Aims: Primary forests are critical for forest biodiversity and provide key ecosystem services. In Europe, these forests are particularly scarce and it is unclear whether they are sufficiently protected. Here we aim to: (a) understand whether extant primary forests are representative of the range of naturally occurring forest types, (b) identify forest types which host enough primary forest under strict protection to meet conservation targets and (c) highlight areas where restoration is needed and feasible.Location: Europe.Methods: We combined a unique geodatabase of primary forests with maps of forest cover, potential natural vegetation, biogeographic regions and protected areas to quantify the proportion of extant primary forest across Europe's forest types and to identify gaps in protection. Using spatial predictions of primary forest locations to account for underreporting of primary forests, we then highlighted areas where restoration could complement protection.Results: We found a substantial bias in primary forest distribution across forest types. Of the 54 forest types we assessed, six had no primary forest at all, and in two‐thirds of forest types, less than 1% of forest was primary. Even if generally protected, only ten forest types had more than half of their primary forests strictly protected. Protecting all documented primary forests requires expanding the protected area networks by 1,132 km² (19,194 km² when including also predicted primary forests). Encouragingly, large areas of non‐primary forest existed inside protected areas for most types, thus presenting restoration opportunities.Main conclusion: Europe's primary forests are in a perilous state, as also acknowledged by EU's “Biodiversity Strategy for 2030.” Yet, there are considerable opportunities for ensuring better protection and restoring primary forest structure, composition and functioning, at least partially. We advocate integrated policy reforms that explicitly account for the irreplaceable nature of primary forests and ramp up protection and restoration efforts alike.

Languages employ different strategies to transmit structural and grammatical information. While, for example, grammatical dependency relationships in sentences are mainly conveyed by the ordering of the words for languages like Mandarin Chinese, or Vietnamese, the word ordering is much less restricted for languages such as Inupiatun or Quechua, as these languages (also) use the internal structure of words (e.g. inflectional morphology) to mark grammatical relationships in a sentence. Based on a quantitative analysis of more than 1,500 unique translations of different books of the Bible in almost 1,200 different languages that are spoken as a native language by approximately 6 billion people (more than 80% of the world population), we present large-scale evidence for a statistical trade-off between the amount of information conveyed by the ordering of words and the amount of information conveyed by internal word structure: languages that rely more strongly on word order information tend to rely less on word structure information and vice versa. Or put differently, if less information is carried within the word, more information has to be spread among words in order to communicate successfully. In addition, we find that–despite differences in the way information is expressed–there is also evidence for a trade-off between different books of the biblical canon that recurs with little variation across languages: the more informative the word order of the book, the less informative its word structure and vice versa. We argue that this might suggest that, on the one hand, languages encode information in very different (but efficient) ways. On the other hand, content-related and stylistic features are statistically encoded in very similar ways.

The high tree mortality during the dry and hot years of 2018–2019 in Europe has triggered concerns on the future of European beech (Fagus sylvaticaL.) forests under climate change and raised questions as to whether forest management may increase tree mortality. We compared long‐term mortality rates of beech between managed and unmanaged stands including the years 2018‐2019 at 11 sites in Hesse, Germany. We hypothesized that mortality would increase with climate water deficits during the growing season, initial stand density, decreasing dominance of trees, and decreasing intensity of tree removals. Initial stand density, tree removals, the climate water balance and the competitive status of trees were used as predictor variables. Mean annual natural mortality rates ranged between 0.5% and 2.1%. Even in the drought years, we observed no signs of striking canopy disintegration. The significantly higher mortality (1.6–2.1%) in unmanaged stands during the drought years 2018 and 2019 was largely confined to suppressed trees. There was no significant increase of mortality in managed stands during the drought years, but a shift in mortality towards larger canopy trees. Our study did not confirm a general influence of management, in the form of tree removals, on mortality rates. Yet, we found that during drought years, management changed the distribution of mortality within the tree community. To analyse the effects of management on mortality rates more comprehensively, a wider gradient in site moisture conditions, including sites drier than in this study, and longer post‐drought periods should be employed.

Tree regeneration is a key process for long‐term forest dynamics, determining changes in species composition and shaping successional trajectories. While tree regeneration is a highly stochastic process, tree regeneration studies often cover narrow environmental gradients only, focusing on specific forest types or species in distinct regions. Thus, the larger‐scale effects of temperature, water availability, and stand structure on tree regeneration are poorly understood. We investigated these effects in respect of tree recruitment (in‐growth) along wide environmental gradients using forest inventory data from Flanders (Belgium), northwestern Germany, and Switzerland covering more than 40 tree species. We employed generalized linear mixed models to capture the abundance of tree recruitment in response to basal area, stem density, shade casting ability of a forest stand as well as site‐specific degree‐day sum (temperature), water balance, and plant‐available water holding capacity. We grouped tree species to facilitate comparisons between species with different levels of tolerance to shade and drought. Basal area and shade casting ability of the overstory had generally a negative impact on tree recruitment, but the effects differed between levels of shade tolerance of tree recruitment in all study regions. Recruitment rates of very shade‐tolerant species were positively affected by shade casting ability. Stem density and summer warmth (degree‐day sum) had similar effects on all tree species and successional strategies. Water‐related variables revealed a high degree of uncertainty and did not allow for general conclusions. All variables had similar effects independent of the varying diameter thresholds for tree recruitment in the different data sets. Synthesis:Shade tolerance and stand structure are the main drivers of tree recruitment along wide environmental gradients in temperate forests. Higher temperature generally increases tree recruitment rates, but the role of water relations and drought tolerance remains uncertain for tree recruitment on cross‐regional scales.

Dead wood and stand structural parameters were sampled in eleven countries using standardized methods at about 90 intensive forest monitoring sites across large parts of Europe. Besides descriptions and correlation analyses of dead wood and stand structure parameters, a joint evaluation of both fields was performed by principal component analysis (PCA). The extracted principal components were subsequently regressed against important numerical and categorical site-related parameters like soil pH, altitude, or forest type. Dead wood volumes varied largely across plots, however, 77 percent of them had volumes below 25 cubic meter per hectare. While all fractions of dead wood - except cut stumps - reveal high intercorrelation, different aspects of stand structure varied more independently. Clark-Evans index, number of tree species and standard deviation of tree trunk diameters revealed as most self-contained. The 1st PCA axis covered 46 percent of the total variance and was mostly loaded by total dead wood volume denoting it as the feature differentiating forests most. The 2nd axis was primarily loaded by tree species diversity together with stem density and the Clark-Evans index. On the 3rd axis diameter differentiation of trees together with the volume of cut stumps prevailed, while the 4th was mainly related to the decay class of woody debris. Bivariate ex post analyses revealed country as a significant predictor of all PCA axes, underlining national forest legislations and management rules as crucial for all investigated structural features of forests. Forest type was related only to the 3rd and 2nd axis. Only the 3rd axis revealed significant relationships with some ecological site factors (age, number of tree layers, latitude, altitude). The outcome underlines the significance of nationally enacted forest legislations for both important structural and biodiversity-relevant features of forest ecosystems and encourages similar approaches with data from national forest inventories or monitoring systems.

Currently, the dynamics underlying the storage and acquisition of biomass, and thus carbon, in naturally developing forests are under debate. A better understanding of the biomass dynamics of forests is needed to clarify the role played by naturally developing forests in the mitigation of climate change. Long‐term monitoring data from unmanaged strict forest reserves (SFRs) in north‐western Germany were used to analyse the biomass dynamics of pure beech, mixed beech and mixed oak forests. A complete balance of above‐ground woody biomass (biomass) and growth, density‐dependent and ‐independent mortality, as well as deadwood decay was derived. Density‐independent mortality served as a proxy for disturbance severity. After a time since abandonment (TSA) of 50 years, the average biomass ranged between 334 t/ha in mixed oak and 478 t/ha in pure beech stands. The net change in biomass was positive in all forest types. Density‐independent mortality and decay rates were much lower than the growth rates. Pure beech forests reached higher levels of biomass, a higher net change in biomass, and more growth than either of the mixed forest types. Biomass increased linearly with TSA in pure beech stands but followed an asymptotic course in the mixed forests. In the latter, the net change in biomass and growth were consistent with a unimodal development pattern. The development of biomass could not be explained by the ageing of the tree communities. Synthesis. We hypothesized that the observed biomass dynamics are a result of the interaction between resource supply within a limited growing space and the resource‐use efficiency of the tree stand in conjunction with disturbances. The still‐linear increase in the biomass of pure beech forests was assumed to reflect the high resource‐use efficiency of beech, especially its use of light. The above‐ground capacity of naturally developing broadleaved forests to store and acquire carbon is substantial. Accordingly, allowing broadleaved forests to develop naturally can contribute substantially to carbon storage and sequestration. However, our study also suggests that the above‐ground carbon sink decreases after several decades.