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.

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

Mixed mountain forests of European beech (Fagus sylvatica L.), Norway spruce (Picea abies (L.) Karst), and silver fir (Abies alba Mill.) cover a total area of more than 10 million hectares in Europe. Due to altitudinal zoning, these forests are particularly vulnerable to climate change. However, as little is known about the long-term development of the productivity and the adaptation and mitigation potential of these forest systems in Europe, reliable information on productivity is required for sustainable forest management. Using generalized additive mixed models this study investigated 60 long-term experimental plots and provides information about the productivity of mixed mountain forests across a variety of European mountain areas in a standardized way for the first time. The average periodic annual volume increment (PAI) of these forests amounts to 9.3 m3ha−1y−1. Despite a significant increase in annual mean temperature the PAI has not changed significantly over the last 30 years. However, at the species level, we found significant changes in the growth dynamics. While beech had a PAI of 8.2 m3ha−1y−1 over the entire period (1980–2010), the PAI of spruce dropped significantly from 14.2 to 10.8 m3ha−1y−1, and the PAI of fir rose significantly from 7.2 to 11.3 m3ha−1y−1. Consequently, we observed stable stand volume increments in relation to climate change.

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.

In Europe, mixed mountain forests, primarily comprised of Norway spruce (Picea abies (L.) Karst.), silver fir (Abies alba Mill.), and European beech (Fagus sylvatica L.), cover about 10 × 10⁶ ha at elevations between ∼600 and 1600 m a.s.l. These forests provide invaluable ecosystem services. However, the growth of these forests and the competition among their main species are expected to be strongly affected by climate warming. In this study, we analyzed the growth development of spruce, fir, and beech in moist mixed mountain forests in Europe over the last 300 years. Based on tree-ring analyses on long-term observational plots, we found for all three species (i) a nondecelerating, linear diameter growth trend spanning more than 300 years; (ii) increased growth levels and trends, the latter being particularly pronounced for fir and beech; and (iii) an elevation-dependent change of fir and beech growth. Whereas in the past, the growth was highest at lower elevations, today’s growth is superior at higher elevations. This spatiotemporal pattern indicates significant changes in the growth and interspecific competition at the expense of spruce in mixed mountain forests. We discuss possible causes, consequences, and silvicultural implications of these distinct growth changes in mixed mountain forests.

The productivity and functioning of mixed-species forests often differs from that of monocultures. However, the magnitude and direction of these differences are difficult to predict because species interactions can be modified by many potentially interacting climatic and edaphic conditions, stand structure and previous management. Process-based forest growth models could potentially be used to disentangle the effects of these factors and thereby improve our understanding of mixed forest functioning while facilitating their design and silvicultural management. However, to date, the predicted mixing effects of forest growth models have not been compared with measured mixing effects. In this study, 26 sites across Europe, each containing a mixture and monocultures of Fagus sylvatica and Pinus sylvestris, were used to calculate mixing effects on growth and yield and compare them with the mixing effects predicted by the forest growth model 3-PG(mix). The climate and edaphic conditions, stand structures and ages varied greatly between sites. The model performed well when predicting the stem mass and total mass (and mixing effects on these components), with model efficiency that was usually > 0.7. The model efficiency was lower for growth or smaller components such as foliage mass and root mass. The model was also used to predict how mixing effects would change along gradients in precipitation, temperature, potential available soil water, age, thinning intensity and soil fertility. The predicted patterns were consistent with measurements of mixing effects from published studies. The 3-PG model is a widely used management tool for monospecific stands and this study shows that 3-PGmb, can be used to examine the dynamics of mixed-species stands and determine how they may need to be managed