A challenge in continuous cover forestry is converting even-aged stands to uneven-aged stands, which can be achieved by releasing advance regeneration through partial harvesting. Partial harvesting increases light and reduces competition, but its effects on oak seedling growth depend on harvesting intensity, direction, understory vegetation, and browsing pressure. Here, we conducted a three-year experiment (2016-2019) in central France, applying partial harvesting with varying intensities (7-30% of the initial basal area) within an 18 m radius around 217 naturally regenerated sessile oak (Quercus petraea (Matt.) Liebl.) seedlings in fenced and unfenced plots. We monitored seedling height growth, understory vegetation, and browsing impacts. Results showed that south-side harvesting uniquely influenced seedling growth, with positive effects only above a threshold of understory vegetation cover and for taller seedlings. Browsing exclusion further enhanced growth. Understory vegetation cover remained unaffected by partial harvesting, possibly due to drought conditions. These findings suggest that targeted south-side harvesting, combined with assessing seedling height and understory competition, can improve oak regeneration. Still forest managers should consider climatic conditions and may need complementary understory interventions to optimize outcomes.

Severe droughts and heat waves have increased over the last decades, leading to forest dieback worldwide. In this context, reducing stand density and mixing tree species are two silvicultural options being investigated by forest managers to limit the impact of climate change on forest health and productivity. We studied tree size effect and stand composition and density effects on crown defoliation in adult trees belonging to two species: sessile oak (Quercus petraea, Matt. (Liebl.)) and Scots pine (Pinus sylvestris, L.). We monitored tree crown defoliation at 18 experimental plots located in central France for five consecutive years following the extreme heat wave of 2019. Our results highlight two different dynamics for crown defoliation in the two tree species. For Scots pine, we observed an abrupt severe increase in defoliation followed by a significant decrease in 2024, while sessile oak, crown defoliation continued to increase steadily after 2020. For both species, larger trees had significantly less crown defoliation than small trees. We found higher defoliation for sessile oak in mixtures and conversely, higher defoliation for Scots pine in pure stands. Finally, we found no difference in crown defoliation between lowdensity and medium-density stands for either species.

European forests have experienced several droughts and heat waves in recent years leading to an increasing level of tree dieback. Quantifying the effects of dieback on tree growth is a crucial issue for forest managers. We monitored tree crown defoliation and tree growth for sessile oak and Scots pine from 2020 to 2023 in the OPTMix experimental network located in central France. We quantified the relationship between tree growth and tree crown defoliation under varying neighborhood conditions. We hypothesized that crown defoliation, tree size and neighborhood competition (local density and composition) would all affect tree growth. We carried out a magnitude analysis to quantify the importance of these different effects on tree growth. We showed that crown defoliation had a strong negative effect for sessile oak (-19.5 % of growth from 42 % to 57 % of defoliation) and a very strong negative effect for Scots pine (-29.5 % of growth from 39 % to 51 % of defoliation). Tree size and neighborhood density had the strongest effects on tree growth for both species, with, respectively, a positive and a negative effect. The proportion of heterospecific species in the neighborhood had the weakest effect, and was positive for sessile oak and negative for Scots pine. We found no interaction between crown defoliation and other predictors, thus indicating that they do not modify the relationship between crown defoliation and radial growth for either species. Forest managers can use defoliation as a rapid indicator of tree health and to help them assess the impact of defoliation on tree growth in various oak-pine forests.

Aim: Climate is a major driver of large-scale variability in biodiversity, as a likely result of more intense biotic interactions under warmer conditions. This idea fuelled decades of research on plant-herbivore interactions, but much less is known about higher-level trophic interactions. We addressed this research gap by characterizing both bird diversity and avian predation along a climatic gradient at the European scale. Location: Europe. Taxon: Insectivorous birds and pedunculate oaks. Methods: We deployed plasticine caterpillars in 138 oak trees in 47 sites along a 19 degrees latitudinal gradient in Europe to quantify bird insectivory through predation attempts. In addition, we used passive acoustic monitoring to (i) characterize the acoustic diversity of surrounding soundscapes; (ii) approximate bird abundance and activity through passive acoustic recordings; and (iii) infer both taxonomic and functional diversity of insectivorous birds from recordings. Results: The functional diversity of insectivorous birds increased with warmer climates. Bird predation increased with forest cover and bird acoustic activity but decreased with mean annual temperature and functional richness of insectivorous birds. Contrary to our predictions, climatic clines in bird predation attempts were not directly mediated by changes in insectivorous bird diversity or acoustic activity, but climate and habitat still had independent effects on predation attempts. Main Conclusions: Our study supports the hypothesis of an increase in the diversity of insectivorous birds towards warmer climates but refutes the idea that an increase in diversity would lead to more predation and advocates for better accounting for activity and abundance of insectivorous birds when studying the large-scale variation in insect-tree interactions.

Adapting forests to climate change is a major challenge for forest ecology and forestry. Among the management options available, encouraging the use of mixtures is a promising way forward. However, this practice requires a thorough understanding of how species respond to mixing. In this article, we analyzed species dominant height responds to mixing and how species ontogeny and traits drive this response. We compared species observed dominant height in mixed even-aged stands with the expected dominant height of the same species in a monospecific stand under the same environmental conditions. We then related this dominant height variation due to mixing to between-species dominant height difference and to species traits linked to competition (shade tolerance, wood density, specific leaf area). We focused our analyses on 76 pairs of forest tree species. We used data from the French National Forest Inventory to calculate species dominant height in 1368 mixed stands. We then used previously developed models to estimate the expected dominant height in virtual monospecific stands with the same environmental conditions. We found that mixture had a significant impact on species dominant height for 15 out of 50 species-combination considered. Dominant height of a given species was higher in mixture than in pure stands when this species had a lower dominant height in pure stands, a lower shade tolerance, a lower specific leaf area or a higher wood density than its companion species. Our results suggest that species dominant height response to mixing depends on how mixture influences the competition for light. Our results will help inform strategies aiming to diversify species in forests, and will be especially useful in anticipating a given species' behavior in response to competition for light when it is mixed with other species.

The European biodiversity and forest strategies rely on forest sustainable management (SFM) to conserve forest biodiversity. However, current sustainability assessments hardly account for direct biodiversity indicators. We focused on forest multi-taxon biodiversity to: i) gather and map the existing information; ii) identify knowledge and research gaps; iii) discuss its research potential. We established a research network to fit data on species, standing trees, lying deadwood and sampling unit description from 34 local datasets across 3591 sampling units. A total of 8724 species were represented, with the share of common and rare species varying across taxonomic classes: some included many species with several rare ones (e.g., Insecta); others (e.g., Bryopsida) were repre-sented by few common species. Tree-related structural attributes were sampled in a subset of sampling units (2889; 2356; 2309 and 1388 respectively for diameter, height, deadwood and microhabitats). Overall, multi-taxon studies are biased towards mature forests and may underrepresent the species related to other develop-mental phases. European forest compositional categories were all represented, but beech forests were over-represented as compared to thermophilous and boreal forests. Most sampling units (94%) were referred to a habitat type of conservation concern. Existing information may support European conservation and SFM stra-tegies in: (i) methodological harmonization and coordinated monitoring; (ii) definition and testing of SFM in-dicators and thresholds; (iii) data-driven assessment of the effects of environmental and management drivers on multi-taxon forest biological and functional diversity, (iv) multi-scale forest monitoring integrating in-situ and remotely sensed information.

The increasing disturbances in monocultures around the world are testimony to their instability under global change. Many studies have claimed that temporal stability of productivity increases with species richness, although the ecological fundamentals have mainly been investigated through diversity experiments. To adequately manage forest ecosystems, it is necessary to have a comprehensive understanding of the effect of mixing species on the temporal stability of productivity and the way in which it is influenced by climate conditions across large geographical areas. Here, we used a unique dataset of 261 stands combining pure and two-species mixtures of four relevant tree species over a wide range of climate conditions in Europe to examine the effect of species mixing on the level and temporal stability of productivity. Structural equation modelling was employed to further explore the direct and indirect influence of climate, overyielding, species asynchrony and additive effect (i.e. temporal stability expected from the species growth in monospecific stands) on temporal stability in mixed forests. We showed that by adding only one tree species to monocultures, the level (overyielding: +6%) and stability (temporal stability: +12%) of stand growth increased significantly. We identified the key effect of temperature on destabilizing stand growth, which may be mitigated by mixing species. We further confirmed asynchrony as the main driver of temporal stability in mixed stands, through both the additive effect and species interactions, which modify between-species asynchrony in mixtures in comparison to monocultures. Synthesis and applications. This study highlights the emergent properties associated with mixing two species, which result in resource efficient and temporally stable production systems. We reveal the negative impact of mean temperature on temporal stability of forest productivity and how the stabilizing effect of mixing two species can counterbalance this impact. The overyielding and temporal stability of growth addressed in this paper are essential for ecosystem services closely linked with the level and rhythm of forest growth. Our results underline that mixing two species can be a realistic and effective nature-based climate solution, which could contribute towards meeting EU climate target policies.

Heterogeneity of structure can increase mechanical stability, stress resistance and resilience, biodiversity and many other functions and services of forest stands. That is why many silvicultural measures aim at enhancing structural diversity. However, the effectiveness and potential of structuring may depend on the site conditions. Here, we revealed how the stand structure is determined by site quality and results from site-dependent partitioning of growth and mortality among the trees. We based our study on 90 mature, even-aged, fully stocked monocultures of Scots pine (Pinus sylvestris L.) sampled in 21 countries along a productivity gradient across Europe. A mini-simulation study further analyzed the site-dependency of the interplay between growth and mortality and the resulting stand structure. The overarching hypothesis was that the stand structure changes with site quality and results from the site-dependent asymmetry of competition and mortality. First, we show that Scots pine stands structure across Europe become more homogeneous with increasing site quality. The coefficient of variation and Gini coefficient of stem diameter and tree height continuously decreased, whereas Stand Density Index and stand basal area increased with site index. Second, we reveal a site-dependency of the growth distribution among the trees and the mortality. With increasing site index, the asymmetry of both competition and growth distribution increased and suggested, at first glance, an increase in stand heterogeneity. However, with increasing site index, mortality eliminates mainly small instead of all-sized trees, cancels the size variation and reduces the structural heterogeneity. Third, we modelled the site-dependent interplay between growth partitioning and mortality. By scenario runs for different site conditions, we can show how the site-dependent structure at the stand level emerges from the asymmetric competition and mortality at the tree level and how the interplay changes with increasing site quality across Europe. Our most interesting finding was that the growth partitioning became more asymmetric and structuring with increasing site quality, but that the mortality eliminated predominantly small trees, reduced their size variation and thus reversed the impact of site quality on the structure. Finally, the reverse effects of mode of growth partitioning and mortality on the stand structure resulted in the highest size variation on poor sites and decreased structural heterogeneity with increasing site quality. Since our results indicate where heterogeneous structures need silviculture interventions and where they emerge naturally, we conclude that these findings may improve system understanding and modelling and guide forest management aiming at structurally rich forests.

The translation of genetic gains into economic gains is important for evaluating the impact of using genetically improved forest reproductive material (FRM) in the forest sector. However, few studies based on European forests have been published to date. Here, we analyse the stand-level wood production and financial performance of planting genetically improved FRM in even-aged planted forests focusing on four European case studies with advanced breeding programme material and different management contexts: Scots pine (Pinus sylvestris L.) in southern Finland, central Sweden and central France, and maritime pine (Pinus pinaster Ait.) in southwestern France. The growth of improved stands was simulated using species-specific growth models by incorporating two levels of expected genetic gains (present and next generations of seed orchards, varying from 7 to 40 per cent depending on the breeding programme) into the estimated mean annual volume increment over a rotation (m3 ha−1 yr−1). For each level of genetic gain, we tested the plantation of improved FRM managed with two silvicultural scenarios (maintaining the standard baseline rotation and thinning regime vs shorter rotation through the earlier achievement of the recommended felling criteria) in comparison with the plantation of the reference unimproved material (absence of genetic gain) managed according to the standard silvicultural regime. The use of improved FRM resulted in a larger financial performance in terms of soil expectation value (SEV € ha−1, discount rate 3 per cent) than planting unimproved reference material in all case studies and silvicultural scenarios for different wood price contexts (SEV gain from +20 to +190 per cent depending on the genetic and silvicultural context). The challenges associated with the economic assessment of realized gains from genetically improved FRM are discussed. We argue that silvicultural guidelines should be adapted to the use of improved FRM in order to gain better financial performance and flexible silvicultural response of planted forests to future environmental and socio-economic changes.

A growing body of research suggests mixed-species stands are generally more productive than pure stands as well as less sensitive to disturbances. However, these effects of mixture depend on species assemblages and environmental conditions. Here, we present the Salem simulator, a tool that can help forest managers assess the potential benefit of shifting from pure to mixed stands from a productivity perspective. Salem predicts the dynamics of pure and mixed even-aged stands and makes it possible to simulate management operations. Its purpose is to be a decision support tool for forest managers and stakeholders as well as for policy makers. It is also designed to conduct virtual experiments and help answer research questions. In Salem, we parameterised the growth in pure stand of 12 common tree species of Europe and we assessed the effect of mixture on species growth for 24 species pairs (made up of the 12 species mentioned above). Thus, Salem makes it possible to compare the productivity of 36 different pure and mixed stands depending on environmental conditions and user-defined management strategies. Salem is essentially based on the analysis of National Forest Inventory data. A major outcome of this analysis is that we found species mixture most often increases species growth, in particular at the poorest sites. Independently from the simulator, foresters and researchers can also consider using the species-specific models that constitute Salem: the growth models including or excluding mixture effect, the bark models, the diameter distribution models, the circumference-height relationship models, as well as the volume equations for the 12 parameterised species. Salem runs on Windows, Linux, or Mac. Its user-friendly graphical user interface makes it easy to use for non-modellers. Finally, it is distributed under a LGPL license and is therefore free and open source.

Climate change is impacting temperate tree species phenology, especially the timing of budburst, which is mainly driven by air temperature. However, interactions with biotic or other environmental factors also influence the timing of budburst and are usually overlooked. We studied the influence of forest stand composition on the budburst date of adult trees belonging to two species: sessile oak (Quercus petraea, Matt. (Liebl.)) and Scots pine (Pinus sylvestris, L.). We monitored their budburst dates for seven consecutive years at 18 experimental plots located in central France. We compared the budburst dates of oaks and pines growing in monospecific stands with those of their counterparts in an oak-pine mixture. Our results show that sessile oak budburst date in mixed stands with Scots pine was delayed by 2.2 days on average (SE = 0.6) compared to its budburst date in monospecific stands. In years with early budburst, the delay was more pronounced - up to four days. For Scots pine, our results showed no difference between budburst dates in monospecific and mixed stands. We hypothesize that the persistent foliage of the Scots pine in the mixed stand intercepted a part of the solar radiation, which affected the temperature perceived by the oak buds, thereby delaying the heat accumulation needed for sessile oak budburst. This effect may be of interest for the management of sessile oak in the context of global warming. In the future, sessile oak may experience more frequent frost damage due to an earlier budburst. Managing sessile oak with an evergreen species could limit late frost damage to some extent by delaying budburst. Stand composition must obviously be taken into account when monitoring the phenology of temperate tree species and to enable robust comparisons of phenological events for a given tree species at different sites.

A growing body of research suggests mixed-species stands are generally more productive than pure stands as well as less sensitive to disturbances. However, these effects of mixture depend on species assemblages and environmental conditions. Here, we present the Salem simulator, a tool that can help forest managers assess the potential benefit of shifting from pure to mixed stands from a productivity perspective. Salem predicts the dynamics of pure and mixed even-aged stands and makes it possible to simulate management operations. Its purpose is to be a decision support tool for forest managers and stakeholders as well as for policy makers. It is also designed to conduct virtual experiments and help answer research questions. In Salem, we parameterised the growth in pure stand of 12 common tree species of Europe and we assessed the effect of mixture on species growth for 24 species pairs (made up of the 12 species mentioned above). Thus, Salem makes it possible to compare the productivity of 36 different pure and mixed stands depending on environmental conditions and user-defined management strategies. Salem is essentially based on the analysis of National Forest Inventory data. A major outcome of this analysis is that we found species mixture most often increases species growth, in particular at the poorest sites. Independently from the simulator, foresters and researchers can also consider using the species-specific models that constitute Salem: the growth models including or excluding mixture effect, the bark models, the diameter distribution models, the circumference-height relationship models, as well as the volume equations for the 12 parameterised species. Salem runs on Windows, Linux, or Mac. Its user-friendly graphical user interface makes it easy to use for non-modellers. Finally, it is distributed under a LGPL license and is therefore free and open source.

The leaf area index (LAI) is a key characteristic of forest stand aboveground net productivity (ANP), and many methods have been developed to estimate the LAI. However, every method has flaws, e.g., methods may be destructive, require means or time and/or show intrinsic bias and estimation errors. A relationship using basal area (G) and stand age to estimate LAI was proposed by Sonohat et al. (2004). We used literature data in addition to data form measurements campaign made in the northern half of France to build a data set with large ranges of pedoclimatic conditions, stand age and measured LAI. We validated the Sonohat et al. (2004) relationship and attempted to improve or modify it using other stand/dendrometric characteristics that could be predictors of the LAI. The result is a series of three models using the G, age and/or quadratic mean diameter (Dg), and the models were able to estimate the LAI of an oak only even-aged forest stand with good confidence (root mean square error, RMSE < 0.75) While G is the main predictor here, age and Dg could be used conjointly or exclusively given the available data, with variable precision in the estimations. Although these models could not, by construction, relate to the interannual variability of the LAI, they may provide the theoretical LAI of an untouched forest (no meteorological, biotic or anthropogenic perturbation) in recent years. additionally, the use of this model may be more interesting than an LAI measurement campaign, depending on the means to be invested in such a campaign.

Positive productivity-diversity relationships, pest-effect mitigation and increased resilience and stability maintain an ongoing interest for mixed stands in forestry. However, how mixing species affects forest regeneration is yet to be further explored. We used data from the French National Forest Inventory (from 2006 to 2016) to model Quercus petraea (Matt.) Liebl regeneration cover in pure and mixed Quercus petraea stands; we included the effects of abiotic and biotic factors as well as mixture. We hypothesized that the characteristics of the companion species would prevent or facilitate the regeneration of oak. Quercus petraea regeneration cover in this study responded negatively to total canopy cover and herbivory pressure. Mean July potential evapotranspiration (PET), mean December maximal temperature and soil pH are variables whose spatial variations over a given territory structure regeneration cover; all three of these variables have optimum values. Quercus petraea regeneration cover is linked to the proportion of Quercus petraea in the canopy layer in all mixed stands, except when the oak is mixed with shade-intolerant conifers: in this case, Quercus petraea regeneration is enhanced. The shade tolerance of admixed broadleaved species did not affect the Quercus petraea regeneration. This suggests that oak regeneration was facilitated with a shade-intolerant coniferous companion species due to better light transmittance through the crown or the competitive advantage of Quercus petraea over coniferous shade-intolerant species. These results are of interest for oak mixtures since Quercus petraea regeneration cover benefits from mixtures with shade-intolerant conifers and is at least equal to that of pure stands.