Many studies show that mixed species stands can have higher gross growth, or so-called overyielding, compared with monocultures. However, much less is known about mortality in mixed stands. Knowledge is lacking, for example, of how much of the gross growth is retained in the standing stock and how much is lost due to mor-tality. Here, we addressed this knowledge gap of mixed stand dynamics by evaluating 23 middle-aged, unthinned triplets of monospecific and mixed plots of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) repeatedly surveyed over 6-8 years throughout Europe. For explanation of technical terms in this abstract see Box 1.First, mixed stands produced more gross growth (+10%) but less net growth (-28%) compared with the weighted mean growth of monospecific stands. In monospecific stands, 73% of the gross growth was accumu-lated in the standing stock, whereas only 48% was accumulated in mixed stands. The gross overyielding of pine (2%) was lower than that of beech (18%). However, the net overyielding of beech was still 10%, whereas low growth and dropout of pine caused a substantial reduction from gross to net growth.Second, the mortality rates, the self-and alien-thinning strength, and the stem volume dropout were higher in mixed stands than monospecific stands. The main reason was the lower survival of pine, whereas beech persisted more similarly in mixed compared with monospecific stands.Third, we found a 10% higher stand density in mixed stands compared with monospecific stands at the first survey. This superiority decreased to 5% in the second survey.Fourth, the mixing proportion of Scots pine decreased from 46% to 44% between the first and second survey. The more than doubling of the segregation index (S) calculated by Pielou index (S increased from 0.2 to 0.5), indicated a strong tendency towards demixing due to pine.Fifth, we showed that with increasing water supply the dropout fraction of the gross growth in the mixture slightly decreased for pine, strongly increased for beech, and also increased for the stand as a whole. We discuss how the reduction of inter-specific competition by thinning may enable a continuous benefit of diversity and overyielding of mixed compared with monospecific stands of Scots pine and European beech.

Process-based models and empirical modelling techniques are frequently used to (i) explore the sensitivity of tree growth to environmental variables, and (ii) predict the future growth of trees and forest stands under climate change scenarios. However, modelling approaches substantially influence predictions of the sensitivity of trees to environmen-tal factors. Here, we used tree-ring width (TRW) data from 1630 beech trees from a network of 70 plots established across European mountains to build empirical predictive growth models using various modelling approaches. In addi-tion, we used 3-PG and Biome-BGCMuSo process-based models to compare growth predictions with derived empirical models. Results revealed similar prediction errors (RMSE) across models ranging between 3.71 and 7.54 cm2 of basal area increment (BAI). The models explained most of the variability in BAI ranging from 54 % to 87 %. Selected explan-atory variables (despite being statistically highly significant) and the pattern of the growth sensitivity differed between models substantially. We identified only five factors with the same effect and the same sensitivity pattern in all empir-ical models: tree DBH, competition index, elevation, Gini index of DBH, and soil silt content. However, the sensitivity to most of the climate variables was low and inconsistent among the empirical models. Both empirical and process -based models suggest that beech in European mountains will, on average, likely experience better growth conditions under both 4.5 and 8.5 RCP scenarios. The process-based models indicated that beech may grow better across European mountains by 1.05 to 1.4 times in warmer conditions. The empirical models identified several drivers of tree growth that are not included in the current process-based models (e.g., different nutrients) but may have a sub-stantial effect on final results, particularly if they are limiting factors. Hence, future development of process-based models may build upon our findings to increase their ability to correctly capture ecosystem dynamics.

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.

Increasing species diversity is considered a promising strategy to mitigate the negative impacts of global change on forests. However, the interactions between regional climate conditions and species-mixing effects on climate-growth relationships and drought resistance remain poorly documented. In this study, we investigated the patterns of species-mixing effects over a large gradient of environmental conditions throughout Europe for European beech (Fagus sylvatica L.) and Scots pine (Pinus sylvestris L.), two species with contrasted ecological traits. We hypothesized that across large geographical scales, the difference of climate-growth relationships and drought resistance between pure and mixed stands would be dependent on regional climate. We used tree ring chronologies derived from 1143 beech and 1164 pine trees sampled in 30 study sites, each composed of one mixed stand of beech and pine and of the two corresponding pure stands located in similar site conditions. For each site and stand, we used Bootstrapped Correlation Coefficients (BCCs) on standardized chronologies and growth reduction during drought years on raw chronologies to analyze the difference in climate-tree growth relationships and resistance to drought between pure and mixed stands. We found consistent large-scale spatial patterns of climate-growth relationships. Those patterns were similar for both species. With the exception of the driest climates where pure and mixed beech stands tended to display differences in growth correlation with the main climatic drivers, the mixing effects on the BCCs were highly variable, resulting in the lack of a coherent response to mixing. No consistent species-mixing effect on drought resistance was found within and across climate zones. On average, mixing had no significant effect on drought resistance for neither species, yet it increased pine resistance in sites with higher climatic water balance in autumn. Also, beech and pine most often differed in the timing of their drought response within similar sites, irrespective of the regional climate, which might increase the temporal stability of growth in mixed compared to pure stands. Our results showed that the impact of species mixing on tree response to climate did not strongly differ between groups of sites with distinct climate characteristics and climate-growth relationships, indicating the interacting influences of species identity, stand characteristics, drought events characteristics as well as local site conditions.

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.

Recent studies show that several tree species are spreading to higher latitudes and elevations due to climate change. European beech, presently dominating from the colline to the subalpine vegetation belt, is already present in upper montane subalpine forests and has a high potential to further advance to higher elevations in European mountain forests, where the temperature is predicted to further increase in the near future. Although essential for adaptive silviculture, it remains unknown whether the upward shift of beech could be assisted when it is mixed with Norway spruce or silver fir compared with mono-specific stands, as the species interactions under such conditions are hardly known. In this study, we posed the general hypotheses that the growth depending on age of European beech in mountain forests was similar in mono-specific and mixed-species stands and remained stable over time and space in the last two centuries. The scrutiny of these hypotheses was based on increment coring of 1240 dominant beech trees in 45 plots in mono-specific stands of beech and in 46 mixed mountain forests. We found that (i) on average, mean tree diameter increased linearly with age. The age trend was linear in both forest types, but the slope of the age–growth relationship was higher in mono-specific than in mixed mountain forests. (ii) Beech growth in mono-specific stands was stronger reduced with increasing elevation than that in mixed-species stands. (iii) Beech growth in mono-specific stands was on average higher than beech growth in mixed stands. However, at elevations > 1200 m, growth of beech in mixed stands was higher than that in mono-specific stands. Differences in the growth patterns among elevation zones are less pronounced now than in the past, in both mono-specific and mixed stands. As the higher and longer persisting growth rates extend the flexibility of suitable ages or size for tree harvest and removal, the longer-lasting growth may be of special relevance for multi-aged silviculture concepts. On top of their function for structure and habitat improvement, the remaining old trees may grow more in mass and value than assumed so far.

Tree species-mixing has been suggested as one option to counteract the adverse effects of global change on tree mineral nutrition, yet the effect of mixing on nutrient availability remains poorly documented. We therefore analyzed the current foliar nutrient (N, P, K, Ca, Mg) quantities and ilr balances (isometric log transformed ratios between elements or groups of elements) for 261 European beech and 248 Scots pine trees from 15 sites, each consisting of one beech-pine mixed stand and the respective monocultures, across a gradient of environmental conditions in Europe. We hypothesized an overall positive effect of mixing on tree foliar nutrient content, and that this mixing effect would be stronger on nutrient-poor sites. Using linear mixed models and multivariate linear regression models, we first tested for the effects of species (beech/pine) and composition (pure/mixed) across all sites; we then investigated whether the species-mixing effect was related to site fertility. The nutrient composition of beech leaves and pine needles differed significantly for all ilr balances. For both species, significant mixing effects were detected for some nutrients and ilr balances; those effects, however, could not be consistently related to contrasted nutrient composition between species. For most nutrients and ilr balances, the mixing effect was influenced by the site nutritional status, but the pattern differed from expectation: absence or minor differences between monocultures and mixtures at the lower end of the chemical fertility gradient, and maximum differences in rich soils. The contrasting foliar nutrient composition of pine and beech trees and the site nutrient status only partly explained the mixing effects on tree mineral nutrition. Our results claim for a better understanding of nutrientrelated mechanisms associated with complementarity and points towards the need to further expand the existing frameworks to account for the multivariate nature of tree nutrition.

Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) dominate many of the European forest stands. Also, mixtures of European beech and Scots pine more or less occur over all European countries, but have been scarcely investigated. The area occupied by each species is of high relevance, especially for growth evaluation and comparison of different species in mixed and monospecific stands. Thus, we studied different methods to describe species proportions and their definition as proportion by area. 25 triplets consisting of mixed and monospecific stands were established across Europe ranging from Lithuania to Spain in northern to southern direction and from Bulgaria to Belgium in eastern to western direction. On stand level, the conclusive method for estimating the species proportion as a fraction of the stand area relates the observed density (tree number or basal area) to its potential. This stand-level estimation makes use of the potential from comparable neighboring monospecific stands or from maximum density lines derived from other data, e.g. forest inventories or permanent observations plots. At tree level, the fraction of the stand area occupied by a species can be derived from the proportions of their crown projection area or of their leaf area. The estimates of the potentials obtained from neighboring monospecific stands, especially in older stands, were poorer than those from the maximum density line depending on the Martonne aridity index. Therefore, the stand-level method in combination with the Martonne aridity index for potential densities can be highly recommended. The species' proportions estimated with this method are best approximated by the proportions of the species' leaf areas. In forest practice, the most commonly applied method is an ocular estimation of the proportions by crown projection area. Even though the proportions of pine were calculated here by measuring crown projection areas in the field, we found this method to underestimate the proportion by 25% compared to the stand-level approach.

Summary There is increasing evidence that species diversity enhances the temporal stability (TS) of community productivity in different ecosystems, although its effect at the population and tree levels seems to be negative or neutral. Asynchrony in species responses to environmental conditions was found to be one of the main drivers of this stabilizing process. However, the effect of species mixing on the stability of productivity, and the relative importance of the associated mechanisms, remain poorly understood in forest communities. We investigated the way mixing species influenced the TS of productivity in Pinus sylvestris L. and Fagus sylvatica L. forests, and attempted to determine the main drivers among overyielding, asynchrony between species annual growth responses to environmental conditions, and temporal shifts in species interactions. We used a network of 93 experimental plots distributed across Europe to compare the TS of basal area growth over a 15‐year period (1999–2013) in mixed and monospecific forest stands at different organizational levels, namely the community, population and individual tree levels. Mixed stands showed a higher TS of basal area growth than monospecific stands at the community level, but not at the population or individual tree levels. The TS at the community level was related to asynchrony between species growth in mixtures, but not to overyielding nor to asynchrony between species growth in monospecific stands. Temporal shifts in species interactions were also related to asynchrony and to the mixing effect on the TS. Synthesis. Our findings confirm that species mixing can stabilize productivity at the community level, whereas there is a neutral or negative effect on stability at the population and individual tree levels. The contrasting findings regarding the relationships between the temporal stability and asynchrony in species growth in mixed and monospecific stands suggest that the main driver in the stabilizing process may be the temporal niche complementarity between species rather than differences in species’ intrinsic responses to environmental conditions.

Key message This data set provides unique empirical data from triplets of Scots pine ( Pinus sylvestris L.) and European beech ( Fagus sylvatica L.) across Europe. Dendrometric variables are provided for 32 triplets, 96 plots, 7555 trees and 4695 core samples. These data contribute to our understanding of mixed stand dynamics. Dataset access at http://dx.doi.org/10.5061/dryad.8v04m . Associated metadata available at https://metadata-afs.nancy.inra.fr/geonetwork/apps/georchestra/?uuid=b3e098ca-e681-4910-9099-0e25d3b4cd52&hl=eng .

The mixing of tree species with complementary ecological traits may modify forest functioning regarding productivity, stability, or resilience against disturbances. This may be achieved by a higher heterogeneity in stand structure which is often addressed but rarely quantified. Here, we use 32 triplets of mature and fully stocked monocultures and mixed stands of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) located along a productivity and water availability gradient through Europe to examine how mixing modifies the stand structure in terms of stand density, horizontal tree distribution pattern, vertical stand structure, size distribution pattern, and variation in tree morphology. We further analyze how site conditions modify these aspects of stand structure. For this typical mixture of a light demanding and shade tolerant species we show that (i) mixing significantly increases many aspects of structural heterogeneity compared with monocultures, (ii) mixing effects such as an increase of stand density and diversification of vertical structure and tree morphology are caused by species identity (additive effects) but also by species interactions (multiplicative effects), and (iii) superior heterogeneity of mixed stands over monocultures can increase from dry to moist sites. We discuss the implications for analyzing the productivity, for modelling and for the management of mixed species stands. (C) 2016 Elsevier B.V. All rights reserved.

Mixing of complementary tree species may increase stand productivity, mitigate the effects of drought and other risks, and pave the way to forest production systems which may be more resource-use efficient and stable in the face of climate change. However, systematic empirical studies on mixing effects are still missing for many commercially important and widespread species combinations. Here we studied the growth of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) in mixed versus pure stands on 32 triplets located along a productivity gradient through Europe, reaching from Sweden to Bulgaria and from Spain to the Ukraine. Stand inventory and taking increment cores on the mainly 60–80 year-old trees and 0.02–1.55 ha sized, fully stocked plots provided insight how species mixing modifies the structure, dynamics and productivity compared with neighbouring pure stands. In mixture standing volume (+12 %), stand density (+20 %), basal area growth (+12 %), and stand volume growth (+8 %) were higher than the weighted mean of the neighbouring pure stands. Scots pine and European beech contributed rather equally to the overyielding and overdensity. In mixed stands mean diameter (+20 %) and height (+6 %) of Scots pine was ahead, while both diameter and height growth of European beech were behind (−8 %). The overyielding and overdensity were independent of the site index, the stand growth and yield, and climatic variables despite the wide variation in precipitation (520–1175 mm year−1), mean annual temperature (6–10.5 °C), and the drought index by de Martonne (28–61 mm °C−1) on the sites. Therefore, this species combination is potentially useful for increasing productivity across a wide range of site and climatic conditions. Given the significant overyielding of stand basal area growth but the absence of any relationship with site index and climatic variables, we hypothesize that the overyielding and overdensity results from several different types of interactions (light-, water-, and nutrient-related) that are all important in different circumstances. We discuss the relevance of the results for ecological theory and for the ongoing silvicultural transition from pure to mixed stands and their adaptation to climate change.

Methodological approach to determining an ecological-economic efficiency of reforestation and its approbation for forest enterprises of Maley Polissya (state enterprise “Brody Forestry” and “Radekhiv Forestry and Hunting”), which have typical for the region a forest site quality in terms of climate, hydrology, and soil conditions, are described. To investigate a dynamics of forest stands increment and productivity with respect to the forest stands’ origin (planted and natural) we developed a seven-stage methodological approach: formulation of the problem; creation of a research database; justification of models specification; estimation of regression models parameters; verification of the models; forecasting of a forest stocks growth dynamics and an average increment; formulation of conclusions and recommendations for effective reforestation in vulnerable natural conditions. In order to estimate the dynamics of stocks and average increment of tree species typical to Maley Polissya conditions, the system of models was estimated. Comparative analysis of pine (Pinus sylvestris L.) stands’ dynamics and productivity with respect to their origin confirmed the need to careful consideration of forest conditions and forest types for each forest site. Thus, in a wet forest conditions (the most common site conditions for SE “Brody Forestry”) the difference in the stock of pine stands with different origin is insignificant, while in the fresh forest conditions (richer conditions, which are typical for the SE “Radekhiv Forestry and Hunting”) the stock of artificial stands is much higher in comparison to natural ones. However, taking into consideration a financial cost of stand formation, a total economic value of forest ecosystem services like biodiversity conservation, landscape amenity etc., as well as forest stand resistance makes the choice of reforestation methods more grounded and holistic. From this perspective natural reforestation becomes more attractive whereas artificial reforestation loses its priority.