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There is a rising interest in the role of species diversity in ecosystem functioning and services, including productivity. Yet, how the diversity-productivity relationship depends on species identity and abiotic conditions remains a challenging issue. We analysed mixture effects on species productivity along site productivity gradients, calculated from a set of abiotic factors, in two biogeographic contexts (highlands and lowlands). We compared the productivity of 5 two-species mixtures (i.e. 10 cases of mixed species) with that of monocultures of the same species. Five main European tree species were considered: sessile oak (Quercus petraea Liebl.), Scots pine (Pinus sylvestris L.), European beech (Fagus sylvatica L.), silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) H. Karst). Our data set was compiled from the 2006 to 2010 French National Forest Inventory data base and covers 2361 plots including pure and mixed stands. Overall productivity of mixtures in highlands, that is European beech-Norway spruce, European beech-silver fir and to a lesser extent, silver fir-Norway spruce, was found to be higher than expected from the productivity of corresponding monospecific stands. Overyielding was mainly due to European beech for the first two mixtures and to silver fir for the third one. No effect of mixture was found for sessile oak-Scots pine and sessile oak-European beech stands in lowlands. Overyielding of sessile oak mixed with Scots pine was not strong enough to significantly increase overall stand productivity. Overyielding of European beech was balanced by an underyielding of sessile oak. The mixture effect changed along site productivity gradients for six cases out of the 10 studied, with a stronger and positive effect on sites with low productivity. The magnitude of this change along site productivity gradients varied up to 89% depending on the tree species.Synthesis. The nature of species interaction in mixtures with regard to productivity changes with species assemblage and abiotic conditions. Overyielding is strongest when species grow in highlands on less productive sites. A negative link between mixture effect and site productivity was found, in line with the stress-gradient hypothesis.

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.

Aims The growing demand from forest managers is to identify silvicultural practices to overcome projected water scarcity during the next decades. One solution is to mix tree species in the same stand, thereby increasing resource partitioning and minimizing competition for limited soil water. This study investigates the mixture approach for Quercus petraea (Matt.) Liebl. and Pinus sylvestris L. during an extreme summer drought event. Methods During the summer drought event in 2016, we analyzed the isotopic signatures of large- and small-tree xylem and soil water throughout the soil profile to assess the depth of water uptake for both tree species. We also measured predawn leaf water potentials (PLWP) to assess water availability for individual tree species. Results When grown in pure stands, both species primarily utilized soil water near the surface. In contrast, partial niche complementarity for limited water resources between the two species in mixed stands resulted in less water constraint (i.e., less negative PLWP) for oak trees compared to pure stands, especially for small trees. Conclusions Results from this study show that contrasting water use strategies can change water availability for trees and could help some species, though not all, to cope with the water scarcity predicted in a changing climate.

1. National and international forest biodiversity assessments largely rely on indirect indicators, based on elements of forest structure that are used as surrogates for species diversity. These proxies are reputedly easier and cheaper to assess than biodiversity. Tree microhabitats - tree-borne singularities such as cavities, conks of fungi or bark characteristics - have gained attention as potential forest biodiversity indicators. However, as with most biodiversity indicators, there is a lack of scientific evidence documenting their quantitative link with the biodiversity they are supposed to assess. 2. We explored the link between microhabitat indices and the richness and abundance of three taxonomic groups: bats, birds, and saproxylic beetles. Using a nation-wide multi-taxon sampling design in France, we compared 213 plots located inside and outside strict forest reserves. We hypothesized that the positive effect setting aside forest reserves has on biodiversity conservation is indirectly due to an increase in the proportion of large structural elements (e.g. living trees, standing and lying deadwood). These, in turn, are likely to favour the quantity and diversity of microhabitats. We analysed the relationship between the abundance and species richness of different groups and guilds (e.g. red-listed species, forest specialists, cavity dwellers) and microhabitat density and diversity. We then used confirmatory structural equation models to assess the direct and indirect effects of management abandonment, large structural elements and microhabitats on the biodiversity of the target species. 3. For several groups of birds and bats, the indirect effect of management abandonment and large structural elements on biodiversity was mediated by microhabitats. However, the magnitude of the link between microhabitat indices and biodiversity was moderate. In particular, saproxylic beetles' biodiversity was poorly explained by microhabitats, large structural elements or management abandonment. 4. Synthesis and applications: Tree microhabitats may serve as indicators for bats and birds, but they are not a universal biodiversity indicator. Rather, compared to large structural elements, they most likely have a complementary role to biodiversity. In terms of forest management and conservation, preserving diversity of microhabitats at the local scale benefits several groups of both bats and birds.

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.

With the aim of wood production with negligible negative effects on biodiversity and ecosystem processes, a silvicultural practice of selective logging with natural regeneration has been implemented in European beech forests (Fagus sylvatica) during the last decades. Despite this near-to-nature strategy, species richness of various taxa is lower in these forests than in unmanaged forests. To develop guidelines tominimize the fundamental weaknesses in the current practice, we linked functional traits of saproxylic beetle species to ecosystem characteristics. We used continental-scale data from 8 European countries and regional-scale data from a large forest in southern Germany and forest-stand variables that represented a gradient of intensity of forest use to evaluate the effect of current near-to-nature management strategies on the functional diversity of saproxylic beetles. Forest-stand variables did not have a statistically significant effect on overall functional diversity, but they did significantly affect community mean and diversity of single functional traits. As the amount of dead wood increased the composition of assemblages shifted toward dominance of larger species and species preferring dead wood of large diameter and in advanced stages of decay. The mean amount of dead wood across plots in which most species occurred was from 20 to 60 m3/ha. Species occurring in plots with mean dead wood >60 m3/ha were consistently those inhabiting dead wood of large diameter and in advanced stages of decay. On the basis of our results, to make current wood-production practices in beech forests throughout Europe more conservation oriented (i.e., promoting biodiversity and ecosystem functioning), we recommend increasing the amount of dead wood to >20 m3/ha; not removing dead wood of large diameter (50 cm) and allowing more dead wood in advanced stages of decomposition to develop; and designating strict forest reserves, with their exceptionally high amounts of dead wood, that would serve as refuges for and sources of saproxylic habitat specialists.

Aim: Biodiversity monitoring and conservation are extremely complex, and surrogate taxa may represent proxies to test methods and solutions. However, cross-taxon correlations in species diversity (i.e., cross-taxon congruence) may vary widely with spatial scale. Our goal is to assess how cross-taxon congruence varies with spatial scale in European temperate forests. We expect that congruence in species diversity increases when shifting from fine to coarse spatial scales, with differences between species richness and composition, and across pairs of taxonomic groups. Location: European temperate forests. Time period: Present. Major taxa studied: Vascular plants, bryophytes, birds, epiphytic lichens, saproxylic beetles and wood-inhabiting fungi. Methods: We used field data (354 plots across 23 sites) encompassing Italy, France and Hungary, with species information for six taxonomic groups. We accounted separately for spatial grain (the size of elementary sampling unit) and extent (the geographical area included in the survey) and evaluated the relationships within all the possible pairs of taxa. Results: Although no pair of taxa had its species richness consistently correlated across scales, we found no changes in the direction of correlations when analysing species composition. However, when increasing grain and extent, we did find a general increase in the magnitude of correlations in species composition and partial changes in significance, with plants having the highest number of significant correlations. Main conclusions: Species richness congruence among taxa is strongly scale dependent owing to differences in the relative contribution of large- and small-scale processes across taxa. Cross-taxon congruence in species composition is scale dependent only for its magnitude, because life-history traits of individual species make responses to environmental factors similar across scales. Forest monitoring should consider multi-taxon sampling and limit the use of surrogates at specific spatial scales, especially for species richness. Sampling plant species composition in scattered plots across different sites may effectively summarize the whole community composition.

1. Assessing how species productivity in mixtures is influenced by species shade tolerance (ST) and phylogeny would be helpful to improve our general understanding of the relationship between tree species diversity and productivity in forests. 2. We investigated the effects of differences in ST and phylogenetic distances (PDs) between pairs of species on the productivity of Quercus petraea growing in 18 mixtures in lowland temperate forests. We calculated the mixture effect as the difference in productivity of Q. petraea in mixed vs. pure stands. Our analyses were based on data from seven annual campaigns of the French National Forest Inventory covering 1,573 plots. 3. The mixture effect on Q. petraea productivity increased when the ST of the companion species decreased. Compared to its productivity in pure stands, Q. petraea productivity in mixed coniferous stands varied from from

The current decline of wild bees puts important ecosystem services such as pollination at risk. Both inventory and monitoring programs are needed to understand the causes of wild bee decline. Effective insect monitoring relies on both mass-trapping methods coupled with rapid and accurate identifications. Identifying wild bees using only morphology can be challenging, in particular, specimens from mass-trapped samples which are often in poor condition. We generated DNA barcodes for 2931 specimens representing 157 species (156 named and one unnamed species) and 28 genera. Automated cluster delineation reveals 172 BINs (Barcodes Index Numbers). A total of 36 species (22.93%) were found in highly urbanized areas. The majority of specimens, representing 96.17% of the species barcoded form reciprocally exclusive groups, allowing their unambiguous identification. This includes several closely related species notoriously difficult to identify. A total of 137 species (87.26%) show a \"one-to-one\" match between a named species and the BIN assignment. Fourteen species (8.92%) show deep conspecific lineages with no apparent morphological differentiation. Only two species pairs shared the same BIN making their identification with DNA barcodes alone uncertain. Therefore, our DNA barcoding reference library allows reliable identification by non-experts for the vast majority of wild bee species in the Loire Valley.

1. Semi-natural grasslands are rich in biodiversity and thus important habitats for conservation, yet they are experiencing rapid declines due to agricultural intensification and abandonment. Promoting a more diverse mammalian herbivore community, including large and megaherbivores, may result in positive cascade effects for biodiversity and ecosystem functioning. Therefore, reintroducing an ecologically functional substitute of an extinct large herbivore could mitigate current biodiversity declines and foster semi-natural grassland conservation. 2. To test this hypothesis, we set up a 3-year rewilding experiment where 12 feral horses were introduced in three 10-hectare enclosure replicates (four horses per enclosure). We used community-weighted mean plant functional traits to elucidate plant community changes induced by grazing through time. We also investigated the effects of this experimental treatment on insect pollinated plants and on pollinator habitat use. 3. The grassland community exerted a mixed tolerance/avoidance response to grazing. This resulted in plant functional compositional changes which favoured prostrate plant species with higher specific leaf area, characteristic of ruderal communities. 4. Plant species richness was higher in grazed compared to ungrazed areas. Butterfly and bumblebee habitat use, as well as feeding and resting activities were also higher in grazed areas. Moreover, the number of pollinators increased with plant species richness. 5. Synthesis and applications. This study demonstrates that, to enhance the diversity of a given herbivore community with ecological replacements of extinct wild horses can have significant effects on the functional composition of grasslands. It can also mitigate plant species declines, in particular bee-dependent plants, and boost pollinator habitat use. Novel management alternatives are urgently needed to reverse the negative effect of land abandonment in European agricultural landscapes. Thus, rewilding interventions with large mammalian herbivores may offset current biodiversity declines by maintaining important functional links between plants and pollinators in grassland ecosystems.