Thomas A. Nagel

Forest disturbances under climate change

Forest disturbances are sensitive to climate. However, our understanding of disturbance dynamics in response to climatic changes remains incomplete, particularly regarding large-scale patterns, interaction effects and dampening feedbacks. Here we provide a global synthesis of climate change effects on important abiotic (fire, drought, wind, snow and ice) and biotic (insects and pathogens) disturbance agents. Warmer and drier conditions particularly facilitate fire, drought and insect disturbances, while warmer and wetter conditions increase disturbances from wind and pathogens. Widespread interactions between agents are likely to amplify disturbances, while indirect climate effects such as vegetation changes can dampen long-term disturbance sensitivities to climate. Future changes in disturbance are likely to be most pronounced in coniferous forests and the boreal biome. We conclude that both ecosystems and society should be prepared for an increasingly disturbed future of forests.

A dendroecological reconstruction of disturbance in an old-growth Fagus-Abies forest in Slovenia

The scarcity of large old-growth forests has made it challenging to quantify disturbance regimes in Central Europe. The objective of this study was to reconstruct the history of disturbance in an old-growth Fagus-Abies forest reserve in Slovenia using a dendroecological approach. We extracted cores from canopy trees blown down during a recent windthrow event and identified growth releases in the tree-ring series using boundary-line release criteria to infer past disturbances. A total of 216 release events were identified from 88 trees. Between 1790 and 1990, moderate, asynchronous release events were present in nearly every decade of the disturbance chronology, suggesting a history of frequent, low severity disturbance. However, there were also peaks in the chronology corresponding to synchronous release events in a large proportion of the trees, suggesting that less frequent, intermediate severity disturbance events played an important role in forest development. These events are likely caused from wind damage associated with local thunderstorms, which seem to occur at intervals between 20–80 years on the study site. Thus, in addition to the small-scale gap phase processes operating in the forest, the results indicate that periodic intermediate severity disturbance events are an important component of the disturbance regime in mountain forests of Central Europe.RésuméLe faible nombre de forêts anciennes suffisamment étendues rend difficile la quantification des régimes de perturbation en Europe centrale. L’objectif de cette étude est de reconstituer l’histoire de ces perturbations dans une hêtraie-sapinière ancienne mise en réserve en Slovénie en utilisant une approche dendroécologique. Des carottes ont été extraites du tronc d’arbres dominants renversés lors d’un récent chablis. Les séries temporelles de cernes ont été analysées pour identifier des périodes de relâchement de contraintes de croissance et pour en déduire l’intensité des perturbations passées. Un total de 216 événements de relâchement de contrainte a été identifié pour 88 arbres. Entre 1790 et 1990, des événements d’intensité modérée et asynchrones ont été détectés dans les séries chronologiques au cours de pratiquement toutes les décennies. Cela suggère une fréquence élevée de perturbations de faible intensité. Cependant, des pics d’enregistrement d’événements synchrones ont été identifiés dans la chronologie dans une proportion importante d’arbres. Cela suggère que des perturbations de sévérité intermédiaire ont joué un rôle important dans le développement de la forêt, malgré leur plus faible fréquence. Ces événements ont probablement été causés par des chablis provoqués par des orages locaux violents, qui semblent se produire à des intervalles de 20 à 80 ans sur le site d’étude. Les résultats montrent ainsi qu’en sus de perturbations locales de faible intensité conduisant à la formation de petites trouées, les perturbations périodiques de sévérité intermédiaire constituent une composante importante du régime de perturbations dans les montagnes en Europe centrale.

A walk on the wild side: disturbance dynamics and the conservation and management of European mountain forest ecosystems

Mountain forests are among the most important ecosystems in Europe as they support numerous ecological, hydrological, climatic, social, and economic functions. They are unique relatively natural ecosystems consisting of long-lived species in an otherwise densely populated human landscape. Despite this, centuries of intensive forest management in many of these forests have eclipsed evidence of natural processes, especially the role of disturbances in long-term forest dynamics. Recent trends of land abandonment and establishment of protected forests have coincided with a growing interest in managing forests in more natural states. At the same time, the importance of past disturbances highlighted in an emerging body of literature, and recent increasing disturbances due to climate change are challenging long-held views of dynamics in these ecosystems. Here, we synthesize aspects of this Special Issue on the ecology of mountain forest ecosystems in Europe in the context of broader discussions in the field, to present a new perspective on these ecosystems and their natural disturbance regimes. Most mountain forests in Europe, for which long-term data are available, show a strong and long-term effect of not only human land use but also of natural disturbances that vary by orders of magnitude in size and frequency. Although these disturbances may kill many trees, the forests themselves have not been threatened. The relative importance of natural disturbances, land use, and climate change for ecosystem dynamics varies across space and time. Across the continent, changing climate and land use are altering forest cover, forest structure, tree demography, and natural disturbances, including fires, insect outbreaks, avalanches, and wind disturbances. Projected continued increases in forest area and biomass along with continued warming are likely to further promote forest disturbances. Episodic disturbances may foster ecosystem adaptation to the effects of ongoing and future climatic change. Increasing disturbances, along with trends of less intense land use, will promote further increases in coarse woody debris, with cascading positive effects on biodiversity, edaphic conditions, biogeochemical cycles, and increased heterogeneity across a range of spatial scales. Together, this may translate to disturbance-mediated resilience of forest landscapes and increased biodiversity, as long as climate and disturbance regimes remain within the tolerance of relevant species. Understanding ecological variability, even imperfectly, is integral to anticipating vulnerabilities and promoting ecological resilience, especially under growing uncertainty. Allowing some forests to be shaped by natural processes may be congruent with multiple goals of forest management, even in densely settled and developed countries.

Disturbance, life history traits, and dynamics in an old‐growth forest landscape of southeastern Europe

Much of our understanding of natural forest dynamics in the temperate region of Europe is based on observational studies in old-growth remnants that have emphasized small-scale gap dynamics and equilibrium stand structure and composition. Relatively little attention has been given to the role of infrequent disturbance events in forest dynamics. In this study, we analyzed dendroecological data from four stands and three windthrow patches in an old-growth landscape in the Dinaric Mountains of Bosnia and Herzegovina to examine disturbance history, tree life history traits, and compositional dynamics. Over all stands, most decades during the past 340 years experienced less than 10% canopy loss, yet each stand showed evidence of periodic intermediate-severity disturbances that removed > 40% of the canopy, some of which were synchronized over the study area landscape. Analysis of radial growth patterns indicated several life history differences among the dominant canopy trees; beech was markedly older than fir, while growth patterns of dead and dying trees suggested that fir was able to tolerate longer periods of suppressed growth in shade. Maple had the fastest radial growth and accessed the canopy primarily through rapid early growth in canopy gaps, whereas most beech and fir experienced a period of suppressed growth prior to canopy accession. Peaks in disturbance were roughly linked to increased recruitment, but mainly of shade-tolerant beech and fir; less tolerant species (i.e., maple, ash, and elm) recruited successfully on some of the windthown sites where advance regeneration of beech and fir was less abundant. The results challenge the traditional notions of stability in temperate old-growth forests of Europe and highlight the nonequilibrial nature of canopy composition due to unique histories of disturbance and tree life history differences. These findings provide valuable information for developing natural disturbance-based silvicultural systems, as well as insight into maintaining less shade-tolerant, but valuable broadleaved trees in temperate forests of Europe.

The historical disturbance regime of mountain Norway spruce forests in the Western Carpathians and its influence on current forest structure and composition

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 m2) 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.

Patterns and drivers of ice storm damage in temperate forests of Central Europe

Ice storms are important natural disturbances in temperate forests, yet have received little attention outside North America. Following an extreme ice storm in Slovenia, we examined patterns of ice damage within and among temperate forest sites and quantified differences in susceptibility to damage with respect to tree species and size across a gradient of storm intensity and site conditions. Based on a damage survey of 60 plots distributed across six unmanaged forest sites, ordinal logistic regression was used to examine patterns of ice damage as a function of storm intensity, species, tree size, and slope steepness. Our findings indicate that a complex interaction among these drivers gave rise to substantial variation in damage type and severity among species, plots, and stands. Fagus sylvatica, the most dominant species, was one of the most susceptible species to severe ice damage, while conifers (Abies alba and Picea abies) were least susceptible. Crown damage was the most common damage type at low storm intensity, while uprooting increased at higher intensity, particularly for large trees on steep slopes. Differences in species susceptibility to ice damage, combined with variation in storm intensity and site conditions, gave rise to heterogeneous damage patterns that have the potential to alter successional pathways. Based on an analysis of historical records, moderate-to-severe ice storms recur relatively frequently in the region, suggesting that they play a more important role in forest dynamics than previously thought.

Contrasting effects of environmental change on the radial growth of co-occurring beech and fir trees across Europe

Under predicted climate change, native silver fir (Abies alba) and European beech (Fagus sylvatica) are the most likely replacement species for the Norway spruce (Picea abies) monocultures planted across large parts of continental Europe. Our current understanding of the adaptation potential of fir-beech mixed forests to climate change is limited because long-term responses of the two species to environmental changes have not yet been comprehensively quantified. We compiled and analysed tree-ring width (TRW) series from 2855 dominant, co-dominant, sub-dominant and suppressed fir and beech trees sampled in 17 managed and unmanaged mixed beech-fir forest sites across Continental Europe, including Bosnia and Herzegovina, Germany, Italy, Romania and Slovakia. Dendroecological techniques that combine various detrending methods were used to investigate variation in radial growth of co-occurring fir and beech trees. Coincidental with peak SO2 emissions, the growth of silver fir declined between 1950 and 1980 at most sites, whereas beech growth increased during this period. Correspondent to a significant warming trend from 1990-2010, average beech growth declined, but silver fir growth increased. Long-term growth patterns and growth-climate sensitivity of fir and beech trees did not significantly differ between managed and unmanaged forests. Multi-decadal changes in the growth rate of all vertical tree classes were similar. In contrast to previous indications of limited drought susceptibility of beech mixed stands, this study suggests that the mixture of tree species in forest stands does not necessarily prevent growth depressions induced by long-term environmental change. Our results further imply that forest management does not necessarily alter their sensitivity to environmental changes.

Challenges for Uneven-Aged Silviculture in Restoration of Post-Disturbance Forests

Forest managers are often required to restore forest stands following natural 17 disturbances, a situation that may become more common and more challenging under global 18 change. In parts of Central Europe, particularly in mountain regions dominated by mixed 19 temperate forests, the use of relatively low intensity, uneven-aged silviculture is a common 20 management approach. Because this type of management is based on mimicking less intense 21 disturbances, the restoration of more severe disturbance patches within forested landscapes has 22 received little attention within the context of uneven-aged silviculture in the region. The goal of this 23 paper is to synthesize research on the restoration of forests damaged by disturbances in temperate 24 forests of Slovenia and neighbouring regions of Central Europe, where uneven-aged silviculture is 25 practiced. We place particular emphasis on the most important biotic and abiotic drivers of 26 post-disturbance regeneration, and use this information to inform silvicultural decisions about 27 applying natural or artificial regeneration in disturbed areas. We conclude with guidelines for 28 restoration silviculture in uneven-aged forest landscapes. 29