Importance of tree species size dominance and heterogeneity on the productivity of spruce-fir-beech mountain forest stands in Europe

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.