In order to gauge ongoing and future changes to disturbance regimes, it is necessary to establish a solid baseline of historic disturbance patterns against which to evaluate these changes. Further, understanding how forest structure and composition respond to variation in past disturbances may provide insight into future resilience to climate-driven alterations of disturbance regimes. We established 184 plots (mostly 1000 m) in 14 primary mountain Norway spruce forests in the Western Carpathians. On each plot we surveyed live and dead trees and regeneration, and cored around 25 canopy trees. Disturbance history was reconstructed by examining individual tree growth trends. The study plots were further aggregated into five groups based on disturbance history (severity and timing) to evaluate and explain its influence on forest structure. These ecosystems are characterized by a mixed severity disturbance regime with high spatiotemporal variability in severity and frequency. However, periods of synchrony in disturbance activity were also found. Specifically, a peak of canopy disturbance was found for the mid-19th century across the region (about 60% of trees established), with the most important periods of disturbance in the 1820s and from the 1840s to the 1870s. Current stand size and age structure were strongly influenced by past disturbance activity. In contrast, past disturbances did not have a significant effect on current tree density, the amount of coarse woody debris, and regeneration. High mean densities of regeneration with height >50 cm (about 1400 individuals per ha) were observed. Extensive high severity disturbances have recently affected Central European forests, spurring a discussion about the causes and consequences. We found some evidence that forests in the Western Carpathians were predisposed to recent severe disturbance events as a result of synchronized past disturbance activity, which partly homogenized size and age structure and made recent stands more vulnerable to bark beetle outbreak. Our data suggest that these events are still part of the range of natural variability. The finding that regeneration density and volume of coarse woody debris were not influenced by past disturbance illustrates that vastly different past disturbance histories are not likely to change the future trajectories of these forests. These ecosystems currently have high ecological resilience to disturbance. In conclusion, we suggest that management should recognize disturbances as a natural part of ecosystem dynamics in the mountain forests of Central Europe, account for their stochastic occurrence in management planning, and mimic their patterns to foster biodiversity in forest landscapes.
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
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