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.

Methane is a powerful greenhouse gas, more potent than carbon dioxide, and emitted from a variety of natural sources including wetlands, permafrost, mammalian guts and termites. As increases in global temperatures continue to break records, quantifying the magnitudes of key methane sources has never been more pertinent. Over the last 40 years, the contribution of termites to the global methane budget has been subject to much debate. The most recent estimates of termite emissions range between 9 and 15 Tg CH4 year-1, approximately 4% of emissions from natural sources (excluding wetlands). However, we argue that the current approach for estimating termite contributions to the global methane budget is flawed. Key parameters, namely termite methane emissions from soil, deadwood, living tree stems, epigeal mounds and arboreal nests, are largely ignored in global estimates. This omission occurs because data are lacking and research objectives, crucially, neglect variation in termite ecology. Furthermore, inconsistencies in data collection methods prohibit the pooling of data required to compute global estimates. Here, we summarise the advances made over the last 40 years and illustrate how different aspects of termite ecology can influence the termite contribution to global methane emissions. Additionally, we highlight technological advances that may help researchers investigate termite methane emissions on a larger scale. Finally, we consider dynamic feedback mechanisms of climate warming and land-use change on termite methane emissions. We conclude that ultimately the global contribution of termites to atmospheric methane remains unknown and thus present an alternative framework for estimating their emissions. To significantly improve estimates, we outline outstanding questions to guide future research efforts.

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.

BACKGROUND: Carbon (C) sink and stock are among the most important ecosystem services provided by forests in climate change mitigation policies. In this context, old-growth forests constitute an essential reference point for the development of close-to-nature silviculture, including C management techniques. Despite their small extent in Europe, temperate old-growth forests are assumed to be among the most prominent in terms of biomass and C stored. However, monitoring and reporting of C stocks is still poorly understood. To better understand the C stock amount and distribution in temperate old-growth forests, we estimated the C stock of two old-growth stands in the Dinaric Alps applying different assessment methods, including direct and indirect approaches (e.g., field measurements and allometric equations vs. IPCC standard methods). This paper presents the quantification and the distribution of C across the five main forest C pools (i.e., aboveground, belowground, deadwood, litter and soil) in the study areas and the differences between the applied methods.
RESULTS: We report a very prominent C stock in both study areas (507 Mg C ha- 1), concentrated in a few large trees (36% of C in 5% of trees). Moreover, we found significant differences in C stock estimation between direct and indirect methods. Indeed, the latter tended to underestimate or overestimate depending on the pool considered.
CONCLUSIONS: Comparison of our results with previous studies and data collected in European forests highlights the prominence of temperate forests, among which the Dinaric Alps old-growth forests are the largest. These findings provide an important benchmark for the development of future approaches to the management of the European temperate forests. However, further and deeper research on C stock and fluxes in old-growth stands is of prime importance to understand the potential and limits of the climate mitigation role of forests.

Conservation of biodiversity requires in-depth knowledge of trait-environment interactions to understand the influence the environment has on species assemblages. Saproxylic beetles exhibit a wide range of traits and functions in the forest ecosystems. Understanding their responses to surrounding environment thus improves our capacity to identify habitats that should be restored or protected. We investigated potential interactions between ecological traits in saproxylic beetles (feeding guilds and habitat preferences) and environmental variables (deadwood, type and age of surrounding forest). We sampled beetles from 78 plots containing newly created high stumps of Scots pine and Silver birch in boreal forest landscapes in Sweden for three consecutive years. Using a model based approach, our aim was to explore potential interactions between ecological traits and the surrounding environment at close and distant scale (20 m and 500 m radius). We found that broadleaf-preferring beetle species are positively associated with the local broadleaf-originated deadwood and broadleaf-rich forests in the surrounding landscapes. Conifer-preferring species are positively associated with the local amount of coniferous deadwood and young and old forests in the surrounding landscape. Fungivorous and predatory beetles are positively associated with old forests in the surrounding landscapes. Our results indicate that both local amounts of deadwood and types of forests in the landscape are important in shaping saproxylic beetle communities. We particularly highlight the need to increase deadwood amounts of various qualities in the landscape, exempt older forests from production and to increase broadleaf-rich habitats in order to meet different beetle species\' habitat requirements. Trait responses among saproxylic beetles provide insights into the significance of broadleaf forest and dead wood as essential attributes in boreal forest restoration, which helps conservation planning and management in forest landscapes.

Deadwood is a key old-growth element in European forests and a cornerstone of biodiversity conservation practices in the region, recognized as an important indicator of sustainable forest management. Despite its importance as a legacy element for biodiversity, uncertainties remain on the drivers of deadwood potentials, its spatial distribution in European forests and how it may change in the future due to management and climate change. To fill this gap, we combined a comprehensive deadwood dataset to fit a machine learning and a Bayesian hurdle-lognormal model against multiple environmental and socio-economic predictors. We deployed the models on the gridded predictors to forecast changes in deadwood volumes in Europe under alternative climate (RCP4.5 and RCP8.5) and management scenarios (biodiversity-oriented and production-oriented strategies). Our results show deadwood hotspots in montane forests of central Europe and unmanaged forests in Scandinavia. Future climate conditions may reduce deadwood potentials up to 13% under a mid-century climate, with regional losses amounting to up to 22% in Southern Europe. Nevertheless, changes in management towards more biodiversity-oriented strategies, including an increase in the share of mixed forests and extended rotation lengths, may mitigate this loss to a 4% reduction in deadwood potentials. We conclude that adaptive management can promote deadwood under changing environmental conditions and thereby support habitat maintenance and forest multifunctionality.

Old-growth forests provide a broad range of ecosystem services. However, due to poor knowledge of their spatiotemporal distribution, implementing conservation and restoration strategies is challenging. The goal of this study is to compare the predictive ability of socioecological factors and different sources of remotely sensed data that determine the spatiotemporal scales at which forest maturity attributes can be predicted. We evaluated various remotely sensed data that cover a broad range of spatial (from local to global) and temporal (from current to decades) extents, from Airborne Laser Scanning (ALS), aerial multispectral and stereo-imagery, Sentinel-1, Sentinel-2 and Landsat data. Using random forests, remotely sensed data were related to a forest maturity index available in 688 forest plots across four ranges of the French Alps. Each model also includes socioecological predictors related to topography, socioeconomy, pedology and climatology. We found that the different remotely sensed data provide information on the main forest structural characteristics as defined by ALS, except for Landsat, which has a too coarse resolution, and Sentinel-1, which responds differently to vegetation structure. The predictions were quite similar considering aerial remotely sensed data, on the one hand, and satellite remotely sensed data, on the other hand. Socioecological variables are the most important predictors compared to the remote sensing metrics. In conclusion, our results indicate that a wide range of remotely sensed data can be used to study old-growth forests beyond the use of ALS and despite different abilities to predict forest structure. Accounting for socioecological predictors is indispensable to avoid a significant loss of predictive accuracy. Remotely sensed data can allow for predictions to be made at different spatiotemporal resolutions and extents. This study paves the way to large-scale monitoring of forest maturity, as well as for retrospective analyses which will show to what extent predicted maturity change at different dates.

Functional trait approaches are common in ecology, but a lack of clear hypotheses on how traits relate to environmental gradients (i.e., trait-niche relationships) often makes uncovering mechanisms difficult. Furthermore, measures of community functional structure differ in their implications, yet inferences are seldom compared among metrics. Community-weighted mean trait values (CWMs), a common measure, are largely driven by the most common species and thus do not reflect community-wide trait-niche relationships per se. Alternatively, trait-niche relationships can be estimated across a larger group of species using hierarchical joint species distribution models (JSDMs), quantified by a parameter Γ. We investigated how inferences about trait-niche relationships are affected by the choice of metric. Using deadwood-dependent (saproxylic) beetles in fragmented Finnish forests, we followed a protocol for investigating trait-niche relationships by (1) identifying environmental filters (climate, forest age, and deadwood volume), (2) relating these to an ecological function (dispersal ability), and (3) identifying traits related to this function (wing morphology). We tested 18 hypothesized dispersal relationships using both CWM and Γ estimates across these environmental gradients. CWMs were more likely than Γ to show support for trait-niche relationships. Up to 13% of species\' realized niches were explained by dispersal traits, but the directions of effects were consistent with fewer than 11%-39% of our 18 trait-niche hypotheses (depending on the metric used). This highlights the difficulty in connecting morphological traits and ecological functions in insects, despite the clear conceptual link between landscape connectivity and flight-related traits. Caution is thus warranted in hypothesis development, particularly where apparent trait-function links are less clear. Inferences differ when CWMs versus Γ estimates are used, necessitating the choice of a metric that reflects study questions. CWMs help explain the effects of environmental gradients on community trait composition, whereas the effects of traits on species\' niches are better estimated using hierarchical JSDMs.

Restoration of degraded habitat is frequently used in ecological compensation. However, ecological restoration suffers from innate problems of long delivery times of features shown to be good proxies for biodiversity, e.g., large dead trees. We tested a possible way to circumvent this problem; the translocation of hard-to-come deadwood substrates from an impact area to a compensation area. Following translocation, deadwood density in the compensation area was locally equivalent to the impact area, around 20 m3 ha-1, a threshold for supporting high biodiversity of rare and red-listed species. However, deadwood composition differed between the impact and compensation area, showing a need to include more deadwood types, e.g., late decomposition deadwood, in the translocation scheme. To guide future compensation efforts, the cost for translocation at different spatial scales was calculated. We conclude that translocation of deadwood could provide a cost-efficient new tool for ecological compensation/restoration but that the method needs refinement.

Selective browsing by deer on young trees may impede the management goal of increasing forest resilience against climate change and other disturbances. Deer population density is often considered the main driver of browsing impacts on young trees, however, a range of other variables such as food availability also affect this relationship. In this study, we use browsing survey data from 135 research plots to explore patterns of roe deer (Capreolus capreolus) browsing pressure on woody plants in mountainous forests in central Europe. We fitted species-specific generalised linear mixed models for eight woody taxa, assessing the potential effects of understory characteristics, roe deer abundance and lying deadwood on browsing intensity. Our study reveals conspecific and associational effects for woody taxa that are intermediately browsed by roe deer. Selective browsing pressure was mediated by preferences of plants, in that, browsing of strongly preferred woody taxa as for example mountain ash (Sorbus aucuparia) and of least preferred woody taxa, for example Norway spruce (Picea abies) was not affected by the surrounding understory vegetation, while browsing pressure on intermediately browsed species like for example silver fir (Abies alba) was affected by understory characteristics. Contrary to our expectations, roe deer abundance was only positively associated with browsing pressure on silver fir and bilberry (Vaccinium myrtillus), while all other plants were unaffected by deer abundance. Finally, we did not find an influence of lying deadwood volume on the browsing pressure on any woody-plant species. Overall, our results indicate that patterns in browsing preference and intensity are species-specific processes and are partly affected by the surrounding understory vegetation. Current management strategies that aim to reduce browsing pressure through culling may be inefficient as they do not address other drivers of browsing pressure. However, managers also need to consider the characteristics of the local understory vegetation in addition to deer abundance and design species-specific plans to reduce browsing on woody plant taxa.