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 Communicated by Peter Biber. 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. Electronic supplementary material

Aim of the study: This paper presents the most appropriate ways to estimate the species proportions by area in mixed stands of Norway spruce (Picea abies L. Karst.) and European beech (Fagus sylvatica L.) by comparing stand level and individual tree level approaches. It also investigates whether different ways of describing species proportions by area can result in different judgments on the over-or under-yielding of species in mixtures.Area of the study: Three triplets of pure and mixed stands of Norway spruce and European beech in three locations in the northeast of Austria are investigated. The three locations differ considerably in slope, bedrock and soil type as well as in site index.Material and methods: In all 9 plots the coordinates of all trees, their dbh, height, height to the crown base and five year increment were measured. The potentially available areas of individual trees are calculated by Voronoi-diagrams and potential densities are estimated from the comparable pure stands, yield tables, and published equations for maximum basal area and Reineke's maximum density line.Main results: The species proportions estimated by the individual tree approach with leaf area as growth characteristic gave the best fit with the stand approach with the most appropriate, regional maximum basal area equations. By using various definitions of species proportions, in the worst case the mixing effects on individual species can be seriously over-or underestimated while the mixing effects on the total increment is only negligibly affected. Research highlights:-Measures of species proportions by area are needed for comparing growth per hectare of a species in a mixed stand with that of the same species in a pure stand.-Species proportions at the stand level are based on estimates of the species' potential densities, either in terms of maximum basal area or of maximum stand density index.-Species proportions at the tree level are derived from the area potentially available (APA) to the individual trees, based on the coordinates of trees in the stands, and on their growth characteristics, such as crown projection area or leaf area.-For the examples of Norway spruce -European beech stands, the species proportions derived according to the individual tree approach using leaf area as growth characteristics fits best with the stand approach using the most appropriate maximum basal area equations

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 Forest Res (2017) 136:171-183 DOI 10.1007/s10342-016-1017 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.Eur J

& 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/ ge orche str a/?uuid=b3e0 98 ca-e 68 1-49 10 -909 9-0e25d3b4cd52&hl=eng.

Key messageThe specific leaf area of European larch depends on branch height and canopy depth, indicating that both, the effect of hydraulic limitations and low water potentials in greater branch heights, and light availability affect specific leaf area.AbstractSpecific leaf area (SLA) is defined as the ratio between projected leaf area and needle dry mass. It often serves as parameter in ecosystem modelling as well as indicator for potential growth rate. We explore the SLA of European larch (Larix decidua) and the most important factors which have an influence on it. Data were collected from eight stands in Styria, Austria. The stands varied in age, elevation and species mixture. Four stands were pure larch stands with only minor proportions of Norway spruce (Picea abies), whereas the other four were mixed stands of larch and spruce. In each stand 15 representative sample trees were felled. The crown of each sample tree was divided into three sections of equal length and in each section a random sample of needles was taken for determining projected leaf area and dry mass of 50 needles. The mean SLA of larch was established to be 117 cm2 g−1 with a standard deviation of ±27.9 cm2 g−1. SLA varies within the crown, but neither between different mixtures nor years of observation nor social position of the trees. A mixed-effects model, with the plots as random effect, revealed that SLA of larch decreased with increasing branch height (p = 0.0012) and increased with increasing canopy depth (p = 0.029). We conclude that both the hydraulic limitations due to low water potentials in greater branch heights and light availability affect specific leaf area.