Influence of wild ungulates on the secondary succession of an Apennine silver fir forest after windthrow

Windstorms are natural factors in vegetation dynamics, but their intensity is increasing and undermines the forest resilience. Post-event interventions depend on forest types and management purposes. To promote mixed stands that are less susceptible to windthrow than monocultures, natural tree regeneration is usually recommended. However, wild ungulates attracted by the increased food supply in the new clearings can influence the secondary succession and slow down vegetation recovery. The impact of ungulate pressure on secondary successions in blowdown areas is still poorly known, especially in Mediterranean areas, which is particularly vulnerable to climate stressors. We investigated this topic using a monospecific silver fir (Abies alba) forest of artificial origin in the Apennines as a model system. The forest was left to natural succession after a severe windthrow in 2015, offering the opportunity to establish an ungulate exclosure experiment to analyse vegetation changes over six years. For the whole plant community, cover and height, α-diversity (species richness, Shannon and evenness indexes), and species composition were recorded yearly, together with tree seedling density, in open and fenced plots at increasing distances from the forest edge. Ecological and functional traits (proportion of light-demanding and forest specialists, endo-/epi-zoochorous species), and life-forms of the community samples were also analysed. Overall, we found that ungulates significantly influenced the dynamic trajectories of secondary succession. The effects on plant cover and diversity were negative and increased with distance from the forest edge. The presence of ungulates favoured generalist species and endozoochorous taxa. However, distance from the forest edge strongly reduced the latter effect. The average density of tree seedlings was overall high (ca. 10,000/ha), but it was strongly reduced by ungulates for three deciduous species and A. alba, the most browsed species. Natural recolonization after windthrow in Apennine pure silver fir stands may favour the formation of mixed, less susceptible forests, but the current ungulate pressure slows down this process, especially in the areas furthest from the forest edge.