Mixed-species stands have been found to be more productive than would be expected from the performance of their component species in monocultures due to facilitation and complementarity between species, although these interactions depend on the combination of species present. Our study focuses on monospecific and mixed-species stands of Scots pine and Norway spruce using 20 triplets established in nine countries along a climatic gradient across Europe. Differences in mean tree and stand characteristics, productivity and stand structure were assessed. Basal area increment in mixed stands was 8% higher than expected while volume increment was only 2% greater. Scots pine trees growing in mixed-species stands showed 11% larger quadratic mean diameter, 7% larger dominant diameter, 17% higher basal area and 25% higher stand volume than trees growing in monospecific stands. Norway spruce showed only a non-significant tendency to lower mean values of diameters, heights, basal area, as well standing volume in mixtures than monocultures. Stand structure indices differed between mixed stands and monocultures of Scots pine showing a greater stratification in mixed-species stands. Furthermore, the studied morphological traits showed little variability for trees growing in monospecific stands, except for diameter at breast height, crown length and crown length ratio. For trees growing in mixed stands, all the morphological traits of the trees were identified as different. Some of these morphological traits were associated with relative productivity. Nevertheless, relative productivity in mixed-species stands was not related to site conditions.
Mixed forests are suggested as a strategic adaptation of forest management to climate change. Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) are tree species of high economic and ecological value for European forestry. Both species coexist naturally in a large part of their distributions but there is a lack of knowledge on the ecological functioning of mixtures of these species and how to manage such stands. This paper analyses these species' intra-and inter-specific competition, including size-symmetric vs. size-asymmetric competition, and explore the effect of weather conditions on tree growth and competition. We studied basal area growth at tree level for Scots pine and Norway spruce in mixed versus pure stands in 22 triplets of fully-stocked plots along a broad range of ecological conditions across Europe. Stand inventory and increment cores provided insights into how species mixing modifies tree growth compared with neighbouring pure stands. Five different competition indices, weather variables and their interactions were included and checked in basal area growth models using a linear mixed model approach. Interspecific size-asymmetric competition strongly influenced growth for both tree species, and was modulated by weather conditions. However, species height stratification in mixed stands resulted in a greater tree basal area growth of Scots pine (10.5 cm 2 year − 1) than in pure stands (9.3 cm 2 year − 1), as this species occupies the upper canopy layer. Scots pine growth depended on temperature and drought, whereas Norway spruce growth was influenced only by drought. Interspecific site-asymmetric competition increased in cold winters for Scots pine, and decreased after a drought year for Nor-way spruce. Although mixtures of these species may reduce tree size for Norway spruce, our results suggest that this could be offset by faster growth in Scots pine. How inter-specific competition and weather conditions alter tree growth may have strong implications for the management of Scots pine-Norway spruce mixtures along the rotation period into the ongoing climate change scenario.
Boreal forests constitute a large portion of the global forest area, yet they are undersampled through field surveys, and only a few remotely sensed data sources provide structural information wall-to-wall throughout the boreal domain. ArcticDEM is a collection of high-resolution (2 m) space-borne stereogrammetric digital surface models (DSM) covering the entire land area north of 60° of latitude. The free-availability of ArcticDEM data offers new possibilities for aboveground biomass mapping (AGB) across boreal forests, and thus it is necessary to evaluate the potential for these data to map AGB over alternative open-data sources (i.e., Sentinel-2). This study was performed over the entire land area of Norway north of 60° of latitude, and the Norwegian national forest inventory (NFI) was used as a source of field data composed of accurately geolocated field plots (n=7710) systematically distributed across the study area. Separate random forest models were fitted using NFI data, and corresponding remotely sensed data consisting of either: i) a canopy height model (ArcticCHM) obtained by subtracting a high-quality digital terrain model (DTM) from the ArcticDEM DSM height values, ii) Sentinel-2 (S2), or iii) a combination of the two (ArcticCHM+S2). Furthermore, we assessed the effect of the forest- and terrain-specific factors on the models’ predictive accuracy. The best model (,i.e., ArcticCHM+S2) explained nearly 60% of the variance of the training set, which translated in the largest accuracy in terms of root mean square error (RMSE=41.4 t ha−1 ). This result highlights the synergy between 3D and multispectral data in AGB modelling. Furthermore, this study showed that despite the importance of ArcticCHM variables, the S2 model performed slightly better than ArcticCHM model. This finding highlights some of the limitations of ArcticDEM, which, despite the unprecedented spatial resolution, is highly heterogeneous due to the blending of multiple acquisitions across different years and seasons. We found that both forest- and terrain-specific characteristics affected the uncertainty of the ArcticCHM+S2 model and concluded that the combined use of ArcticCHM and Sentinel-2 represents a viable solution for AGB mapping across boreal forests. The synergy between the two data sources allowed for a reduction of the saturation effects typical of multispectral data while ensuring the spatial consistency in the output predictions due to the removal of artifacts and data voids present in ArcticCHM data. While the main contribution of this study is to provide the first evidence of the best-case-scenario (i.e., availability of accurate terrain models) that ArcticDEM data can provide for large-scale AGB modelling, it remains critically important for other studies to investigate how ArcticDEM may be used in areas where no DTMs are available as is the case for large portions of the boreal zone.
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.