Vascular plant and cryptogam diversity in Fagus sylvatica primeval forests and comparison to production stands in the western Carpathian Mountains, Slovakia

Summary Conflicting evidence exists with respect to the putative effect of forest management on plant species diversity. Various published studies have concluded that primeval forests are not that species-rich and that forest management may increase species richness in certain cases. Yet, it appears that such conclusions have often been drawn from the comparison of production forests to reference forests with a management legacy, i.e. stands in which forest management ceased only a few decades ago. The present study explores the impact of forest management on the diversity of vascular plants, bryophytes and lichens in Fagus sylvatica production forests in comparison to untouched primeval forests without any detectable management legacy in the western Carpathian Mountains, eastern Slovakia. This study in three production and three primeval forests also assesses the role of natural forest dynamics for species diversity and highlights the importance of height in the crown for the diversity and composition of epiphytic bryophyte and lichen communities. The results demonstrate that forest management-related disturbances do not increase landscape-level vascular plant species richness in comparison to untouched primeval forests. Even though mean plot-level diversity (alpha-diversity) was higher in the production forests, rarefaction/extrapolation showed a similarly high landscape-level vascular plant diversity (gamma-diversity) in the primeval forests. Comparing the gamma-diversity of vascular plants in the initial, optimal and terminal stages with the production forests showed no significant differences between the three stages, and to the managed stands. In contrast, mean plot-level species richness of epiphytic cryptogams tended to be higher in the primeval forests. Rarefaction/extrapolation revealed an about 30 and 100 % higher bryophyte and lichen species richness (gamma-diversity) in the primeval forests, respectively. Comparing the epiphyte species diversity of the three stages with the production forests provided a significantly higher species richness in any stage of the forest development. Species turnover between plots (beta-diversity) was in the three systematic groups generally higher in the primeval forests, indicating a greater habitat heterogeneity and spatially more variable species composition of the communities. In addition to habitat diversity, habitat continuity is playing an essential role for maintaining a high higher species richness in the primeval forests. This is visible when comparing the epiphyte species richness on stems of a given stem diameter class between production und primeval forests. Epiphytic bryophyte and lichen diversity per stem size class was significantly higher in the primeval forests. This was attributed to the fact that trees with large diameters in the production stands rarely were older than 100 years, whereas the maximum age of large-sized beeches in the primeval forest was over 400 years. Natural forest dynamics affected the composition, but not the species richness, of the forest floor vegetation. Certain species showed a strong preference for either the initial, optimal or terminal stage, reflecting a high species turnover in primeval forests. Vascular species richness, however, did not differ between the three stages. In contrast to vascular plants, bryophyte and lichen species composition and richness on living trees and standing deadwood was strongly affected by natural forest dynamics. In general, epiphyte diversity increased from the initial to the terminal stage. Several bryophytes and lichens showed a distinct preference for one of the three stages, showing a close association with the ageing of trees and diameter growth. Of the detected bryophytes and lichens, 50 and 22 %, respectively, were significantly associated with large-diameter stems (≥70 cm). Bryophytes and lichens on lying deadwood were not affected by natural forest development, as no stage differences in the species richness were detectable. The analysis of the epiphytic bryophyte and lichen vegetation from the tree base to the crown demonstrated that sampling only the lowermost 2 m results in a marked underestimation of total epiphyte species richness in temperate broadleaf forests. More specifically, about 10 % of the overall bryophyte and 48 % of the lichen species pool would have been missed. The vertical change in the epiphytic bryophyte and lichen flora reflects the increase in light intensity, decrease in atmospheric moisture and the growing input of acids and nitrogen compounds when moving from the trunk base to the canopy. The vertical change in microclimate and microsite conditions also leads to distinct compositional variations of both studied groups along the height gradient. This study clearly shows that forest management markedly reduces overall plant diversity and strongly impacts on the species composition in managed forests in comparison to untouched primeval forests. The higher plant species richness in primeval forests is mainly due to the much longer habitat continuity and greater habitat heterogeneity in horizontal and vertical direction, as generated by the processes of natural forest development. The disappearance of the terminal stage of forest development and a principal reduction in forest structural heterogeneity in production forests promote the loss of many plant species with close association to primeval forest attributes. Protecting the last remaining temperate primeval forests is thus an important element of a global strategy to conserve the biodiversity on earth.