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Forest composed of Picea abies L., Abies alba Mill. and Fagus sylvatica L. cover a large area in the European mountain regions and have a high ecological and socio-economic importance as they supply many ecosystems services. Because of climate change, these forests are exposed to warming, and this effect increases with elevation, which may impact their delivery of goods and services. Previous studies did not find significant changes in the overall productivity of these species over the last 30 years, but they observed changes in species competitiveness at the species and tree levels. In this study, we aimed to link previous results on tree, species and stand level growth in spruce-fir-beech mixed mountain forests by analysing species size distribution dynamics under different climate conditions and their effect on stand growth. We developed a matrix model based on data from 76 long-term experimental plots distributed throughout Europe. We used the change in stand basal area to explore whether temperature modifies species size dominances and proportions, whether the temperature effects on changes in species basal area depend on species size dominance, and whether the effect of species size dominance on changes in the stand basal area varies with temperature. Our results showed that annual mean temperature is an important climatic driver of species dynamics in spruce-fir-beech mixed mountain forests, such that stand basal area growth was favored by higher temperatures, particularly due to positive responses of silver fir which were greater than negative effects of temperature on European beech. The high temperatures also favored the size-dominance of silver fir, while European beech tended to have smaller diameters, independent of the temperature. We also found that the identity of the size-dominant species also influenced changes in stand basal area, with the highest or the lowest changes when Norway spruce and European beech were the size-dominant species, respectively. Silver fir was less influenced by the identity of the size-dominant species than by temperature. Therefore, although mixed mountain forests of spruce-fir-beech were found to be resilient systems in terms of stand productivity, we conclude that increasing temperatures may modify species dynamics and consequently silvicultural interventions will be needed to control species proportions and dominances.

Centro de Investigación Forestal (CIFOR), Spruce-fir-beech mixed forests cover a large area in European mountain regions, with high ecological and socio-economic importance. As elevation-zone systems they are highly affected by climate change, which is modifying species growth patterns and productivity shifts among species. The extent to which associated tree species can access resources and grow asynchronously may affect their resistance and persistence under climate change. Intra-specific synchrony in annual tree growth is a good indicator of species specific dependence on environmental conditions variability. However, little attention has been paid to explore the role of the inter-specific growth asynchrony in the adaptation of mixed forests to climate change. Here we used a database of 1790 tree-ring series collected from 28 experimental plots in spruce-fir-beech mixed forests across Europe to explore how spatio-temporal patterns of the intra- and inter-specific growth synchrony relate to climate variation during the past century. We further examined whether synchrony in growth response to inter-annual environmental fluctuations depended on site conditions. We found that the inter-specific growth synchrony was always lower than the intra-specific synchrony, for both high (inter-annual fluctuations) and low frequency (mid- to long-term) growth variation, suggesting between species niche complementarity at both temporal levels. Intra- and inter-specific synchronies in inter-annual growth fluctuations significantly changed along elevation, being greater at higher elevations. Moreover, the climate warming likely induced temporal changes in synchrony, but the effect varied along the elevation gradient. The synchrony strongly intensified at lower elevations likely due to climate warming and drying conditions. Our results suggest that intra- and inter-specific growth synchrony can be used as an indicator of temporal niche complementarity among species. We conclude that spruce-fir-beech mixtures should be preferred against mono-specific forests to buffer climate change impacts in mountain regions., This study was supported by the COST Action CLIMO (Climate-Smart Forestry in Mountain Regions – CA15226) financially supported by the EU Framework Programme for Research and Innovation HORIZON 2020. This publication is also part of a project that has received funding from the European Union’s HORIZON 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement No 778322 (CARE4C)., 10 Pág.

There is concern in the scientific community and among forest managers about potential reductions in the provisioning of forest ecosystem services due to the loss of tree species diversity. Many studies have shown how species diversity influences forest functioning, especially productivity, but the influence of structural diversity, such as tree size heterogeneity, has received much less attention. This study focused on understanding the relationship between stand productivity and several structural characteristics of spruce-fir-beech mountain forest stands in Europe. We used a dataset of 89 long-term plots in spruce-fir-beech forests distributed along the European mountains where the three species, Norway spruce (Picea abies (L.) Karst.), silver fir (Abies alba Mill.) and European beech (Fagus sylvatica L.), represent at least 75% of the basal area. Site-dependent conditions were accounted for in a linear mixed-effect basic model, which related the stand productivity with the morphological, climatic and pedological characteristics. The influence of tree species diversity, tree size heterogeneity, species size dominance, and species overlapping in the size distribution on stand productivity was analysed by adding variables to the basic model one by one and evaluating the change in the Akaike's Information Criterion (AIC). The variables that resulted in significant reductions in the AIC, and that were not correlated with each other, were used to build a model to estimate stand productivity. The model showed that in spruce-fir-beech mixed mountain forests (i) when Norway spruce, silver fir and European beech are evenly present within the size distribution (high evenness) the productivity decreases, (ii) the stand productivity increases when the diameter distribution is skewed to the right (higher numbers of smaller individuals), (iii) the stand productivity increases as the proportion of basal area that is spruce increases, and (iv) stand productivity increases with the variability in diameter. We discuss the implications of our results for the management of spruce-fir-beech mountain forest in Europe and for preserving and increasing the stand productivity of these mixed forests.

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.

Establishing mixed-species stands is frequently proposed as a strategy to adapt forests to the increasing risk of water scarcity, yet contrasted results have been reported regarding mixing effects on tree drought exposure. To investigate the drivers behind the spatial and temporal variation in water-related mixing effects, we analysed the δ13C variation in 22-year tree ring chronologies for beech and pine trees sampled from 17 pure and mixed pine–beech stands across a large gradient of environmental conditions throughout Europe. In the pure stands, average δ13C values were lower for beech (−27.9‰ to −22.2‰) than for pine (−26.0‰ to −21.1‰), irrespective of site conditions. Decreasing SPEI values (calculated over June to September) were associated with an increase in δ13C for both species, but their effect was influenced by stand basal area for pine and site water availability for beech. Mixing did not change the temporal constancy of δ13C nor the tree reaction to a drought event, for any of the species. While the mixing effect (Δ δ13C = δ13C pure stands − δ13C mixed stands) was on average positive for beech and non-significant for pine across the whole gradient, this effect strongly differed between sites. For both species, mixing was not significant at extremely dry sites and positive at dry sites; on moderately wet sites, mixing was positive for beech and negative for pine; at sites with permanent water supply, no general patterns emerge for any of the species. The pattern of mixing effect along the gradient of water availability was not linear but showed threshold points, highlighting the need to investigate such relation for other combinations of tree species., The networking in this study has been supported by COST Action FP1206 EUMIXFOR.

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.

Centro de Investigación Forestal (CIFOR), 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., Open Access funding enabled and organized by Projekt DEAL.The authors would like to acknowledge networking support by the COST (European Cooperation in Science and Technology) Action CLIMO (Climate-Smart Forestry in Mountain Regions—CA15226) financially supported by the EU Framework Programme for Research and Innovation HORIZON 2020. This publication is part of a project that has received funding from the European Union’s HORIZON 2020 research and innovation programme under the Marie Skłodowska-Curie Grant Agreement No 778322. Thanks are also due to the European Union for funding the project ‘Mixed species forest management. Lowering risk, increasing resilience (REFORM)’ (# 2816ERA02S under the framework of Sumforest ERA-Net). Further, we would like to thank the Bayerische Staatsforsten (BaySF) for providing the observational plots and to the Bavarian State Ministry of Food, Agriculture, and Forestry for permanent support of the Project W 07 ‘Long-term experimental plots for forest growth and yield research’ (#7831-26625-2017). We also thank the Forest Research Institute, ERTI Sárvár, Hungary, for assistance and for providing observational plots. Furthermore, our work was partially supported by the SRDA via Project No. APVV-16-0325 and APVV-15-0265, the Ministry of Science and Higher Education of the Republic of Poland, the Project “EVA4.0” No. CZ.02.1.01/0.0/0.0/16_019/0000803 funded by OP RDE and the Project J4-1765 funded by the Slovenian Research Agency and also by the Bulgarian National Science Fund (BNSF) and the Project No. DCOST 01/3/19.10.2018., 19 Pág.