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Extending the network of strict forest reserves is one of the conservation measures promoted by the French National Strategy for Biodiversity. However, there is a lack of evidence concerning the efficiency of setting aside strict forest reserves to preserve biodiversity in the case of temperate forests. Moreover, there are potentially unexplored underlying ecological mechanisms that forest management could usefully mimic. In order to disentangle the respective roles of management abandonment, stand structural attributes and climatic and topographic variables in determining forest biodiversity, we conducted the first national-scale study in France comparing biodiversity in managed and in unmanaged forests. Here we focus on bryophytes (all species combined and forest specialists separately). We analyzed data from 127 plots in ten lowland forests in France. Our aim was to disentangle the relationships between bryophyte richness and (i) management abandonment per se, (ii) associated forest-structure variables like deadwood volume, and (iii) macroclimatic variables important for bryophytes (temperature, precipitation, relative humidity, solar radiation and vapor pressure deficit). For each studied combination of variables (univariate, additive or interactive models), we compared hierarchical models of several types: linear with a fixed slope, linear with a random slope, quadratic, sigmoid or threshold models. We found that deadwood variables were the main drivers of bryophyte richness in managed as well as in unmanaged stands. We observed a sigmoid relationship of total deadwood volume to overall richness, and a threshold effect of large and very large deadwood volume on forest specialist richness. The effect of management abandonment was globally non-significant, though impact varied strongly among the different forest sites. A combination of deadwood and macroclimatic variables best predicted bryophyte richness, through non-linear relationships: 1) higher solar radiation reinforced the positive effects of large deadwood on forest-specialist bryophyte richness; and 2) higher mean annual temperatures counteracted the positive effects of total deadwood amount on total bryophyte species richness. Maintaining high amounts of deadwood in both managed and unmanaged forests is likely to improve bryophyte richness and will be particularly important under ongoing climate change. • Deadwood is crucial for forest bryophyte richness in managed and in unmanaged stands. • The management abandonment effect was non-significant and varied widely among sites. • Nonlinear combinations of deadwood and climate data best explained species richness. • Solar radiation reinforced large-deadwood positive effect on forest specialists. • Higher temperature counteracted the deadwood positive effect on bryophyte richness. [ABSTRACT FROM AUTHOR]

Balancing increasing demand for wood products while also maintaining forest biodiversity is a paramount challenge. Europe's Biodiversity and Forest Strategies for 2030 attempt to address this challenge. Together, they call for strict protection of 10% of land area, including all primary and old growth forests, increasing use of ecological forestry, and less reliance on monocultural plantations. Using data on country wide silvicultural practices and a new database on strict forest reserves across Europe, we assess how triad forest zoning could help meet these goals. Our analysis reveals that zoning in Europe is overwhelmingly focused on wood production, while there has been little concomitant protection of forests in strict reserves. Moreover, most strict forest reserves are < 50 ha in size, likely too small to capture the minimum dynamic area necessary to sustain many taxa. We outline research priorities to meet future demands for timber while minimizing the impact on native biodiversity., (© 2024. The Author(s).)

We present a data set resulting from the first round of a national monitoring program of forest reserves. It contains 9538 permanent plots, distributed across 111 study sites in mainland France (including Corsica). Notably focusing on dead wood measurement, this protocol has primarily been applied in strict forest reserves and special nature reserves (sensu Bollmann & Braunisch 2013), with 68% (6494) of the plots being currently located in strict forest reserves (unmanaged) and 24.7% (2363 plots) in forests unmanaged for at least 50 years. Sites cover a large variety of ecological conditions, from lowland to subalpine forests, but with an underrepresentation of Mediterranean forests (Table 1). The protocol assesses all the stages of a tree's life cycle, from seedling to decomposed lying dead wood. On each plot, a combination of three sampling techniques was used: (1) fixed-area inventory for regeneration, standing dead trees, living trees, and coarse woody debris (CWD) with diameter over 30 cm; (2) transect lines for CWD with diameter <30 cm; and (3) fixed-angle plot method for living trees with diameter at breast height (DBH) >30 cm (using a relascopic angle of 3%). Measurements include exact tree location (azimuth, distance), species, diameter(s), tree-related microhabitats, decay stage and bark cover, and seedling cover. With ongoing climate change, the program network can also provide important information to monitor changes in forest ecosystems. It can also be used as forest management monitoring or conservation status assessment. These data are freely available for noncommercial scientific use (Creative Commons Attribution 4.0 CC BY SA 4.0) with attribution, and this paper must be cited if this material is reused., (© 2024 The Ecological Society of America.)

Protecting structural features, such as tree-related microhabitats (TreMs), is a cost-effective tool crucial for biodiversity conservation applicable to large forested landscapes. Although the development of TreMs is influenced by tree diameter, species, and vitality, the relationships between tree age and TreM profile remain poorly understood. Using a tree-ring-based approach and a large data set of 8038 trees, we modeled the effects of tree age, diameter, and site characteristics on TreM richness and occurrence across some of the most intact primary temperate forests in Europe, including mixed beech and spruce forests. We observed an overall increase in TreM richness on old and large trees in both forest types. The occurrence of specific TreM groups was variably related to tree age and diameter, but some TreM groups (e.g., epiphytes) had a stronger positive relationship with tree species and elevation. Although many TreM groups were positively associated with tree age and diameter, only two TreM groups in spruce stands reacted exclusively to tree age (insect galleries and exposed sapwood) without responding to diameter. Thus, the retention of trees for conservation purposes based on tree diameter appears to be a generally feasible approach with a rather low risk of underrepresentation of TreMs. Because greater tree age and diameter positively affected TreM development, placing a greater emphasis on conserving large trees and allowing them to reach older ages, for example, through the establishment of conservation reserves, would better maintain the continuity of TreM resource and associated biodiversity. However, this approach may be difficult due to the widespread intensification of forest management and global climate change., (© 2023 Society for Conservation Biology.)

In Europe, forest management has controlled forest dynamics to sustain commodity production over multiple centuries. Yet over-regulation for growth and yield diminishes resilience to environmental stress as well as threatens biodiversity, leading to increasing forest susceptibility to an array of disturbances. These trends have stimulated interest in alternative management systems, including natural dynamics silviculture (NDS). NDS aims to emulate natural disturbance dynamics at stand and landscape scales through silvicultural manipulations of forest structure and landscape patterns. We adapted a "Comparability Index" (CI) to assess convergence/divergence between natural disturbances and forest management effects. We extended the original CI concept based on disturbance size and frequency by adding the residual structure of canopy trees after a disturbance as a third dimension. We populated the model by compiling data on natural disturbance dynamics and management from 13 countries in Europe, covering four major forest types (i.e., spruce, beech, oak, and pine-dominated forests). We found that natural disturbances are highly variable in size, frequency, and residual structure, but European forest management fails to encompass this complexity. Silviculture in Europe is skewed toward even-aged systems, used predominately (72.9% of management) across the countries assessed. The residual structure proved crucial in the comparison of natural disturbances and silvicultural systems. CI indicated the highest congruence between uneven-aged silvicultural systems and key natural disturbance attributes. Even so, uneven-aged practices emulated only a portion of the complexity associated with natural disturbance effects. The remaining silvicultural systems perform poorly in terms of retention compared to tree survivorship after natural disturbances. We suggest that NDS can enrich Europe's portfolio of management systems, for example where wood production is not the primary objective. NDS is especially relevant to forests managed for habitat quality, risk reduction, and a variety of ecosystem services. We suggest a holistic approach integrating NDS with more conventional practices., (© 2022 The Ecological Society of America.)

Conventional conservation policies in Europe notably rely on the passive restoration of natural forest dynamics by setting aside forest areas to preserve forest biodiversity. However, since forest reserves cover only a small proportion of the territory, conservation policies also require complementary conservation efforts in managed forests in order to achieve the biodiversity targets set up in the Convention on Biological Diversity. Conservation measures also raise the question of large herbivore management in and around set-asides, particularly regarding their impact on understory vegetation. Although many studies have separately analyzed the effects of forest management, management abandonment, and ungulate pressure on forest biodiversity, their joint effects have rarely been studied in a correlative framework. We studied 212 plots located in 15 strict forest reserves paired with adjacent managed forests in European France. We applied structural equation models to test the effects of management abandonment, stand structure, and ungulate pressure on the abundance, species richness, and diversity of herbaceous vascular plants and terricolous bryophytes. We showed that stand structure indices and plot-level browsing pressure had direct and opposite effects on herbaceous vascular plant species diversity; these effects were linked with the light tolerance of the different species groups. Increasing canopy cover had an overall negative effect on herbaceous vascular plant abundance and species diversity. The effect was two to three times greater in magnitude than the positive effects of browsing pressure on herbaceous plants diversity. On the other hand, a high stand density index had a positive effect on the species richness and diversity of bryophytes, while browsing had no effect. Forest management abandonment had few direct effects on understory plant communities, and mainly indirectly affected herbaceous vascular plant and bryophyte abundance and species richness and diversity through changes in vertical stand structure. Our results show that conservation biologists should rely on foresters and hunters to lead the preservation of understory vegetation communities in managed forests since, respectively, they manipulate stand structure and regulate ungulate pressure. Their management actions should be adapted to the taxa at stake, since bryophytes and vascular plants respond differently to stand and ungulate factors., (© 2022 The Ecological Society of America.)

Managing forests to preserve biodiversity requires a good knowledge not only of the factors driving its dynamics but also of the structural elements that actually support biodiversity. Tree-related microhabitats (e.g. cavities, cracks, conks of fungi) are tree-borne features that are reputed to support specific biodiversity for at least a part of species' life cycles. While several studies have analysed the drivers of microhabitats number and occurrence at the tree scale, they remain limited to a few tree species located in relatively narrow biogeographical ranges. We used a nationwide database of forest reserves where microhabitats were inventoried on more than 22,000 trees. We analysed the effect of tree diameter and living status (alive or dead) on microhabitat number and occurrence per tree, taking into account biogeoclimatic variables and tree genus. We confirmed that larger trees and dead trees bore more microhabitats than their smaller or living counterparts did; we extended these results to a wider range of tree genera and ecological conditions than those studied before. Contrary to our expectations, the total number of microhabitat types per tree barely varied with tree genus-though we did find slightly higher accumulation levels for broadleaves than for conifers-nor did it vary with elevation or soil pH, whatever the living status. We observed the same results for the occurrence of individual microhabitat types. However, accumulation levels with diameter and occurrence on dead trees were higher for microhabitats linked with wood decay processes (e.g. dead branches or woodpecker feeding holes) than for other, epixylic, microhabitats such as epiphytes (ivy, mosses and lichens). Promoting large living and dead trees of several tree species may be a relevant, and nearly universal, way to favour microhabitats and enhance the substrates needed to support specific biodiversity. In the future, a better understanding of microhabitat drivers and dynamics at the tree scale may help to better define their role as biodiversity indicators for large-scale monitoring., Competing Interests: The authors have declared that no competing interests exist.

Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant-plant interactions and through soil disturbance. In forest ecosystems, researchers' attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation-wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light-demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non-forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape-level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition., (©2017 The Authors. Global Change Biology Published by John Wiley & Sons Ltd.)

Past and present pressures on forest resources have led to a drastic decrease in the surface area of unmanaged forests in Europe. Changes in forest structure, composition, and dynamics inevitably lead to changes in the biodiversity of forest-dwelling species. The possible biodiversity gains and losses due to forest management (i.e., anthropogenic pressures related to direct forest resource use), however, have never been assessed at a pan-European scale. We used meta-analysis to review 49 published papers containing 120 individual comparisons of species richness between unmanaged and managed forests throughout Europe. We explored the response of different taxonomic groups and the variability of their response with respect to time since abandonment and intensity of forest management. Species richness was slightly higher in unmanaged than in managed forests. Species dependent on forest cover continuity, deadwood, and large trees (bryophytes, lichens, fungi, saproxylic beetles) and carabids were negatively affected by forest management. In contrast, vascular plant species were favored. The response for birds was heterogeneous and probably depended more on factors such as landscape patterns. The global difference in species richness between unmanaged and managed forests increased with time since abandonment and indicated a gradual recovery of biodiversity. Clearcut forests in which the composition of tree species changed had the strongest effect on species richness, but the effects of different types of management on taxa could not be assessed in a robust way because of low numbers of replications in the management-intensity classes. Our results show that some taxa are more affected by forestry than others, but there is a need for research into poorly studied species groups in Europe and in particular locations. Our meta-analysis supports the need for a coordinated European research network to study and monitor the biodiversity of different taxa in managed and unmanaged forests.