, 385 μmol (CO2) mol−1] and elevated [ELECO2, 770 μmol (CO2) mol−1] on the stem diameter increment (DI), based on a 7-year monitoring of the diameter growth of European beech (Fagus sylvatica L.) seedlings. Furthermore, to improve our understanding of the influence of climatic variables, we assessed the effects of precipitation and temperature on radial growth. After 7 years’ experience (irrespective of admixture treatment), our data were not able to confirm the effect of elevated CO2 on the diameter growth (p value = 0.35), but the result was changed when the effect of elevated CO2 was considered under the different types of admixture. Elevated CO2 along with single-admixture increased the diameter increment significantly (p value = 0.001). In contrast, we characterized a significant effect of AMBCO2 × G interaction on DI (p value = 0.002). But DI was unaffected by elevated CO2 in M (p value = 0.77). In both [CO2], inter-specific competition with spruce resulted in the highest radial growth. The inter-specific interaction was most pronounced in the lower proportion with spruce (S) in ELECO2, while in ambient CO2, the higher proportion with spruce (G) had the greatest effect on the radial growth. With respect to the climatic factors, our observations showed that temperature had a stronger influence than precipitation in ELECO2, but in AMBCO2, precipitation had the strongest effect on the radial growth as indicated by the statistically significant correlation between climatic variables and radial growth. Overall, our result suggested that seedlings growth was more under the control of climate and edaphic factors than dependent on tree species diversity and CO2 concentrations.

Process-based models are increasingly applied for simulating long-term forest developments in order to capture climate change impacts and to investigate suitable management responses. Regarding dimensional development, however, allometric relations such as the height/diameter ratio, branch and coarse root fractions or the dependency of crown dimension on stem diameter often do not account for environmental influences. While this may be appropriate for even-aged, monospecific forests, serious biases can be expected if stand density or forest structure changes rapidly. Such events occur in particular when forests experience disturbances such as intensive thinning or during early development stages of planted or naturally regenerated trees. We therefore suggest a calculation of allometric relationships that depends primarily on neighbourhood competition. Respective equations have been implemented into a physiology-based ecosystem model that considers asymmetric competition by explicit simulation of resource acquisition and depletion per canopy layer. The new implementation has been tested at two sites in Germany where beech (Fagus sylvatica) saplings have either been planted below a shelterwood of old spruces (Picea abies) or grown under clear-cut conditions. We show that the modified model is able to realistically describe tree development in response to stand density changes and is able to represent regeneration growth beneath a gradually decreasing overstorey of mature trees. In particular, the model could represent the faster crown size development in saplings until full ground coverage is established and a faster height growth afterwards. The effect enhances leaf area and thus assimilation per tree and increases carbon availability for stem growth at early development stages. Finally, the necessity to consider dynamic allometric relations with respect to climate change impacts is discussed, and further improvements are suggested.

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 15N 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.

Both climate and land-use changes, including the introduction and spread of allochthonous species, are forecast to affect forest ecosystems. Accordingly, forests will be affected in terms of species composition as well as their soil chemical and biological characteristics. The possible changes in both tree cover and soil system might impact the amount of carbon that is stored in living plants and dead biomass and within the soil itself. Additionally, such alterations can have a strong impact on the detrital food web that is linked to litter decomposition. Although there are studies on the influence of plant diversity on soil physical and chemical characteristics, the effects on soil biological activity and carbon storage processes remain largely unknown. The aim of this study was to investigate and compare chemical and biological variables in covariation with plant communities in an autochthonous beech forest (Fagus sylvatica L.) and a black pine plantation (Pinus nigra J.F. Arnold subsp. nigra). Our results confirmed that the two communities were considerably different, with the old-growth beech community having a lower number of plant species and the pine community was in development as a consequence of anthropogenic activities. These aspects of the two communities were also reflected in the soil, with the beech soil having higher nitrogen levels and a more specialized microbial community compared to the pine soil, with most extracellular enzymes (such as peroxidase and chitinase) showing lower activity in the pine soil.

International audience; Hightlights:• Predictions of physiological process depends on climate model, species and region.• Predictions were improved after correction for the three models considered.• Processes simulated exhibited large variability at the plot scale.• This variability faded out at larger scales, owing to an aggregation effect.• Process predictions were more variable during the driest years.Abstract: Climate change affects various aspects of ecosystem functioning, especially photosynthesis, respiration and carbon storage. We need accurate modelling approaches (impact models) to simulate forest functioning and vitality in a warmer world so that forest models can estimate multiple changes in ecosystem service provisions (e. g., productivity and carbon storage) and test management strategies to promote forest resilience.Here, we aimed to quantify the bias in these models, addressing three questions: (1) Do the predictions of impact models vary when forcing them with different climate models, and how do the predictions differ under climate model vs. observational climate forcings? (2) Does the climate impact simulation variability caused by climate forcings fade out at large spatial scales? (3) How does using simulated climate data affect process-based model predictions in stressful drought events? To answer these questions, we present historical results for 1960-2010 from the CASTANEA ecophysiological forest model and use the data from three climate models. Our analysis focuses on monospecific stands of European beech (Fagus sylvatica), temperate deciduous oaks (Quercus robur and Q. petraea), Scots pine (Pinus sylvestris) and spruce (Picea abies) in French forests.We show that prediction of photosynthesis, respiration and wood growth highly depends on the climate model used and species and region considered. Predictions were improved after a monthly mean bias or monthly quantile mapping correction for the three models considered. The processes simulated by the impact model exhibited large variability under different climate forcings at the plot scale (i.e., a few hectares). This variability faded out at larger scales (i.e., an ecological region, 100 km(2)), owing to an aggregation effect. Moreover, process predictions obtained under different climate forcings were more variable during the driest years. These results highlight the necessity of quantifying the bias correction effect on process predictions before predicting flux dynamics with a process-based model.