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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.

Questions: How have the historical frequency and severity of natural disturbances in primary Picea abies forests varied at the forest stand and landscape level during recent centuries? Is there a relationship between physiographic attributes and historical patterns of disturbance severity in this system? Location: Primary P. abies forests of the Eastern Carpathian Mountains, Romania; a region thought to hold the largest concentration of primary P. abies forests in Europe's temperate zone.Methods: We used dendrochronological methods applied to many plots over a large area (132 plots representing six stands in two landscapes), thereby providing information at both stand and landscape levels. Evidence of past canopy disturbance was derived from two patterns of radial growth: (1) abrupt, sustained increases in growth (releases) and (2) rapid early growth rates (gap recruitment). These methods were augmented with non-metric multidimensional scaling to facilitate the interpretation of factors influencing past disturbance.Results: Of the two growth pattern criteria used to assess past disturbance, gap recruitment was the most common, representing 80% of disturbance evidence overall. Disturbance severities varied over the landscape, including stand-replacing events, as well as low-and intermediate-severity disturbances. More than half of the study plots experienced extreme-severity disturbances at the plot level, although they were not always synchronized across stands and landscapes. Plots indicating high-severity disturbances were often spatially clustered (indicating disturbances up to 20 ha), while this tendency was less clear for lowand moderate-severity disturbances. Physiographic attributes such as altitude and land form were only weakly correlated with disturbance severity. Historical documents suggest windstorms as the primary disturbance agent, while the role of bark beetles (Ips typographus) remains unclear. Conclusions:The historical disturbance regime revealed in this multi-scale study is characterized by considerable spatial and temporal heterogeneity, which could be seen among plots within stands, among stands within landscapes and between the two landscapes. When the disturbance regime was evaluated at these larger scales, the entire range of disturbance severity was revealed within this landscape.

Questions What historical natural disturbances have shaped the structure and development of an old‐growth, sub‐alpine Picea abies forest? Are large‐scale, high‐severity disturbances (similar to the recent windthrow and bark beetle outbreaks in the region) within the historical range of variability for this forest ecosystem? Can past disturbances explain the previously described gradient in stand structure that had been attributed to an elevation gradient? Location Šumava National Park (the Bohemian Forest) of the southwest Czech Republic. Methods We reconstructed the site's disturbance history using dendroecological methods in a 20‐ha study plot, established to span an elevation gradient. Growth patterns of 400 increment cores were screened for: (1) abrupt increases in radial growth indicating mortality of a former canopy tree and (2) rapid early growth rates indicating establishment in a former canopy gap. Results Spatial and temporal patterns of canopy accession varied markedly over the 20‐ha study area, resulting in disturbance pulses that corresponded to an elevation gradient. On the lower slope of the plot, the majority of the trees reached the canopy during two pulses (1770–1800 and 1820–1840), while most trees on the upper slope accessed the canopy in one pulse (1840–1860). Historically documented windstorms roughly coincide with peaks in our disturbance reconstruction. Conclusions Our study provides strong evidence that these forests were historically shaped by infrequent, moderate‐ to high‐severity natural disturbances. Our methods, however, could not definitively identify the agent(s) of these disturbances. Nevertheless, the recent mid‐1990s windstorm and the ensuing spruce bark beetle outbreak may provide an analogue for past disturbance, as the duration and severity of these events could easily explain past patterns of growth response and recruitment in our results. Thus, it seems reasonable to assume the interaction of windstorms and bark beetles seen in the contemporary landscape has occurred historically. Finally, our results suggest that the previously documented elevation gradient in forest structure may not be related to elevation per se (lower temperatures and shorter growing season) but rather to changes in disturbance severity mediated by elevation.

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