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.

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. 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 ofFagus sylvaticaandPinus sylvestris(63 plots). These sites were distributed across the current distribution of these species within Europe. 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. The mixing effects on light absorption were only correlated with the mixing effects on growth forP. sylvestris, suggesting that the mixing effects on this species were driven by the light‐related interactions, whereas mixing effects onF. sylvaticaor whole community growth were probably driven by non‐light‐related interactions. 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.

Aim of study: We aim at (i) developing a reference definition of mixed forests in order to harmonize comparative research in mixed forests and (ii) review the research perspectives in mixed forests.Area of study: The definition is developed in Europe but can be tested worldwide.Material and Methods: Review of existent definitions of mixed forests based and literature review encompassing dynamics, management and economic valuation of mixed forests.Main results: A mixed forest is defined as a forest unit, excluding linear formations, where at least two tree species coexist at any developmental stage, sharing common resources (light, water, and/or soil nutrients). The presence of each of the component species is normally quantified as a proportion of the number of stems or of basal area, although volume, biomass or canopy cover as well as proportions by occupied stand area may be used for specific objectives. A variety of structures and patterns of mixtures can occur, and the interactions between the component species and their relative proportions may change over time.The research perspectives identified are (i) species interactions and responses to hazards, (ii) the concept of maximum density in mixed forests, (iii) conversion of monocultures to mixed-species forest and (iv) economic valuation of ecosystem services provided by mixed forests.Research highlights: The definition is considered a high-level one which encompasses previous attempts to define mixed forests. Current fields of research indicate that gradient studies, experimental design approaches, and model simulations are key topics providing new research opportunities.Keywords: COST Action; EuMIXFOR; mixed-species forests; admixtures of species.