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Primeval forests in the temperate zone exist only as a few remnants, but theses serve as important reference areas for conservation. As key habitats, tree-related microhabitats (TreMs) are of intense interest to forest ecologists, but little is known about their natural composition and dynamics in different tree species. Beech forms a major part of the temperate forests that extend from Europe, home to European beech Fagus sylvatica L. (Fs), eastward to Iran, where Oriental beech Fagus orientalis Lipsky (Fo) is the dominant species. In this study, we compared TreMs in primeval forests of both species, using data from Fo growing in 25 inventory plots throughout the Hyrcanian forest belt in Iran and from Fs growing in a 9 ha permanent plot in the Uholka Forest of Ukraine. TreMs based on 47 types and 11 subgroups were recorded. Beech trees in the Hyrcanian forest had a higher mean diameter at breast height (dbh) than beech trees in Uholka and contained twice as many TreMs per hectare. Although the mean richness of TreMs per TreM bearing tree was similar in the two species, on the basis of the comparison single trees in two groups (n = 405 vs. 2251), the composition of the TreMs clearly differed, as the proportions of rot holes, root-buttress concavities, and crown deadwood were higher in the Hyrcanian Forest, and those of bark losses, exposed heartwood, and burrs and cankers higher in Uholka Forest. Estimates of TreMs dynamics based on dbh and using Weibull models showed a significantly faster cumulative increase of TreMs in Fo, in which saturation occurred already in trees with a dbh of 70–80 cm. By contrast, the increase in TreMs in Fs was continuous. In both species, the probability density was highest at a dbh of about 30 cm, but was twice as high in Fo. Because of limitations of our study design, the reason behind observed differences of TreM formation and composition between regions remains unclear, as it could be either result of the tree species or the environment, or their interaction. However, the observed differences were more likely the result of differences in the environment than in the two tree species. Nevertheless, our findings demonstrate that the Hyrcanian Forest, recently designated as a natural heritage site in Iran, is unique, not only as a tertiary relict or due to its endemic trees, herbs and arthropods, but also because of its TreMs, which form a distinct and rich habitat for associated taxa, including endemic saproxylic species.

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1. The retention of trees bearing tree-related microhabitats (TreMs) has become an important means of conserving biodiversity in production forests. However, we lack estimates of TreM formation rates and evidence on factors driving TreM formation. 2. Based on the observation of 80,099 living trees from 19 species groups in Europe and Iran, we estimated the probability of TreM occurrence on trees and the associated rate of first TreM formation as a function of tree DBH, management, tree species group and random site effects. We built a separate model for each of 11 TreM groups. 3. The hazard rate of first TreM formation (defined as the probability of formation of a first TreM forming on a tree that is known to have none, during an infinitesimal DBH increment) increased with DBH for some TreM groups like breedingwoodpecker-hole, rot-hole or root-concavity, indicating an acceleration in TreM formation during tree growth. However, it decreased with DBH for TreM groups like bark-loss or dendrotelm, indicating slower formation on very large trees. Most TreM groups had reduced formation rates in managed forests (last logging less than 100 years ago) compared to unmanaged forests (no logging for at least 100 years), with the exception of dendrotelms. No general difference appeared | 493

Unmanaged forests are remnants of natural ecosystems that provide a basis for close-to-nature silvicultural research and applications. These forests have high amounts of dead wood, and although this material is being increasingly studied, the diversity of dead wood in terms of different diameters, decay stages, and spatial distribution patterns is as important as its volume for understanding forest dynamics. Here, we study natural forests in northern Iran to investigate the spatial distribution, decay stages, and volume of dead wood in unmanaged temperate forests at different developmental stages. Three stem-mapped sampling plots (100 m × 100 m) were established in uneven-aged stands dominated by Caspian beech (Fagus orientalis Lispsky). The total dead wood ranged from 37 to 119 m 2 ha −1 . Our results imply a spatial distribution shift from aggregation to OPEN ACCESSForests 2013, 4 752 randomness for dead trees in Caspian beech forest succession. We detected significant spatial interactions (attraction) between living and dead trees at short to medium spatial scales (1-20 m) in the plot with the earlier successional stage, suggesting that intra-specific competition is a prevailing force causing tree mortality at the stem-exclusion phase. By contrast, as trees become dominant with the mortality of other trees, the random tree-mortality pattern prevails. The spatial distribution and volume of dead wood may serve as a management target in near-to-natural Caspian beech forest. On the basis of our results, conservation-oriented management strategies should take into account the increasing amount of dead wood, particularly of large diameter in a late stage of decay.