Drivers of tree regeneration in coniferous monocultures during conversion to mixed forests in Central Europe - Implications for forest restoration management

In Central Europe, anthropogenic coniferous monocultures have been subject to conversion to more diverse mixed forests since the 1990s, however, they are still abundant across many forest landscapes. Artificial and natural tree regeneration both play a key role during conversion by determining the species composition and structure of the future forests. Many abiotic and biotic factors can potentially influence the regeneration process and its specific combinations or interactions may be different among tree species and its developmental stages. Here, we aimed to identify and quantify the effect of the most important drivers on the density of the most abundant regenerating tree species (i.e., Norway spruce and European beech), as well as on species and structural diversity of the tree regeneration. We studied tree regeneration in four former monospecific coniferous stand types (i.e., Norway spruce, Scots pine, European larch, and Douglas fir) in Southwest Germany that have been under conversion to mixed forests since the 1990s. We sampled tree regeneration in four growth height classes together with a variety of potentially influencing factors on 108 sampling plots and applied multivariate analyses. We identified light availability in the understorey, stand structural attributes, browsing pressure, and diaspore source abundance as the most important factors for the density and diversity of tree regeneration. Particularly, we revealed speciesspecific differences in drivers of regeneration density. While spruce profited from increasing light availability and decreasing stand basal area, beech benefited either from a minor reduction or more strikingly from an increase in overstorey density. Increasing diaspore source abundance positively and a high browsing pressure negatively affected both species equally. Our results suggest that humus and topsoil properties were modified during conversion, probably due to changes in tree species composition and silvicultural activities. The species and structural diversity of the tree regeneration benefitted from increasing light availability, decreasing stand basal area, and a low to moderate browsing pressure. We conclude that forest managers may carefully equilibrate among the regulation of overstorey cover, stand basal area, and browsing pressure to fulfil the objectives of forest conversion, i.e., establishing and safeguarding a diverse tree regeneration to promote the development of mature mixed forests in the future.