The old-growth forest remnants of Western Carpathians provide a unique possibility to study the disturbance regimes of forest ecosystems without human influence. This study investigated the gap dynamics in beech-dominated old-growth forest Badínsky prales in Central Slovakia. Considering the decline of silver fir in last decades, the study analyzed the main characteristics of disturbance regime with the emphasis on the role of fir. On a 5-ha research plot, the dominant tree species was beech, the proportion of fir reached about 10%. However, a significantly higher proportion of fir ([30%) was observed in the coarse woody debris. In total, 45 canopy openings were recorded. Canopy gaps and expanded gaps covered 11.3 and 37.9% of the forest area, respectively. Despite the highest frequency of small gaps \100 m 2 , their proportion of the overall gap area reached only 20%, what suggests the important role of intermediate and large gaps in the gap dynamics as well. The analysis of gapmakers' crown projections confirmed a rather low contribution of fir (14.6%) to the gap formation despite its relatively intensive mortality. A high variability of the next generation age between the gaps (6-44 years) was recorded what suggests a large temporal variation of the disturbance events. The lateral expansion of adjacent trees was found to be the determining process for the closure of small canopy openings. The intermediate and large gaps are more likely closed by the height growth of natural regeneration and understory trees that are present on the majority of the area.
Research on old-growth forests is essential for understanding forest ecosystem functioning and for the development of forest management strategies. Their structural dynamics depends on disturbance patterns (White and Pickett, 1985). In recent decades, much research in old-growth forests has focused on natural disturbances. However, over several centuries, the natural disturbance regime has been severely altered by forest fragmentation, changes in fire regime, fauna extinctions, the introduction of forest management into the forest matrix surrounding old-growth, forest road infrastructure, and, in the last few decades, by overbrowsing, air pollution and climate change (Nowacki and Abrams, 1994;Uotila et al., 2002;Oulehle et al., 2010). Present-day old-growth forest structures are often a result of complex interactions between natural and anthropogenic disturbances (McLachlan et al., 2000). In Europe, old-growth forests have been significantly influenced by millennia of civilization. Thus, studies of
The structure of natural subalpine spruce forest in the Zadna´Pol'ana massif of the Western Carpathians was analysed. We focused on the variability of different aspects of stand structure, tree decay and regeneration processes in altitudinal gradient. We used systematic sampling, covering an area of 2 km 2 , to detect even subtle changes in stand structure within one forest type over a range of less than 200 m in elevation. Mean stand density was 290 trees (>7 cm DBH) per hectare, average basal area was 41 m 2 ha À1 , and the volume accumulation in living trees amounted to 500 m 3 /ha À1 . Stand volume decreased by more than 50% between 1,260 and 1,434 m a.s.l. This means for an increase of altitude of 100 m that stand volume decreased by nearly 200 m 3 . Neither stand density nor basal area was related to elevation. Maximum tree height was strongly correlated to elevation, and it decreased on average by 6 m for each 100 m increment of altitude. No significant changes in the maximum spruce diameter were recorded in relation to the elevation gradient. Spatial distribution of trees was biased toward regularity at lower altitudes. Tree clustering increased with increasing altitude. The stock of coarse woody debris (CWD) decreased slightly along the altitudinal gradient, but changes were not significant. Density of spruce saplings and their number growing on CWD significantly increased across the elevation gradient. Despite the fact that the analysed forest tract was relatively large, highly variable in respect to environmental factors, and that stand volume, spatial structure, and tree height displayed strong variability along the elevation gradient, the diameter structure of stands and regeneration measures were uniform. Our results suggest that the recruitment of new trees in the Zadna´Pol'ana subalpine spruce forest is not temporally continuous even at a scale of several square kilometres.
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