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.

1. 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 termed mesophication and 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.2. 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. 3. 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-tolerant Carpinus, Tilia,and Acer began simultaneously with the cessation of oak and pine recruitment. Synthesis. The study reports the first evidence of mesophication in temperateEurope 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. K E Y W O R D Sage structure, disturbance regime, fire suppression, forest history, oak trees, pine trees

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 sylvatica L.) 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.

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.

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Research Highlights: Past disturbances occurred naturally in primary forests in the Southern Carpathians. High- and moderate-severity disturbances shaped the present structure of these ecosystems, which regenerated successfully without forestry interventions. Background and Objectives: Windstorms and bark beetle outbreaks have recently affected large forest areas across the globe, causing concerns that these disturbances lie outside the range of natural variability of forest ecosystems. This often led to salvage logging inside protected areas, one of the main reasons for primary forest loss in Eastern Europe. Although more than two-thirds of temperate primary forests in Europe are located in the Carpathian region of Eastern Europe, knowledge about how natural disturbances shape the forest dynamics in this region is highly essential for future management decisions. Material and Methods: We established our study in a primary forest valley situated in the centre of the largest temperate primary forest landscape in Europe (Făgăraș Mountains). A dendrochronological investigation was carried out to reconstruct the natural disturbance history and relate it to the present forest structure. Results: The dendrochronological analysis revealed high temporal variability in the disturbance patterns both at the patch and stand level. Moderate severity disturbance events were most common (20–40% of canopy disturbed in 60% of the plots) but high severity events did also occur (33% of the plots). Regeneration was spruce-dominated and 71% of the seedlings were found on deadwood microsites. Conclusions: We conclude that the current structure of the studied area is a consequence of the past moderate-severity disturbances and sporadic high-severity events. The peak in disturbances (1880–1910) followed by reduced disturbance rates may contribute to a recent and future increase in disturbances in the Făgăraș Mts. Our findings show that these disturbance types are within the range of natural variability of mountain spruce forests in the Southern Carpathians and should not be a reason for salvage logging in primary forests from this area.

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