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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., The networking in this study has been supported by the COST Action FP1206 EUMIXFOR. All authors thank the numerous persons who contributed to the fieldwork in the different sites across Europe. We acknowledge the E-OBS dataset from the EU-FP6 project ENSEMBLES (http://ensembles-eu.metoffice.com) and the data providers in the ECA&D project (http://www.ecad.eu).

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 nutrient-related mechanisms associated with complementarity and points towards the need to further expand the existing frameworks to account for the multivariate nature of tree nutrition., The main autor obtained a PhD grant from the ‘Fonds National de la Recherche Scientifique’ (FNRS-FRIA) and additional funding support from the Walloon forest service (Service Public de Wallonie—Département de la Nature et des Forêts) through the 5 year research programme “Accord cadre de recherches et de vulgarisation forestières”.

1. When tree‐species mixtures are more productive than monocultures, higher light absorption is often suggested as a cause. However, few studies have quantified this effect and even fewer have examined which light‐related interactions are most important, such as the effects of species interactions on tree allometric relationships and crown architecture, differences in vertical or horizontal canopy structure, phenology of deciduous species or the mixing effects on tree size and stand density. 2. In this study, measurements of tree sizes and stand structures were combined with a detailed tree‐level light model (Maestra) to examine the contribution of each light‐related interaction on tree‐ and stand‐level light absorption at 21 sites, each of which contained a triplet of plots including a mixture and monocultures of Fagus sylvatica and Pinus sylvestris (63 plots). These sites were distributed across the current distribution of these species within Europe. 3. Averaged across all sites, the light absorption of mixtures was 14% higher than the mean of the monocultures. At the whole community level, this positive effect of mixing on light absorption increased as canopy volume or site productivity increased, but was unrelated to climate. At the species population or individual tree levels, the mixing effect on light absorption resulted from light‐related interactions involving vertical canopy structure, stand density, the presence of a deciduous species (F. sylvatica), as well as the effects of mixing on tree size and allometric relationships between diameter and height, crown diameter and crown length. 4. The mixing effects on light absorption were only correlated with the mixing effects on growth for P. sylvestris, suggesting that the mixing effects on this species were driven by the light‐related interactions, whereas mixing effects on F. sylvatica or whole community growth were probably driven by non‐light‐related interactions. 5. Synthesis. The overall positive effect of mixing on light absorption was the result of a range of light‐related interactions. However, the relative importance of these interactions varied between sites and is likely to vary between other species combinations and as stands develop. Czech Republic Ministry of Education, Youth and Sports. Grant Number: COST CZ ‐ LD14063 and and LD14074; COST (European Cooperation in Science and Technology). Grant Number: COST Action EuMIXFOR; German Research Foundation. Grant Number: FO 791/4‐1

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.;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. © 2016 The Authors. Journal of Ecology © 2016 British Ecological Society

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-e681-4910-90990e25d3b4cd52&hl=eng.; 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-e681-4910-90990e25d3b4cd52&hl=eng.

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. The networking in this study has been supported by COST Action FP1206 EUMIXFOR. All contributors thank their national funding institutions to establish, measure, and analyze data from the triplets. The first author thanks the Austrian Science Fund, which supported his work under project number P24433-B16. We also want to thank the two anonymous reviewers for their constructive criticism.