Separating forest continuity from tree age effects on plant diversity in the ground and epiphyte vegetation of a Central European mountain spruce forest

Forest continuity has been identified as an important factor influencing the structure and diversity of forest vegetation. Primary forests with centuries of continuity are usually more diverse than young secondary forests as forest are colonized only slowly and because the former are richer in old tree individuals. In the present study, performed in unmanaged high-elevation spruce forests of the Harz Mountains, Germany, we had the unique opportunity to separate the effects of forest continuity and tree age on plant diversity. We compared an old-growth spruce forest with century-long habitat continuity with an adjacent secondary spruce forest, which had naturally established on a former bog after 1796 when peat exploitation halted. Comparative analysis of the ground and epiphyte vegetation showed that the plant diversity of the old-growth forest was not higher than that of the secondary forest with a similar tree age of >200 years. Our results suggest that a period of >200 years was sufficient for the secondary forest to be colonized by the whole regional species pool of herbaceous and cryptogam forest plants and epiphytes. Therefore, it is likely that habitat structure, including the presence of old and decaying trees, was more important for determining plant diversity than the independent effect of forest continuity. Our results are probably not transferrable to spruce forests younger than 200 years and highly fragmented woodlands with long distances between new stands and old-growth forests that serve as diaspore sources. In addition, our results might be not transferable to remote areas without notable air pollution, as the epiphyte vegetation of the study area was influenced by SO₂ pollution in the second half of the 20th century.