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.

The shift from shade‐intolerant species to shade‐tolerant mesophytic species in deciduous and mixed forests of the temperate zone is well described in studies from North America. This process has been termedmesophicationand it has been linked to changes in fire regime. Fire suppression results in the cessation of establishment of heliophytic, fire‐dependent tree species such as oak (Quercus) and pine (Pinus). Due to the scarcity of old‐growth forests in Europe, data on long‐term compositional changes in mixed forests are very limited, as is the number of studies exploring whether fire played a role in shaping the dynamics. The aim of this study was to reconstruct tree succession in a 43‐ha natural mixed deciduous forest stand in Białowieża Forest (BF), Poland using dendrochronological methods. In addition, the presence of aboveground fire legacies (charred and fire‐scarred deadwood) enabled the fire history reconstruction. Dendrochronological data revealed tree establishment (Quercus) back to the end of the 1500s and fires back to 1659. Under a regime of frequent fires until the end of the 18th century, only oak and pine regenerated, sporadically. A shift in the fire regime in the first half of the 19th century triggered oak and pine cohort regeneration, then gradually spruce (Picea) encroached. Under an increasingly dense canopy and less flammable conditions, regeneration of shade‐tolerantCarpinus,Tilia,andAcerbegan simultaneously with the cessation of oak and pine recruitment. Synthesis. The study reports the first evidence of mesophication in temperate Europe and proves that fire was involved in shaping the long‐term dynamics of mixed deciduous forest ecosystems. Our data suggest that fire exclusion promoted a gradual recruitment of fire‐sensitive, shade‐tolerant species that inhibited the regeneration of oak and pine in BF.

The impact of forest management on biodiversity is difficult to scrutinize along gradients of management. A step towards analyzing the impact of forest management on biodiversity is comparisons between managed and primary forests. The standardized typology of tree-related microhabitats (TreMs) is a multi-taxon indicator used to quantify forest biodiversity. We aim to analyze the influence of environmental factors on the occurrence of groups of TreMs by comparing primary and managed forests. We collected data for the managed forests in the Black Forest (Germany) and for the primary forests in the Western (Slovakia) and Southern Carpathians (Romania). To model the richness and the different groups of TreMs per tree, we used generalized linear mixed models with diameter at breast height (DBH), altitude, slope and aspect as predictors for European beech (Fagus sylvatica (L.)), Norway spruce (Picea abies (L.)) and silver fir (Abies alba (Mill.)) in primary and managed temperate mountain forests. We found congruent results for overall richness and the vast majority of TreM groups. Trees in primary forests hosted a greater richness of all and specific types of TreMs than individuals in managed forests. The main drivers of TreMs are DBH and altitude, while slope and aspect play a minor role. We recommend forest and nature conservation managers to focus: 1) on the conservation of remaining primary forests and 2) approaches of biodiversity-oriented forest management on the selection of high-quality habitat trees that already provide a high number of TreMs in managed forests based on the comparison with primary forests.

Retention of habitat trees is a common biodiversity conservation practice in continuous cover forests of temperate Europe. Commonly, living habitat trees are selected on the basis of their tree-related microhabitats (TreMs) such as cavities or crown deadwood. Owing to the increasing frequency and intensity of climate change-related disturbances, habitat trees in particular are expected to experience increased mortality rates. This may impact the long-term provisioning of TreMs. Here, we compared the TreM occurrence on living and dead trees to investigate whether dead trees support more and other TreMs than living trees. We also hypothesized that a combination of living and dead trees results in the most diverse stand-level TreM composition. We surveyed the TreM composition of living and dead habitat trees in 133 one-hectare plots in the Black Forest region managed according to a continuous cover approach. We fitted generalized linear mixed models to identify the main predictors of TreM occurrence to predict their abundance and richness. Tree identity (as a combination of species and vitality status) and diameter were the main drivers of TreM abundance and richness, which were highest on dead Abies alba. Even though dead A. alba and Picea abies supported TreM numbers similar to those provided by large living trees, their TreM composition was significantly different. This suggests that dead trees cannot substitute the habitat functions of living habitat trees, but they can complement them to increase the overall stand-level TreM diversity, in particular through decayed, large snags.

Retention of structural elements such as deadwood and habitat trees at the level of forest stands has been promoted to integrate biodiversity conservation into multiple‐use forest management. The conservation value of habitat trees is largely determined by the presence, richness, and abundance of tree‐related microhabitats (TreMs). Since TreMs are often lacking in intensively managed forests, an important question of forest conservation is how the abundance and richness of TreMs may be effectively restored. Here, we investigated whether the strict protection of forest through cessation of timber harvesting influenced TreM occurrence at tree and stand levels. For that purpose, we compared four managed and four set‐aside stands (0.25 ha each) in the Białowieża Forest, with identical origin following clear‐cuts approximately 100 years ago. We found that the abundance and richness of TreMs on living trees were not significantly different between stands that were either conventionally managed or where active forest management ceased 52 years ago. Yet, our analysis of TreMs on tree species with contrasting life‐history traits revealed that short‐lived, fast‐growing species (pioneers) developed TreMs quicker than longer‐lived, slower‐growing species. Hence, tree species such as Populus or Betula, which supply abundant and diverse TreMs, can play an important role in accelerating habitat restoration.

Timber‐oriented forest management profoundly alters forest structure and composition, with complex effects on associated biodiversity. However, while species' responses to forest management and resulting structural characteristics have been the subject of numerous studies, direct and indirect effects that cascade through trophic levels are rarely disentangled. As insectivorous bats are particularly sensitive to changes in forest structure, that shape their available flight space, we investigated how forest structure, composition and management also indirectly modify their habitats, for example, by affecting important insect prey groups. We used structural equation models (SEMs) to test bat responses to forest composition, structure (forest heterogeneity, old‐growth attributes) and management intensity, quantifying direct and indirect prey‐mediated effects. For that, three bat guilds—short‐ (SRE), mid‐ (MRE) and long‐range echolocating (LRE) bats—and their prey insects (moths and ground beetles) were analysed from 64 sites in the Black Forest, Germany. We found guild‐specific effects on bats: While the structural heterogeneity of forests directly influenced the activity of bat guilds, the main influence of forest management, composition and structure was mediated through their prey‐groups. SRE activity responded to moths and LRE activity was associated with ground beetles, with positive effects of the insect groups' abundance, but negative effects of the same group's species richness. In addition, the SEM approach revealed a negative top‐down relationship between MRE activities and moths, suggesting predation or avoidance behaviour of moths. While forest management directly or indirectly increased prey insect abundance, it negatively affected the availability of roosting structures for bats. Synthesis and applications. The results highlight the indirect and positive effects of forest management on bats and support the important role of bats in insect regulation within continuous cover forests. Although forest management created small gaps that improved foraging habitats for most bats, it compromised the roosting functionality for bats. The ‘close‐to‐nature forestry’ currently prevalent in Europe mainly promotes continuous‐cover forests in mid‐successional stages. Expanding the forest management portfolio towards open and old‐growth forests would increase roosting opportunities and provide complementary foraging habitats for different bat species, while promoting high biodiversity in managed forest landscapes.