Evaluating dynamic tree-species-shifting and height development caused by ungulate browsing in forest regeneration using a process-based modeling approach

Ungulate browsing can prevent the successful regeneration and development of forest-owner-intended and climate-resilient admixed tree species, compromising the future provision of multiple ecosystem services. A reduction of the height increment of palatable tree species caused by ungulate browsing results in a competition shift in favor of less palatable species. While this key process of losing admixed tree species is well understood, forest growth models have not yet captured its full complexity. Therefore, our objectives were to simulate (i) the loss of susceptible tree species from different browsing regimes and its impact on stand development, (ii) changes in the height increment of saplings and trees, and (iii) the shifts in social status during stand development without pre-defined limits a sapling can resist browsing. To address these objectives, we used the process-based model iLand to simulate the exposure of Norway spruce, Silver fir, and European beech stands to varying browsing densities. We successfully reproduced expected dynamic shifts in species composition and height development 24 years after establishment, depending on ungulate pressure and species-specific competitive strength. Stand composition and growth differences already occur with slight changes in ungulate density. When saplings remained in stand structure, independent of the number being browsed, our results led to more individuals with substantially smaller dimensions. We found that dynamic tree species shifting can be simulated realistically by a process-based modeling approach, which can be used to show the strong impact of ungulates on stand development.