Spatial distribution of tree-related microhabitats in a primeval mountain forest: From natural patterns to landscape planning and forest management recommendations.

Tree-related Microhabitats (TreMs) are essential for sustaining forest biodiversity. Although TreMs represent ephemeral resources that are spread across the landscape, their spatial distribution within temperate forests remains poorly understood. To address this knowledge gap, we conducted a study on 90 sample plots (0.05 ha each) located in a primeval mountain European beech Fagus sylvatica-dominated forest (Bieszczady Mountains, Carpathians). We explored the TreM profile with its link to habitat characteristics and described the spatial distribution of TreM indices. We identified 61 TreM types, with a mean richness of 19.7 ± 4.9 SD TreM types per plot, a mean density of 740.7 ± 292.5 SD TreM-bearing trees ha and a mean TreM diversity of 1.2 ± 0.1 SD. The diameter and living status of trees (living vs dead standing tree) were correlated with TreM richness on an individual tree. The stand structure, i.e. density and/or basal area of living and/or dead standing trees, and topographic conditions, i.e. slope exposure, were correlated with the TreM richness, density and diversity recorded on a study plot. We found no relationship between TreM richness, density and diversity and the presence of canopy gaps, which indicates that the influence of small-scale disturbances on the TreM profile is limited. However, our analysis revealed a clustered spatial pattern of TreM indices, with TreM-rich habitat patches (hot-spots) covering ~20 % of the forest. A moderate TreM richness, density and diversity dominated ~60 % of the forest, while TreM-poor habitat patches (cold-spots) covered ~20 %. Based on our findings, we advise the transfer of knowledge on the spatial distribution of TreMs from primeval to managed forests and advocate the '2:6:2' triad rule: to allocate 20 % of forests as strictly protected areas, to dedicate 60 % to low-intensity forest management with the retention of large living trees and all dead standing trees, and to use the remaining 20 % for intensive timber production. To ensure the continuance of the majority of TreM types, ≥55 living trees ha >60 cm in diameter should be retained. Such an approach will maintain a rich and diverse TreM assemblage across a broad spatial scale, which in turn will support biodiversity conservation and ecosystem restoration in secondary or managed forests.