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Abstract

Mortality, driven by both climate and disturbance legacies, is a key process shaping forest dynamics. Understanding the mortality patterns in primary forests in the absence of severe disturbances provides information on background natural dynamics of a given forest type under ongoing climate change. This can then be compared to mortality rates in severely-disturbed stands. Using a large number of sample plots along a gradient from low to high disturbance, we examined the mortality rates and composition of mortality agents in primary mountain Norway spruce (Picea abies (L.) Karst.) forests on different spatial scales. We evaluated the mortality rates and causes of mortality in 28 stands across a large geographical gradient spanning over 1000 km. We resampled (five-year period) 371 plots (16,287 living trees) in primary Norway spruce forests along the Carpathian mountain chain. The estimated overall annual mortality rate was within the previously reported range of background (ambient) mortality, however, stand-level and plot-level mortality rates varied substantially. Over 18% of plots displayed more than 2% annual mortality and 6% of plots even exceeded 10% per year. Stands in the Western Carpathians showed the highest variability in the mortality rate, with 30% of the stands in this region showing annual mortality rates over 5%. At the plot level, mixed-severity disturbances increased variability of mortality rates within most localities. Overall mortality was evenly distributed among size classes up to 50 cm diameter at breast height (DBH). However, the distributions differ for individual mortality agents. Mortality modes were classified into six categories (broken crown, broken stem, uprooted, competition, bark beetle/fungi, climatic extremes). Bark beetle (Ips typographus L.) infestation was the most frequent mortality agent in all stands, whereas the influence of competition as a mortality agent varied substantially. Mortality from abiotically-caused physical damage was similar to that from competition, yet the distribution among modes of physical damage (uprooted, crown, or stem breakage) varied. The lack of clear evidence of mortality agents in some locations implies that many tree deaths are caused by a combination of contributing factors. The results suggest the role of bark beetle as a mortality agent does not equate to severe mortality at large scales. Prevalence of different size classes affected by individual mortality agents underline the high complexity of the mortality process in primary forests.

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... As a result of dry weather established in Belarus in last decades, decline and dieback of the spruce (Picea abies L.) formation has begun [2] and is still continuing. Colonization of weakened trees by the bark beetle (Ips typographus) [2] [10] was the major reason of spruce dieback in Belarus in 2019 [1] [11], being also the cause of drying out of spruce in 75% of cases in the forests of Central and Eastern Europe [12] [13]. Therefore, prompt monitoring of forest areas employing remote sensing methods is of interest in order to identify dying coniferous trees at the initial stage for follow-up activities concerning preventing and mitigation of dieback [14]. ...
... The present study does not include information on the causes or timing of tree death, which can be ascribed to a multitude of abiotic and biotic factors and often a combination of them (Manion, 1981;Das et al., 2011;Pommerening and Särkkä, 2013;Synek et al., 2020). Moreover, the multitude of abiotic and biotic mortality factors only partly explains the spatial pattern of dead trees, since the mortality events that we analyzed are constrained to occur only within the locations where tree regeneration was successfully reaching the calipering threshold of 4 cm dbh (Aakala et al., 2012). ...
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Spatial patterns can reveal a lot about ecological processes, but our knowledge of the spatial ecology of tree regeneration at a fine scale is quite limited. Therefore, we studied the spatial patterns of living and dead small trees in two subalpine Norway spruce forest reserves in Switzerland (Scatl`e and B¨odmerenwald) using three types of analyses. First, we investigated the distances of small trees to the nearest large neighboring tree and, by using maximum distances as indicator, inferred the size of forest gaps, detecting mainly forest gaps of small size, although with two exceptions that were driven by large-scale disturbances. Second, we accounted for spatial inhomogeneity in the pattern of small and large trees (i.e., variations in local tree densities) by including environmental covariates in point pattern models. Latitude (within the forest reserve), elevation and aspect contributed significantly to explaining the density of living and dead small trees, and partly of living and dead large trees. Yet, the influence of these environmental covariates varied between the two reserves due to their different topography and peculiar site conditions. Third, we analyzed neighborhood interactions between small and large trees based on the vicinity and size of trees. In both forest reserves, small living trees were randomly dispersed around large dead trees over a broad range of distances and, at certain distances in one reserve, even dispersed away from them. Small living trees further showed clustering around large living trees at short distances and dispersion at large distances. Small dead trees featured mainly a random pattern, although with a tendency to cluster around large neighbors at short distances, irrespective whether these were living or dead. Yet, the weakening of clustering with increasing distances indicates that the influence of large trees on small trees varies with spatial scale and thus that these neighborhood interactions are scale-dependent. Overall, our study contributes to a better understanding of the spatial ecology of mortality in small trees and ultimately of tree regeneration processes and stand dynamics in mountain forests.
... In Southeastern Europe convective instability is more common (Furtuna et al., 2018;Nagel et al., 2017;Taszarek et al., 2019), whereas cyclones dominate in Western Europe. Despite the lower prevalence of cyclone-induced winds in Central, Southern, and Eastern Europe, windthrow is still a major disturbance agent driving forest dynamics there Sommerfeld et al., 2018;Synek et al., 2020). Disturbance reconstructions in old-growth European beech (Fagus sylvatica L.) dominated forests from Slovenia to Montenegro show that windthrow is the most prevalent disturbance agent but large-scale stand-leveling windstorms are not the norm (Furtuna et al., 2018;Nagel et al., 2017). ...
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Wind is the leading disturbance agent in European forests, and the magnitude of wind impacts on forest mortality has increased over recent decades. However, the atmospheric triggers behind severe winds in Western Europe (large‐scale cyclones) differ from those in Southeastern Europe (small‐scale convective instability). This geographic difference in wind drivers alters the spatial scale of resulting disturbances and potentially the sensitivity to climate change. Over the 20th century, the severity and prevalence of cyclone‐induced windstorms have increased while the prevalence of atmospheric instability has decreased and thus, the trajectory of Europe‐wide windthrow remains uncertain. To better predict forest sensitivity and trends of windthrow disturbance we used dendrochronological methods to reconstruct 140 years of disturbance history in beech‐dominated primary forests of Central and Eastern Europe. We compared generalized linear mixed models of these disturbance time series to determine whether large‐scale cyclones or small‐scale convective storms were more responsible for disturbance severity while also accounting for topography and stand character variables likely to influence windthrow susceptibility. More exposed forests, forests with a longer absence of disturbance, and forests lacking recent high severity disturbance showed increased sensitivity to both wind drivers. Large‐scale cyclone‐induced windstorms were the main driver of disturbance severity at both the plot and stand scale (0.1–∼100 ha) whereas convective instability effects were more localized (0.1 ha). Though the prevalence and severity of cyclone‐induced windstorms have increased over the 20 century, primary beech forests did not display an increase in the severity of windthrow observed over the same period.
... Eruptive insect species are considered as one of the main disturbance agents causing changes in ecosystems and affecting people's livelihoods, the economy, and hydrology [3][4][5][6]. In Europe, one of the main tree species affected by large-scale insect outbreaks is the Norway spruce (Picea abies, [L.] Karst) [7], whose natural habitat belt spans Scandinavian and Eastern European lowlands, montane areas in Central Europe, and large parts of the Carpathian Mts. [8]. ...
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Research Highlights: Bark beetles are important agents of disturbance regimes in temperate forests, and specifically in a connected wind-bark beetle disturbance system. Large-scale windthrows trigger population growth of the European spruce bark beetle (Ips typographus L.) from endemic to epidemic levels, thereby allowing the killing of Norway spruce trees over several consecutive years. Background and Objectives: There is a lack of evidence to differentiate how outbreaks are promoted by the effects of environmental variables versus beetle preferences of trees from endemic to outbreak. However, little is known about how individual downed-tree characteristics and local conditions such as tree orientation and solar radiation affect beetle colonization of downed trees. Materials and Methods: To answer this question, we investigated the infestation rates and determined tree death categories (uprooted, broken, and stump) in wind-damaged areas in Western Tatra Mts. in Carpathians (Slovakia) from 2014-2016, following a windthrow in May 2014. In total, we investigated 225 trees over eight transects. For every tree, we measured its morphological (tree height, crown characteristics), environmental (solar radiation, terrain conditions, trunk zenith), temporal (time since wind damage), and beetle infestation (presence, location of attack, bark desiccation) parameters. We applied Generalized Additive Mixed Models (GAMM) to unravel the main drivers of I. typographus infestations. Results: Over the first year, beetles preferred to attack broken trees and sun-exposed trunk sides over uprooted trees; the infestation on shaded sides started in the second year along with the infestation of uprooted trees with lower desiccation rates. We found that time since wind damage, stem length, and incident solar radiation increased the probability of beetle infestation, although both solar radiation and trunk zenith exhibited nonlinear variability. Our novel variable trunk zenith appeared to be an important predictor of bark beetle infestation probability. We conclude that trunk zenith as a simple measure defining the position of downed trees over the terrain can anticipate beetle infestation. Conclusions: Our findings contribute to understanding of the bark beetle's preferences to colonize windthrown trees in the initial years after the primary wind damage. Further, our findings can help to identify trees that are most susceptible to beetle infestation and to prioritize management actions to control beetle population while maintaining biodiversity.
... For Norway spruce, this means that the trees may be more susceptible to decline in drought years (Biermayer and Tretter 2016;Kölling et al. 2009) or that the trees have a requirement for facilitated positive interspecific interactions (Brandl and Falk 2019;del Río et al. 2014;Pretzsch et al. 2012). In addition, Norway spruce is generally characterized by rather high mortality rates also in its natural range (Synek et al. 2020). ...
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Key message Under severe drought, growth of Norway spruce suffered much more than European beech. Norway spruce benefited from growing in the environment of beech, and both species acclimated slightly to 5 years of experimentally extended drought. Abstract Recent studies show that the detrimental effects of drought on stand growth are mitigated when the stand contains mixed tree species. We analysed the growth responses of Norway spruce and European beech to episodic and experimentally extended drought in intra- and inter-specific mature stands. We used annual diameter growth records dating back to 1998 to determine the impact of the natural episodic drought in 2003 and 2015. To analyse extended drought, spruce and beech trees were exposed to extreme drought under automatic throughfall exclusion roofs from 2014 to 2018. The growth of spruce in an inter-specific environment with beech was 20–50% less affected by natural episodic drought compared with an intra-specific constellation. When beech grew in an inter-specific environment, it was by 23% more affected by drought compared to intra-specific conditions, but seemed to recover faster. The induced drought from 2014 to 2018 resulted in a strong growth reduction in the first year particularly for spruce, followed by a slight acclimation to the dry conditions. Beech acclimated and recovered faster than spruce across all growing conditions, while spruce only acclimatized faster in the environment of beech. Both species showed a higher mortality under induced drought compared with the controls; for spruce, the mortality rate was fivefold higher compared to the long-term mortality. The long-term moderate-growth stabilization and the growth increase after the 5-year exposure to drought suggest a gradual acclimation to drought by beech. The resistance and acclimation to drought of spruce when growing in mixture should be considered when designing resource efficient and productive mixed conifer-broadleaved stands for future climates.
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Forest typology is yet to get sufficiently integrated into related ecological and geographical sciences. The succession of plant phytocenoses necessitates constant adjustments to forest types. The discussed studies have been conducted to improve the classification of forest types in the Ukrainian Carpathians and bring the description closer to the actual distribution of vegetation within the studied area. The paper provides a detailed analysis of forest typology research for the Ukrainian Carpathians area during late 20 th and early 21 st century. The forest fund areas of the Carpathian region, which are subordinated to the State Agency of Forest Resources of Ukraine, have been classified by forest vegetation types. Total area of subor and coniferous forest types is 1,493.1 ha and 28,910.2 ha, respectively. The study involved establishing of permanent sample plots on the territory of nature protection institutions and forest management enterprises. According to the findings, it is proposed to complement the classification of subor and coniferous forest type as defined by Z.Yu. Herushynskyi on the territory of the Ukrainian Carpathians with the following types: fresh pine subor forest type, wet pine subor forest type, and wet pine coniferous type. The paper defines the main diagnostic features of the suggested forest types. These subor and coniferous forest types can be clearly distinguished from other forest types by soil and hydrological conditions, and can be used to describe the corresponding forest vegetation types. The correctness of definition of new forest types is confirmed with a set of plant indicator species that have been identified within the relevant areas. The findings provide a better understanding of forest ecology and make a significant contribution to forest typology studies on the territory of the Ukrainian Carpathians. Another step towards researching the patterns of the establishment of plant complexes in the Ukrainian Carpathians has been taken. Forest formations of the Carpathians are presented in more detail in forest typological science
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The increase in mean annual temperature and reduction in summer rainfall from climate change seem to increase the frequency of natural and human-made disturbances to forest vegetation. This type of rapid vegetation change also significantly affects bat diversity. The aim of our study was to document differences in the ecological parameters of bat assemblages in different types of temperate mountain forests, particularly between disturbed and undisturbed coniferous and deciduous forests. Additionally, these issues were considered along an elevation gradient. We mist netted bats on 73 sites, between 931 and 1453 m elevation, in the forests of the Tatra Mountains in southern Poland. During 2016–2020, 745 bats, representing 15 species, were caught. The most abundant were Myotis mystacinus (Kuhl, 1817) (53.0%) and M. brandtii (Eversmann, 1845) (21.5%). We observed differences in species diversity, elevational distribution, and dominance between different types of forests and forest zones. Species richness peaked at around 1000–1100 m elevation. The highest species richness and other indices were observed in undisturbed beech stands, although they constituted only about 2.7% of the forest area. The lowest species diversity was observed in disturbed coniferous forests, in both the lower and upper forest zone. The species richness and dominance structure of bat assemblages were also found to depend on the location above sea level. In some bat species, the sex ratio was higher at higher elevations, and differences in the sex ratio in a few bat species, between different types of forests, were observed. Our findings suggest that disturbed, beetle-killed spruce forests are an unsuitable environment for some bat species.
Article
Tree mortality is an important ecological process influencing multiple functions of forest ecosystems. Previous studies have shown two basic size‐mortality patterns, including a competition‐driven declining and a disturbance‐driven increasing mortality rate with tree size. Subtropical forests, which have a high species diversity and subject to frequent monsoon disturbances, are widely distributed in eastern Asia. However, the tree size‐mortality pattern in the mature subtropical forests remains unclear. Here we analyzed patterns of size‐dependent mortality from tree species to forest community using a 5‐year inventory data from 117 species and 163,612 individuals in a 20‐ha forest dynamic plot in a mature subtropical monsoon evergreen forest in eastern China. To explain the spatial variability in mortality patterns, two major biotic drivers (competition and tree size) and multiple local‐scale environmental factors were further analyzed. Our results showed that tree size was the best predictor of tree mortality at the scales of both species and community. A species‐level analysis identified four size‐mortality patterns that are shaped by species‐specific attributes such as maximum size and life form. For 27 out of 92 species that comprised 59% of tree individuals, the relationship between size and mortality exhibited a U‐shaped pattern of a first decline followed by an increase. An overall community‐scale size‐dependent mortality also showed a U‐shaped pattern. Tree mortality was also influenced by the competition and environmental conditions, but the relative importance varied widely across tree sizes and species. The competition showed significant correlations with the mortality of small trees, while the effect of environmental conditions on mortality was strongest for large trees. A principal component analysis showed that a combination of biotic and abiotic factors explained 42.3% of the spatial variation in mortality at large sizes. Synthesis. Our results reveal four identifiable size‐dependent mortality patterns that differ across diverse species, jointly leading to a U‐shaped size‐mortality pattern at the community level. This finding calls for the need to establish the details of every potential size‐mortality pattern with consideration of the different effects of biotic and abiotic factors on tree mortality of specific size.
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Models are pivotal for assessing future forest dynamics under the impacts of changing climate and management practices, incorporating representations of tree growth, mortality, and regeneration. Quantitative studies on the importance of mortality submodels are scarce. We evaluated 15 dynamic vegetation models (DVMs) regarding their sensitivity to different formulations of tree mortality under different degrees of climate change. The set of models comprised eight DVMs at the stand scale, three at the landscape scale, and four typically applied at the continental to global scale. Some incorporate empirically derived mortality models, and others are based on experimental data, whereas still others are based on theoretical reasoning. Each DVM was run with at least two alternative mortality submodels. Model behavior was evaluated against empirical time series data, and then, the models were subjected to different scenarios of climate change. Most DVMs matched empirical data quite well, irrespective of the mortality submodel that was used. However, mortality submodels that performed in a very similar manner against past data often led to sharply different trajectories of forest dynamics under future climate change. Most DVMs featured high sensitivity to the mortality submodel, with deviations of basal area and stem numbers on the order of 10–40% per century under current climate and 20–170% under climate change. The sensitivity of a given DVM to scenarios of climate change, however, was typically lower by a factor of two to three. We conclude that (1) mortality is one of the most uncertain processes when it comes to assessing forest response to climate change, and (2) more data and a better process understanding of tree mortality are needed to improve the robustness of simulated future forest dynamics. Our study highlights that comparing several alternative mortality formulations in DVMs provides valuable insights into the effects of process uncertainties on simulated future forest dynamics.
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Aim Primary forests have high conservation value but are rare in Europe due to historic land use. Yet many primary forest patches remain unmapped, and it is unclear to what extent they are effectively protected. Our aim was to (1) compile the most comprehensive European‐scale map of currently known primary forests, (2) analyse the spatial determinants characterizing their location and (3) locate areas where so far unmapped primary forests likely occur. Location Europe. Methods We aggregated data from a literature review, online questionnaires and 32 datasets of primary forests. We used boosted regression trees to explore which biophysical, socio‐economic and forest‐related variables explain the current distribution of primary forests. Finally, we predicted and mapped the relative likelihood of primary forest occurrence at a 1‐km resolution across Europe. Results Data on primary forests were frequently incomplete or inconsistent among countries. Known primary forests covered 1.4 Mha in 32 countries (0.7% of Europe’s forest area). Most of these forests were protected (89%), but only 46% of them strictly. Primary forests mostly occurred in mountain and boreal areas and were unevenly distributed across countries, biogeographical regions and forest types. Unmapped primary forests likely occur in the least accessible and populated areas, where forests cover a greater share of land, but wood demand historically has been low. Main conclusions Despite their outstanding conservation value, primary forests are rare and their current distribution is the result of centuries of land use and forest management. The conservation outlook for primary forests is uncertain as many are not strictly protected and most are small and fragmented, making them prone to extinction debt and human disturbance. Predicting where unmapped primary forests likely occur could guide conservation efforts, especially in Eastern Europe where large areas of primary forest still exist but are being lost at an alarming pace.
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Determining the drivers of shifting forest disturbance rates remains a pressing global change issue. Large-scale forest dynamics are commonly assumed to be climate driven, but appropriately scaled disturbance histories are rarely available to assess how disturbance legacies alter subsequent disturbance rates and the climate sensitivity of disturbance. We compiled multiple tree-ring based disturbance histories from primary Picea abies forest fragments distributed throughout five European landscapes spanning the Bohemian Forest and the Carpathian Mountains. The regional chronology includes 11 595 tree cores, with ring dates spanning the years 1750 to 2000, collected from 560 inventory plots in 37 stands distributed across a 1000 km geographic gradient, amounting to the largest disturbance chronology yet constructed in Europe. Decadal disturbance rates varied significantly through time and declined after 1920, resulting in widespread increases in canopy tree age. Approximately 75% of current canopy area recruited prior to 1900. Long-term disturbance patterns were compared to an historical drought reconstruction, and further linked to spatial variation in stand structure and contemporary disturbance patterns derived from LANDSAT imagery. Historically, decadal Palmer drought severity index minima corresponded with higher rates of canopy removal. The severity of contemporary disturbances increased with each stand's estimated time since last major disturbance, increased with mean diameter and declined with increasing within-stand structural variability. Reconstructed spatial patterns suggest that high small-scale structural variability has historically acted to reduce large-scale susceptibility and climate sensitivity of disturbance. Reduced disturbance rates since 1920, a potential legacy of high 19th century disturbance rates, have contributed to a recent region-wide increase in disturbance susceptibility. Increasingly common high-severity disturbances throughout primary Picea forests of Central Europe should be reinterpreted in light of both legacy effects (resulting in increased susceptibility) and climate change (resulting in increased exposure to extreme events). This article is protected by copyright. All rights reserved.
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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 ( re, drought, wind, snow and ice) and biotic (insects and pathogens) disturbance agents. Warmer and drier conditions particularly facilitate re, 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.
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ContextGrowing evidence suggests that climate change could substantially alter forest disturbances. Interactions between individual disturbance agents are a major component of disturbance regimes, yet how interactions contribute to their climate sensitivity remains largely unknown. Objectives Here, our aim was to assess the climate sensitivity of disturbance interactions, focusing on wind and bark beetle disturbances. Methods We developed a process-based model of bark beetle disturbance, integrated into the dynamic forest landscape model iLand (already including a detailed model of wind disturbance). We evaluated the integrated model against observations from three wind events and a subsequent bark beetle outbreak, affecting 530.2 ha (3.8 %) of a mountain forest landscape in Austria between 2007 and 2014. Subsequently, we conducted a factorial experiment determining the effect of changes in climate variables on the area disturbed by wind and bark beetles separately and in combination. ResultsiLand was well able to reproduce observations with regard to area, temporal sequence, and spatial pattern of disturbance. The observed disturbance dynamics was strongly driven by interactions, with 64.3 % of the area disturbed attributed to interaction effects. A +4 °C warming increased the disturbed area by +264.7 % and the area-weighted mean patch size by +1794.3 %. Interactions were found to have a ten times higher sensitivity to temperature changes than main effects, considerably amplifying the climate sensitivity of the disturbance regime. Conclusions Disturbance interactions are a key component of the forest disturbance regime. Neglecting interaction effects can lead to a substantial underestimation of the climate change sensitivity of disturbance regimes.
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Bark beetle outbreaks have had major impacts on Norway spruce forests in Europe. The large majority of these forests are located in areas under forest management; thus, few studies have investigated outbreak-driven spruce mortality patterns unaffected by humans. Our study examined spruce mortality resultant from a beetle outbreak in a high-elevation, unmanaged forest over a 17-year span. We analyzed three tree-level survivorship and DBH datasets collected during pre-, mid-, and post-outbreak conditions to evaluate long-term mortality dynamics. We measured changes in mortality severity, topographic and stand structure characteristics, and stand complexity using ANOVAs, and we assessed five topographic and stand structure mortality predictors by employing boosted regression trees. Our results showed that though spruce mortality increased significantly over time, such increases were disproportionate with spatial synchrony. Moreover, the outbreak did not significantly alter the living stand structure and had little effect on stand complexity, exhibiting the effects of an outbreak that spread throughout the forest without causing major stand-level damage. Larger trees at higher elevations on south-facing slopes were targeted most frequently, particularly during the later stages of the outbreak. Aspect, elevation, and slope were the best predictors of mortality, demonstrating moderate forecasting ability. We showed that bark beetle outbreaks can operate on patch-scale gradients, affecting microhabitat conditions, without resulting in sweeping, stand-altering mortality. Small-scale outbreaks may increase forest resilience against more severe outbreaks in the future by creating canopy gaps that facilitate regeneration, which leads to more complex age and size structures within the stand.
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Climate change is expected to drive increased tree mortality through drought, heat stress, and insect attacks, with manifold impacts on forest ecosystems. Yet, climate-induced tree mortality and biotic disturbance agents are largely absent from process-based ecosystem models. Using data sets from the western USA and associated studies, we present a framework for determining the relative contribution of drought stress, insect attack, and their interactions, which is critical for modeling mortality in future climates. We outline a simple approach that identifies the mechanisms associated with two guilds of insects - bark beetles and defoliators - which are responsible for substantial tree mortality. We then discuss cross-biome patterns of insect-driven tree mortality and draw upon available evidence contrasting the prevalence of insect outbreaks in temperate and tropical regions. We conclude with an overview of tools and promising avenues to address major challenges. Ultimately, a multitrophic approach that captures tree physiology, insect populations, and tree-insect interactions will better inform projections of forest ecosystem responses to climate change. © 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.
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Forest insects and pathogens (FIPs) have enormous impacts on community dynamics, carbon storage and ecosystem services, however, ecosystem modelling of FIPs is limited due to their variability in severity and extent. We present a general framework for modelling FIP disturbances through their impacts on tree ecophysiology. Five pathways are identified as the basis for functional groupings: increases in leaf, stem and root turnover, and reductions in phloem and xylem transport. A simple ecophysiological model was used to explore the sensitivity of forest growth, mortality and ecosystem fluxes to varying outbreak severity. Across all pathways, low infection was associated with growth reduction but limited mortality. Moderate infection led to individual tree mortality, whereas high levels led to stand-level die-offs delayed over multiple years. Delayed mortality is consistent with observations and critical for capturing biophysical, biogeochemical and successional responses. This framework enables novel predictions under present and future global change scenarios.
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QuestionsWhat are the spatial patterns of adult trees and recruits in natural Picea abies forests? How are these patterns related to each other? Does the relationship differ before and after stand-replacing disturbance? What are the ecological processes behind these spatial patterns? LocationMountain Picea abies forests in the Sumava Mts. (Bohemian Forest), Czech Republic, affected by a high-severity outbreak of bark beetle (Ips typographus) that caused large-scale dieback of the forest canopy. Methods We measured the spatial coordinates and heights of all recruits and the coordinates and DBH of all adult trees in nine plots across a wide range of recruit densities. We distinguished pre- and post-disturbance recruits, trees killed by the disturbance and trees already dead before it. To analyse spatial relationship among these groups, we used univariate and bivariate pair-correlation functions. To provide further insight into the mechanisms behind the observed patterns, we fitted Thomas and Matern point processes to the observed data. ResultsRecruits formed tight clusters (2-9m), whereas trees were distributed randomly or weakly clustered at short distances (1-2m). Both pre-disturbance and post-disturbance recruits were highly clustered (cluster radii<2m) around trees before and after the stand-replacing disturbance. This fine-scale pattern was likely driven by a combination of: (1) seed accumulation in tree wells during winter; (2) nurse effects of tree trunks extending the vegetation period, suppressing competitive vegetation and enhancing nutrient supply from decomposed litter; and (3) suitable seedbeds on some decaying wood. The Thomas point process fitted the observed pattern of decreasing recruit density with increasing distance from mature trees better than the Matern process. Conclusions Tree spatial pattern in mountain P.abies forests showed high resilience to stand-replacing disturbance. After a self-thinning of recruits tightly clustered around parental trees, their spatial pattern will mirror the pattern of trees that formed the stand before the disturbance. This memory' of tree spatial patterns is an important biological legacy and should be viewed as a fundamental property of natural P.abies forests.
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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). Thesemethods were augmented with non-metricmultidimensional 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.
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Bark beetle dispersal and host selection behaviour are a complex and poorly understood process, resulting in specific spatio-temporal infestation patterns in forests. Aerial images from the Bavarian Forest National Park (Germany) provide a high-resolution, that is, tree-scale data set for the period 2001–2010, including information about Ips typographus (Col., Curculio., Scolytinae) infestation, the application of sanitary logging, natural forest edges and the area of living spruce susceptible to bark beetle infestation. We combined methods of GIS and image analysis to investigate the infestation probabilities at three types of forest edges under spatial and temporal aspects and compared them to the corresponding probabilities at the stand interior. Our results showed a pronounced infestation predisposition of such edge trees delimiting infestation patches cleared by sanitary logging measures, in particular at the south-facing edge sector. In contrast, edges adjacent to non-cleared infestation were revealed as less attractive for subsequent infestations, but nonetheless more attractive than permanent forest edges or the stand interior. Additionally, we measured near-bark surface air temperature to determine microclimatic differences at those edge- or non-edge sites and related them to predisposition results. Finally, our study emphasized favourable microclimatic conditions—summarized as the “sun-effect”—as a decisive factor enhancing the local infestation probability at recent forest edges in multiple ways. Both insect- and host tree-related reactions to suddenly altered microclimate are supposed to bias arbitrary colonization behaviour at patch and tree level, thereby mainly explaining observed infestation patterns. From the forester’s point of view, our results may contribute to precise bark beetle risk assessment and thus facilitate decision making in forest management.
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The characteristics of spruce individuals, which survived a massive bark beetle outbreak, were compared with the characteristics of neighbouring attacked trees in Šumava National Park (Czech Republic). Selected parameters related to crown geometry, stand conditions and distances between trees were measured or estimated. Significant differences were found between the surviving trees and the neighbouring trees attacked by I. typographus. Trees with a higher level of stem shading (longer crown length) tended to survive. The attacked trees were usually located in areas with larger basal area, especially southwards from them. A shorter distance to a previously attacked tree increased the probability of additional attack. Spruce trees with more progressive crown structure transformation (primary structure defoliation) were significantly more frequently attacked by spruce bark beetle. Superior and taller trees had a clearly longer life expectancy than dominant ones. These results show that the attack of trees by bark beetle can be predicted to a certain degree, which can be used in management of endangered spruce forests.
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In densely populated regions, forests can help protect communities and infrastructures from natural hazards such as avalanches and rockfall. To promote the protective function, substantial efforts are made to actively manage forest stands. In 2009 alone the Swiss government invested more than 60 million sfr for the maintenance of protection forests. However, to date there has been no comprehensive evaluation of how the structural development of actively managed stands differs from that of passively managed stands in the Alps. Over the past century the structure of Norway-spruce dominated subalpine forests of the Swiss Alps has been changing and it is not clear how these changes affect the potential protective function of these forests, as well as other forest functions such as wildlife habitat. Furthermore, it is not clear how stand dynamics and structural changes differ between stands that are actively managed and those that are passively managed, and thus to what degree active efforts of forest management are contributing to stands that actually have a greater protective function than passively managed forests.
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Keywords: individual-tree mortality model / logistic regression / mixed model / rebollo oak / Mediteranean oak Abstract • Tree mortality is an important process in forest ecosystem dynamics and is one of the least un-derstood phenomena, because of the complex interactions between different environmental stresses, minimal understanding of whole-plant mortality processes, and a chronic shortage of data. • A multilevel logistic regression model was developed for predicting the probability of mortality in individual trees with the objective of improving long-term planning in Spanish pyrenean oak forests. The data came from one 10-year re-measurement of the permanent plot network belonging to the Spanish National Forest Inventory distributed throughout north-west Spain. • The probability of mortality decreased with increasing individual diameter at breast height and in-creasing ratio of the height of subject tree to the dominant height of the sample plot. The resulting mortality model was evaluated using an independent data set from a region close to the study area. • The regeneration of pyrenean oak generally takes place through stump and/or root sprouting; so stand dynamics differ from those of others species. The model developed is expected to improve the accuracy of stand forecasts in northwest Spain.
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1Disturbance histories derived from old-growth forest remnants in Europe and eastern North America have shaped many of our current theories of forest dynamics and succession. Yet the small size typical of these remnants suggests they might not capture the full range of variability that may emerge at larger scales.2We investigated the frequency and severity of natural disturbance in a 2000-ha old-growth landscape (Big Reed Forest Reserve) in northern Maine, USA. Given its size, the Reserve provides an ideal opportunity to study, at multiple scales, natural forest processes in a region that has otherwise been dramatically altered by human activities. Using dendrochronological methods, we reconstructed disturbance histories for 37 randomly located plots stratified by five forest types (hardwood forests, mixed woods forests, red spruce forests, northern white-cedar seepage forests and northern white-cedar swamps).3We found no evidence of stand replacing disturbance on any plot during the last 120–280 years (depending on plot). The overall mean disturbance rate was 9.6% canopy loss per decade (median 6.5%, maximum 55%, plots pooled), yet the distribution was strongly skewed toward the lower rates.4We found little differences in disturbance rates between forest types, save a slightly lower rate in the northern white-cedar swamps. However, if we ignore forest-type classifications, we see that disturbance rates are clearly influenced by gradients in the relative abundance of component tree species, owing to species’ relative susceptibilities to particular disturbance agents.5Synthesis. Relatively low rates of canopy disturbance allow the accrual of shade-tolerant saplings. The abundance of this advance regeneration, coupled with the absence of stand-replacing disturbance, has maintained canopy dominance by shade-tolerant species in all plots, all forest types and throughout the entire landscape. Disturbance histories from individual plots coalesce to form a picture of a landscape in which pulses of moderate-severity disturbance are interposed upon a background of scattered small-scale canopy gaps. The landscape-level mosaic resulting from this disturbance regime consists of patches in various stages of structural development, not various stages of compositional succession.
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Forest gap models, initially conceived in 1969 as a special case of individual-tree based models, have become widely popular among forest ecologists for addressing a large number of applied research questions, including the impacts of global change on long-term dynamics of forest structure, biomass, and composition. However, they have been strongly criticized for a number of weaknesses inherent in the original model structure. In this paper, I review the fundamental assumptions underlying forest gap models, the structure of the parent model JABOWA, and examine these criticisms in the context of the many alternative formulations that have been developed over the past 30 years.Four assumptions originally underlie gap models: (1) The forest is abstracted as a composite of many small patches of land, where each can have a different age and successional stage; (2) patches are horizontally homogeneous, i.e., tree position within a patch is not considered; (3) the leaves of each tree are located in an indefinitely thin layer (disk) at the top of the stem; and (4) successional processes are described on each patch separately, i.e., there are no interactions between patches. These simplifications made it possible to consider mixed-species, mixed-age forests, which had been difficult previously mainly because of computing limitations.The structure of JABOWA is analysed in terms of the functional relationships used for formulating the processes of tree establishment, growth, and mortality. It is concluded that JABOWA contains a number of unrealistic assumptions that have not been questioned strongly to date. At the same time, some aspects of JABOWA that were criticized strongly in the past years are internally consistent given the objectives of this specific model.A wide variety of formulations for growth processes, establishment, and mortality factors have been developed in gap models over the past 30 years, and modern gap models include more robust parameterizations of environmental influences on tree growth and population dynamics as compared to JABOWA. Approaches taken in more recent models that led to the relaxation of one or several of the four basic assumptions are discussed. It is found that the original assumptions often have been replaced by alternatives; however, no systematic analysis of the behavioral effects of these conceptual changes has been attempted to date.The feasibility of including more physiological detail (instead of using relatively simple parameterizations) in forest gap models is discussed, and it is concluded that we often lack the data base to implement such approaches for more than a few commercially important tree species. Hence, it is important to find a compromise between using simplistic parameterizations and expanding gap models with physiology-based functions and parameters that are difficult to estimate. While the modeling of tree growth has received a lot of attention over the past years, much less effort has been spent on improving the formulations of tree establishment and mortality, although these processes are likely to be just as sensitive to global change as tree growth itself. Finally, model validation issues are discussed, and it is found that there is no single data source that can reliably be used for evaluating the behavior of forest gap models; instead, I propose a combination of sensitivity analyses, qualitative examinations of process formulations, and quantitative tests of gap models or selected submodels against various kinds of empirical data to evaluate the usefulness of these models for assessing their utility for predicting the impacts of global change on long-term forest dynamics.
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Greenhouse gas emissions have significantly altered global climate, and will continue to do so in the future. Increases in the frequency, duration, and/or severity of drought and heat stress associated with climate change could fundamentally alter the composition, structure, and biogeography of forests in many regions. Of particular concern are potential increases in tree mortality associated with climate-induced physiological stress and interactions with other climate-mediated processes such as insect outbreaks and wildfire. Despite this risk, existing projections of tree mortality are based on models that lack functionally realistic mortality mechanisms, and there has been no attempt to track observations of climate-driven tree mortality globally. Here we present the first global assessment of recent tree mortality attributed to drought and heat stress. Although episodic mortality occurs in the absence of climate change, studies compiled here suggest that at least some of the world's forested ecosystems already may be responding to climate change and raise concern that forests may become increasingly vulnerable to higher background tree mortality rates and die-off in response to future warming and drought, even in environments that are not normally considered water-limited. This further suggests risks to ecosystem services, including the loss of sequestered forest carbon and associated atmospheric feedbacks. Our review also identifies key information gaps and scientific uncertainties that currently hinder our ability to predict tree mortality in response to climate change and emphasizes the need for a globally coordinated observation system. Overall, our review reveals the potential for amplified tree mortality due to drought and heat in forests worldwide.
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Climatic constraints on tree growth mediate an important link between terrestrial and atmospheric carbon pools. Tree rings provide valuable information on climate‐driven growth patterns, but existing data tend to be biased towards older trees on climatically extreme sites. Understanding climate change responses of biogeographic regions requires data that integrate spatial variability in growing conditions and forest structure. We analyzed both temporal (c. 1901‐2010) and spatial variation in radial growth patterns in 9 876 trees from fragments of primary Picea abies forests spanning the latitudinal and altitudinal extent of the Carpathian arc. Growth was positively correlated with summer temperatures and spring moisture availability throughout the entire region. However, important seasonal variation in climate responses occurred along geospatial gradients. At northern sites, winter precipitation and October temperatures of the year preceding ring formation were positively correlated with ring width. In contrast, trees at the southern extent of the Carpathians responded negatively to warm and dry conditions in autumn of the year preceding ring formation. An assessment of regional synchronization in radial growth variability showed temporal fluctuations throughout the 20th century linked to the onset of moisture limitation in southern landscapes. Since the beginning of the study period, differences between high and low elevations in the temperature sensitivity of tree growth generally declined, while moisture sensitivity increased at lower elevations. Growth trend analyses demonstrated changes in absolute tree growth rates linked to climatic change, with basal area increments in northern landscapes and lower altitudes responding positively to recent warming. Tree growth has predominantly increased with rising temperatures in the Carpathians, accompanied by early indicators that portions of the mountain range are transitioning from temperature to moisture limitation. Continued warming will alleviate large‐scale temperature constraints on tree growth, giving increasing weight to local drivers that are more challenging to predict. This article is protected by copyright. All rights reserved.
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Disentangling the importance of developmental vs. environmental drivers of variation in forest biomass is key to predicting the future of forest carbon sequestration. At coarse scales, forest biomass is likely to vary along major climatic and physiographic gradients. Natural disturbance occurs along these broad biophysical gradients, and depending on their extent, severity and frequency, could either amplify or dampen spatial heterogeneity in forest biomass. Here we evaluate spatial variation in the basal area of late-successional Picea abies (L./Karst.) forests across the Carpathian Mountain Range of central Europe and compare the roles of coarse-scale biophysical gradients and natural disturbances in driving that variation across a hierarchy of scales (landscapes, stands, and plots). We inventoried forest composition and structure, and reconstructed disturbance histories using tree cores collected from 472 plots nested within 30 late-successional stands, spanning the Carpathian Mountains (ap-proximately 4.5 degrees of latitude). We used linear mixed-effects models to compare the effect of disturbance regimes and site conditions on stand basal area at three hierarchical scales. We found that the basal area of late-successional Picea abies forests varied across a range of spatial scales, with climatic drivers being most important at coarse scales and natural disturbances acting as the primary driver of forest heterogeneity at fine scales. For instance, the stand-level basal area varied among landscapes, with the highest values (48-68 m 2 ha −1) in the warmer southern Carpathian Mountains, and lower values (37-52 m 2 ha −1 on average) in cooler areas of the eastern and western Carpathians. Finer-scale variation was driven by local disturbances (mainly bark beetle and windstorms) and the legacies of disturbances that occurred more than a century ago. Our findings suggest that warming could increase the basal area of northern sites, but potential increasing disturbances could disrupt these environmental responses.
Book
Many terms often used to describe old-growth forests imply that these forests are less vigorous, less productive and less stable than younger forests. But research in the last two decades has yielded results that challenge the view of old-growth forests being in decline. Given the importance of forests in battling climate change and the fact that old-growth forests are shrinking at a rate of 0.5% per year, these new results have come not a moment too soon. This book is the first ever to focus on the ecosystem functioning of old-growth forests. It is an exhaustive compendium of information that contains original work conducted by the authors. In addition, it is truly global in scope as it studies boreal forests in Canada, temperate old-growth forests in Europe and the Americas, and global tropical forests. Written in part to affect future policy, this eminently readable book is as useful for the scientist and student as it is for the politician and politically-interested layman.
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Natural disturbance regimes are changing substantially in forests around the globe. However, large-scale disturbance change is modulated by a considerable spatiotemporal variation within biomes. This variation remains incompletely understood particularly in the temperate forests of Europe, for which consistent large-scale disturbance information is lacking. Here our aim was to quantify the spatiotemporal patterns of forest disturbances across temperate forest landscapes in Europe using remote sensing data, and determine their underlying drivers. Specifically, we tested two hypotheses: (1) Topography determines the spatial patterns of disturbance, and (2) climatic extremes synchronize natural disturbances across the biome. We used novel Landsat-based maps of forest disturbances 1986-2016 in combination with landscape analysis to compare spatial disturbance patterns across five unmanaged forest landscapes with varying topographic complexity. Furthermore, we analyzed annual estimates of disturbance change for synchronies and tested the influence of climatic extremes on temporal disturbance patterns. Spatial variation in disturbance patterns was substantial across temperate forest landscapes. With increasing topographic complexity, natural disturbance patches were smaller, more complex in shape, more dispersed, and affected a smaller portion of the landscape. Temporal disturbance patterns, however, were strongly synchronized across all landscapes, with three distinct waves of high disturbance activity between 1986 and 2016. All three waves followed years of pronounced drought and high peak wind speeds. Natural disturbances in temperate forest landscapes of Europe are thus spatially diverse but temporally synchronized. We conclude that the ecological effect of natural disturbances (i.e., whether they are homogenizing a landscape or increasing its heterogeneity) is strongly determined by the topographic template. Furthermore, as the strong biome-wide synchronization of disturbances was closely linked to climatic extremes, large-scale disturbance episodes are likely in Europe's temperate forests under climate changes. This article is protected by copyright. All rights reserved.
Article
Tree mortality as a crucial element of natural forest dynamics is still a poorly understood process. Abrupt growth decreases are known to occur several years or decades before complete cessation of growth. Hence, identifying and linking these growth decreases to potential inciting factors such as drought and frost will improve our understanding of mortality processes. We analyzed nine Central European tree species including six coniferous species (Abies alba, Picea abies, Larix decidua, Pinus sylvestris, Pinus cembra, Pinus montana) and three broadleaved species (Fagus sylvatica, Quercus spp., Acer pseudoplatanus). Tree-ring data from 848 standing dead trees from 14 forest reserves all over Switzerland were sampled. We applied distributed lag non-linear models to relate abrupt growth decreases to drought and frost. The results indicate for many species that both drought and frost have a moderate to major impact on abrupt growth decreases prior to tree death. While late frost in spring may instantaneously result in sustained abrupt growth decreases in most species except Scots pine and mountain pine, severe drought over several months in spring may either show an immediate negative impact on growth, such as in beech, or feature negative reactions that are lagged by several years, such as in oak and Scots pine. Thus, extreme climatic conditions have an essential influence on abrupt growth decreases that finally result in tree death, although variability of the reactions within and among species is high.
Article
The drivers of background tree mortality rates – the typical low rates of tree mortality found in forests in the absence of acute stresses like drought – are central to our understanding of forest dynamics, the effects of ongoing environmental changes on forests, and the causes and consequences of geographical gradients in the nature and strength of biotic interactions. To shed light on factors contributing to background tree mortality, we analyzed detailed pathological data from 200,668 tree-years of observation and 3729 individual tree deaths, recorded over a 13-year period in a network of permanent forest plots in California's Sierra Nevada mountain range. We found that: (1) Biotic mortality factors (mostly insects and pathogens) dominated (58%), particularly in larger trees (86%). Bark beetles were the most prevalent (40%), even though there were no outbreaks during the study period; in contrast, the contribution of defoliators was negligible. (2) Relative occurrences of broad classes of mortality factors (biotic, 58%; suppression, 51%; and mechanical, 25%) are similar among tree taxa, but may vary with tree size and growth rate. (3) We found little evidence of distinct groups of mortality factors that predictably occur together on trees. Our results have at least three sets of implications. First, rather than being driven by abiotic factors such as lightning or windstorms, the “ambient” or “random” background mortality that many forest models presume to be independent of tree growth rate is instead dominated by biotic agents of tree mortality, with potentially critical implications for forecasting future mortality. Mechanistic models of background mortality, even for healthy, rapidly-growing trees, must therefore include the insects and pathogens that kill trees. Second, the biotic agents of tree mortality, instead of occurring in a few predictable combinations, may generally act opportunistically and with a relatively large degree of independence from one another. Finally, beyond the current emphasis on folivory and leaf defenses, studies of broad-scale gradients in the nature and strength of biotic interactions should also include biotic attacks on, and defenses of, tree stems and roots. This article is protected by copyright. All rights reserved.
Article
1.Unprecedented bark beetle outbreaks have been observed for a variety of forest ecosystems recently, and damage is expected to further intensify as a consequence of climate change. In Central Europe, the response of ecosystem management to increasing infestation risk has hitherto focused largely on the stand level, while the contingency of outbreak dynamics on large-scale drivers remains poorly understood.2.To investigate how factors beyond the local scale contribute to the infestation risk from Ips typographus (Col., Scol.) we analysed drivers across seven orders of magnitude in scale (from 103 to 1010 m²) over a 23 year period, focusing on the Bavarian Forest National Park. Time-discrete hazard modelling was used to account for local factors and temporal dependencies. Subsequently, beta regression was applied to determine the influence of regional and landscape factors, the latter characterized by means of graph theory.3.We found that in addition to stand variables large-scale drivers also strongly influenced bark beetle infestation risk. Outbreak waves were closely related to landscape-scale connectedness of both host and beetle populations as well as to regional bark beetle infestation levels. Furthermore, regional summer drought was identified as an important trigger for infestation pulses. Large-scale synchrony and connectivity are thus key drivers of the recently observed unprecedented bark beetle outbreak in the area.4.Synthesis and applications. Our multi-scale analysis provides evidence that the risk for biotic disturbances is highly dependent on drivers beyond the control of traditional stand-scale management. This finding highlights the importance of fostering the ability to cope with and recover from disturbance. It furthermore suggests that a stronger consideration of landscape and regional processes is needed to address changing disturbance regimes in ecosystem management.This article is protected by copyright. All rights reserved.
Article
This study investigates temporal shifts in Norway spruce (Picea abies) mortality, stand structure characteristics, and stand complexity facilitated by a bark beetle (Ips typographus) outbreak that affected an unmanaged subalpine forest region in Tatra National Park, Poland in the late 2000s. Changes in survivorship and stand structure characteristics (diameter (DBH), basal area, height, age, and crown length ratio) of nearly 2500 spatially-referenced trees located in 64 long-term survey plots were compared over four time periods that spanned the duration of the outbreak disturbance event. Stand structure characteristics, topographic factors (slope, elevation, and aspect), and solar equinox radiation were tested as predictors of mortality for multiple stages in the outbreak using boosted regression tree modeling. Our findings showed that: (1) spatial synchrony was not reflective of mortality severity; (2) mortality rates increased significantly as the outbreak progressed; (3) the stand’s structure was altered significantly by the outbreak (larger trees were killed most frequently); (4) stand structure characteristics were the best predictors of mortality in all stages of the outbreak, though topographic factors and solar equinox radiation also exhibited moderate to strong predictive power in some stages; and (5) stand complexity decreased significantly as the outbreak progressed. This illustrates the inherently complex nature of bark beetle outbreaks on fine spatial scales and suggests that the extent and severity of spruce mortality during an outbreak event is largely dependent on the relative stage of the outbreak and the structure of the stand.
Article
Dynamic models are pivotal for projecting forest dynamics in a changing climate, from the local to the global scale. They encapsulate the processes of tree population dynamics with varying resolution. Yet, almost invariably, tree mortality is modeled based on simple, theoretical assumptions that lack a physiological and/or empirical basis. Although this has been widely criticized and a growing number of empirically derived alternatives are available, they have not been tested systematically in models of forest dynamics. We implemented an inventory-based and a tree-ring-based mortality routine in the forest gap model ForClim v3.0. We combined these routines with a stochastic and a deterministic approach for the determination of tree status (alive vs. dead). We tested the four new model versions for two Norway spruce forests in the Swiss Alps, one of which was managed (inventory time series spanning 72 years) and the other was unmanaged (41 years). Furthermore, we ran long-term simulations (∼400 years) into the future under three climate scenarios to test model behavior under changing environmental conditions. The tests against inventory data showed an excellent match of simulated basal area and stem numbers at the managed site and a fair agreement at the unmanaged site for three of the four empirical mortality models, thus rendering the choice of one particular model difficult. However, long-term simulations under current climate revealed very different behavior of the mortality models in terms of simulated changes of basal area and stem numbers, both in timing and magnitude, thus indicating high sensitivity of simulated forest dynamics to assumptions on tree mortality. Our results underpin the potential of using empirical mortality routines in forest gap models. However, further tests are needed that span other climatic conditions and mixed forests. Short-term simulations to benchmark model behavior against empirical data are insufficient; long-term tests are needed that include both nonequilibrium and equilibrium conditions. Thus, there is the potential to greatly improve the robustness of future projections of forest dynamics via more reliable tree mortality submodels.
Article
Climate conditions and forest structure interact to determine the extent and severity of bark beetle outbreaks, yet the relative importance of each may vary though the course of an outbreak. In 2008, we conducted field surveys and reconstructed forest conditions at multiple stages within a recent mountain pine beetle (MPB) outbreak in Rocky Mountain National Park, Colorado. At each stage in the outbreak, we examined changes in (1) lodgepole pine mortality and surviving stand structure, (2) the influence of topographic versus stand structure variables on mortality rates, and (3) stand complexity and landscape heterogeneity. Lodgepole pine mortality reduced basal area by 71 %, but only 47 % of stems were killed. Relative to pre-outbreak stands, surviving stands had lower mean dbh (11.0 vs. 17.4 cm), lower basal area (8.5 vs. 29.3 m2 ha−1), lower density (915 vs. 1,393 stems ha−1), and higher proportions of non-host species (23.1 vs. 10.6 % m2 ha−1). Factors predicting mortality rates changed through the course of the outbreak. Tree mortality during the early stage of the outbreak was associated with warm, dry sites and abundant large trees. During the middle and late stages, mortality was associated with stand structure alone. Stand complexity increased, as defined by stand-scale variability in density, basal area, and the proportion of susceptible trees. Landscape heterogeneity decreased according to semi-variograms of tree diameter and basal area. Increased stand complexity may inhibit future MPB population development, but decreased landscape heterogeneity may facilitate outbreak spread across the landscape if a future outbreak were to irrupt.
Article
Spruce bark beetle outbreaks are common in Norway spruce forests following windstorm damage, due to ample availability of brood material. The realization of an outbreak depends on factors regulating the Ips typographus population dynamics, such as weather conditions and salvage cutting. In this study, we take an ecosystem modelling approach to analyse the influence of multiple environmental factors on the risk for I. typographus outbreaks. Model calculations of I. typographus phenology and population dynamics as a function of weather and brood tree availability were developed and implemented in the LPJ-GUESS ecosystem modelling framework. The model simulations were driven by gridded climate data covering Sweden with a spatial resolution of 0.5° and a daily temporal resolution. Records on storm damage and I. typographus outbreak periods in Sweden for the period of 1960–2009 were used for model evaluation, and a sensitivity analysis was performed to examine the model behaviour. The model simulations replicated the observed pattern in outbreak frequency, being more common in southern and central Sweden than in northern Sweden. A warmer climate allowing for more than one generation per year can increase the risk for attacks on living trees. The effect of countermeasures, aiming at either reduce the availability of brood material or the I. typographus population size, is dependent on a non-linear relation between I. typographus attack density and reproductive success. The sensitivity analysis indicated a major reduction in the risk of attacks on living trees by timely salvage cutting and cutting of infested trees. Knowledge uncertainties associated with attacks on standing trees, i.e. factors influencing tree defence capacity and I. typographus reproductive success, should be further addressed.
Article
a b s t r a c t Correct knowledge of disturbance ecology is essential for understanding the characteristic behavior of forest ecosystems and for guiding appropriate management strategies. However, the role of natural dis-turbances in shaping European mountain forest ecosystems has not been adequately studied, possibly because of the perception that the development of most European forests is primarily shaped by human influences and/or fine-scale gap-phase dynamics. In the present study, we investigate the long-term disturbance history of old protected forest dominated by Norway spruce in the Parangalitsa Reserve, Bulgaria. We used aerial photo interpretation and dendro-ecological methods to reconstruct the history of wind, insect, and fire disturbances across a topographically complex landscape. Over the past 150 years wind has been the most important disturbance agent in this ecosystem and at least 18% of the forested area shows evidence of high-severity blowdowns. Windthrow patches ranged in size from <1 ha to >10 ha (minimum 0.11 ha, mean 0.16 ha, maximum 10 ha). Although small disturbances were much more frequent, few larger blowdowns accounted for most of the disturbed area. Pure coniferous and single-cohort coniferous forest patches were more affected by blowdowns than mixed coniferous–deciduous and multi-cohort coniferous forest patches. Although bark beetle (Ips typhog-raphus) populations were large enough to cause mortality of some live trees, the populations did not grow to epidemic proportions during recent decades. Fire disturbance was of limited importance in the last 200 years and only two patches (4% of the study area) showed evidence of fire. The present research indicates that wind disturbances have been characteristic of these ecosystems at least over the past decades to centuries. Thus, blowdowns appear integral to the normal function and struc-ture of the Picea-dominated mountain forests in the region and such events, in and of themselves, do not represent unhealthy forest conditions or environmental emergencies. Management strategies that aim to maintain these ecosystems within a natural range of variation should incorporate wind disturbances into the management strategy. The frequency and magnitude of future wind disturbances may be considered within the historical framework described in the current study to assess potential effects of climate change on altered disturbance regimes.
Article
Several methods (discriminant analysis, Probit analysis, Logit analysis) were compared for the prediction of individual overstory tree mortality in northern hardwood stands in Wisconsin. A generalized form of the logistic equation provided the greatest discriminating power for predicting live and dead trees. Independent variables were tree diameter and diameter increment, competition index, and length of growth period. Forest Sci. 22:438-444.
Article
Abstract 1 To maintain biodiversity in forests more wind-felled trees must be left where they fall. However, there is concern among forest owners that this may result in higher tree mortality caused by the spruce bark beetle, Ips typographus (L.) (Col.: Scolytidae). 2 In the 5 years following a major storm disturbance the number of standing spruces killed by I. typographus was determined in a total of 53 stands. In five of the stands all wind-thrown trees were left (unmanaged stands) and in 48 of the stands, which were situated at distances of 1.4–10.0 km from each focal unmanaged stand, the wind-felled trees were removed directly after the storm (managed stands). In the winter preceding the fifth summer new storm-fellings occurred in the study area. 3 In the 4-year period between the first and second storm-fellings, 50–322 standing trees were killed by I. typographus per unmanaged stand. There was a direct linear relationship between the number of storm-felled spruces colonized by I. typographus and the number of trees subsequently killed in the unmanaged stands. 4 Tree mortality caused by I. typographus in the unmanaged stands was almost nil in the first year, peaked in the second or third year, and decreased markedly to a low level in the fourth and fifth years. 5 In the 4-year period between the first and second storm-fellings twice as many trees were killed per ha in the unmanaged stands than in the managed stands: the average difference being 6.2 killed trees per ha, equivalent to 19% of the number of spruce trees felled by the first storm in the unmanaged stands. 6 Much higher numbers of trees were killed per ha in the stand edges than in the interiors of both the unmanaged and the managed stands.
Article
The intensity of bark beetle Ips typographus L. (Col., Scolytidae) attack on Norway spruce (Picea abies Karst.) is known to vary greatly among stands. In a control strategy approach, previous studies investigated the relationships between the variability in intensity of I. typographus attack and site characteristics such as stand age and altitude, mean tree circumference, growth rate and nearest-neighbour distance, soil moisture, pH in H2O and KCl, and soil contents of C, N, K, P, Mg, Ca, Fe, Cu, Zn and Mn. The data analysis method used in these studies was mainly the multiple linear regression, with the mean number of attacks per spruce tree in a stand as variable to explain. Previous results showed that the expected vulnerability of a Norway spruce stand to attack by I. typographus can be estimated on the basis of simple information of easy access to the forester, when the data on the stand in question is used with others for fitting the regression model. Prediction of the vulnerability of a stand, without including its data in the fitting of the model, was shown to be more approximate. Therefore, the objectives of this study were: (1) to improve the performance of models predicting the vulnerability of Norway spruce stands to attack by I. typographus, based on site characteristics; (2) to assess the stability of such predictive models when these are built using a moderate number of stands; and (3) to incorporate the resulting information in a global approach to control and prevention. Published data were re-analysed for these purposes. A jackknifed multiple linear regression procedure, in which each stand in turn is discarded when fitting the model (jackknife replication), is presented. A great variability in the models fitted, depending on the stand discarded, is observed. For instance, the number of explanatory variables retained ranges from one (i.e. soil P content, for five jackknife replications) to 10 (for one jackknife replication), for R2-values ranging from 0.5 to 1.0 and for one influential stand (i.e. the same stand characterized by an atypically low number of insect attacks compared to other stands with similar soil P content) against many influential stands. Differences between the model finally selected here using the revisited data and the models proposed earlier are discussed. A path analysis diagram is proposed for a more comprehensive modelling of Norway spruce stand vulnerability to I. typographus attack, based on site characteristics.
Article
Montane Norway spruce forests of Central Europe have a very long tradition of use for timber production; however, recently there has been increasing concern for their role in maintaining biological diversity. This concern, coupled with recent severe windstorms that led to wide-spread bark beetle outbreaks, has brought the management of montane spruce forests to the forefront of public policy discussions in Central Europe. In order to shed light on the natural development and current structure of mature montane spruce forests, we established four 0.25 ha research plots in a semi-natural montane spruce forest in the Šumava Mountains (The Bohemian Forest), Czech Republic. We mapped all trees, extracted increment cores for age and growth-pattern analyses, and inventoried all current tree regeneration, including the substrates on which it was found. Stands were characterized by uni-modal tree diameter distributions and high basal areas (56.6 m2 ha−1 on average), indicating a natural transition from the stem exclusion phase towards the understory reinitiation phase. The stands showed largely single-cohort recruitment age structures, however, with recruitment spanning seven decades. Our analyses suggest that this cohort existed as advance regeneration prior to major disturbances in the late 1800s, which included post-bark beetle salvage logging. Spatial pattern analyses of living and dead stems combined, showed an increase in uniformity of living trees, pointing to the role of natural density-dependent mortality. However, past growth patterns and historical documentation suggest that low intensity canopy disturbances (wind and snow) also caused mortality and diversified canopy structure. Because the stands developed naturally over the past 120+ years and thus escaped thinning operations, high volumes of coarse woody debris (94 m3 ha−1) and snag densities (546 stems ha−1) have accrued. Advance spruce regeneration was quite abundant and existed primarily on deadwood substrates, even though these occupied only a small percent of stand area. Because of salvage logging in the late 1880s, these stands do not qualify, according to the traditional paradigm, as natural spruce forests. As a result, they are recently subject to active management practices including salvage logging that remove dead and dying trees. Given the importance of deadwood for forest regeneration and recovery from disturbance, as demonstrated in this study, we argue that dead wood removal may limit future natural regeneration in these stands. Thus, the purported benefits of removing dead and dying trees from semi-natural forests must be carefully weighed against the potential detrimental impacts on natural spruce forest regeneration and biodiversity.
Article
Summary 1 An old-growth mesic forest in northern Michigan, USA, experienced an unusually intense storm in July 2002. Permanent inventory plots and a 2.9-ha mapped stand allow comparison of effects of this rare disturbance with patterns of 'baseline' mortality. 2 Tree mortality attributable to immediate effects of the storm was of similar magnitude to mortality over the previous decade, amounting to about 9% of basal area and 7% of stem density (mean of 88 inventory plots). 3 Storm mortality differed from baseline mortality in patterns related to species and size. Tsuga canadensis suffered little mortality of either type. Betula alleghaniensis had high baseline mortality in all size classes, but very low storm mortality. Acer spp. and Fagus grandifolia showed similar overall levels of baseline and storm mortality, but storm mortality was higher for larger stems. 4 Spatial patterns of mortality differed between baseline and storm-caused disturbance, with storm mortality patterns related to composition and to substrate variation over the stand. Baseline canopy mortality was hyperdispersed at local scales, but mortality due to the storm was strongly clustered at distances up to 30 m. 5 Properties of rare, intermediate disturbances cannot be predicted by simply scaling up patterns due to frequent, less intense, events. They may have distinctive influences on community dynamics, countering trends towards dominance by shade-tolerant species, generating demographic and spatial structure in the canopy, and distinctively affecting understorey environment. 6 These results suggest that climate change may induce changes in forest ecosystems by changing disturbance patterns even when species are not close to limits of physiological tolerance. Forest management for natural regeneration may need to take the effects of rare, intermediate disturbance into account.
Article
Despite extensive forest destruction in the Middle Ages and later intensive commercial forest management, remnants of virgin forests remained spared in some Central, Eastern and South-Eastern European countries. These virgin forests are the last examples of original forests in this part of Europe. That is why their protection becomes an important issue of current European forestry and nature protection policy. But the knowledge about the location and the area of virgin forests in these countries is incomplete up till now. This article has the prime goal to present a conceptual framework what virgin forests might be (“A conceptual framework for defining of virgin forests” section). Based on this framework, a working methodology has been tested in Bulgaria and Romania (“Results of the two national projects in Romania and in Bulgaria” section and further). For this reason two projects have been carried out by the Royal Dutch Society of Nature Conservation (KNNV) in close co-operation with the Forestry Institutes in Romania and in Bulgaria. The results of these projects are described in general terms and further analysis in the future is necessary to describe specific features like forest structure and spatial heterogeneity of these forests. Based on the results of the inventory, principles of sustainable protection and management of the mapped virgin forests were defined and described in the research reports. The usefulness of the inventory became evident already during the EU pre-accession period of both countries while preparing the NATURA 2000 network. The remaining virgin forests of temperate Europe are an inexhaustible source of ecological information about biodiversity, structure, natural processes and overall functioning of undisturbed forest ecosystems. Their research will reveal information which can be used for ecological restoration of man-made forests which are degraded through intensive forestry practices over the last centuries. The last virgin forests of temperate Europe represent an irreplaceable part of the natural capital of Europe and are worth to be protected by law. Their last remnants in South-Eastern and Eastern Europe are endangered by commercial activities. A full inventory of remaining virgin forests in all countries of temperate Europe is a matter of highest urgency. A representative selection of virgin forest sites should be declared by UNESCO as World Heritage Sites. KeywordsForests-Virgin forests-Forest management-Forest protection-Nature protection-Natura 2000 network-Mapping habitats-Forest policy
Article
Mortality as a basic process should be an integral part of any model of woody plant dynamics. But progress in developing mortality algorithms is slow. The aim of this paper is to provide a state-of-the-art description of mortality algorithms in published plant growth models, their structure, and the problems they face. The subsequent objective is to assess how models attempt to solve the problems, and provide some insight on how modelling mortality can be improved. Sixty-one models, concerned with forest yield and dieback and forest/shrubland dynamics, were reviewed, the greatest proportion being the gap type. The algorithms could be broadly divided into stochastic and deterministic. Despite considerable variability, the basic premise of every algorithm was the same: beyond a specified threshold a plant either dies (deterministic), or has a greater probability of dying (stochastic). Factors used to predict mortality include size, age, competition, carbon balance (growth) and abiotic influences. Some algorithms were random, and one-third used more than one predictor. Although the classification is far from clear-cut, mortality algorithms can be placed in one or more of the following classes: carbon-based, abiotic/age-based, competitive, gap-type, statistically fitted and progressive-stress type. It is clear both that mortality is difficult to model, and that there is no best way to model it for all applications. The complexity of environmental stresses and lack of process information of woody plant mortality has however led to the wide use of empirical algorithms. These have many problems, including data intensity and spatial and temporal specificity. Such problems render mortality simulation prone to error and weak in response to environmental change. Test data and procedures also are lacking. In the literature suggested improvements to the problem include the use of mechanistic algorithms. But this approach also has associated problems. So here it is suggested here that the use of a biologically reasonable predictor in a simple stochastic algorithm, used within a mechanistic model of plant growth, is a preferable improvement. The approach provides a mechanistic-empirical consensus. A shift in emphasis towards modelling for exploration and explanation, rather than to predict with high levels of accuracy is also suggested.
Article
The outbreaks of Ips typographus (L.) in Central Europe after severe storms in the 1990s triggered extensive research. Molecular techniques were used to analyze the relations and origins of European Ips species. The biological characteristics of I. typographus such as the influence of temperature on life history parameters and flight behavior have been analyzed in detail. The spruce bark beetle was found to disperse well beyond 500 m. However, new attacks mostly occurred in the vicinity of old ones.Many studies refer to invertebrate natural enemies of I. typographus such as predatory beetles and flies as well as various parasitoids. While the species assemblages of antagonists have been extensively investigated their dynamics and impact on I. typographus populations are controversial.The susceptibility and defense mechanisms of host trees are crucial for a successful attack by bark beetles. Newly attacked trees respond with preformed resin, local wound reactions and eventually with systemic changes in their physiology. Risk assessments have been performed at both tree and stand level. Risk of attack seems to be mainly related to the exposition, age, and nutrient and water supply of the trees. The dynamics of outbreaks largely depends on insect abundance, tree susceptibility, weather conditions, and human measures. This renders predictions difficult. Various control techniques are reviewed and the need for more sophisticated risk assessment tools is stressed.
Article
In the 1990s, a bark beetle (Ips typographus) outbreak caused a dieback of semi-natural mountain spruce (Picea abies) forests in the Šumava National Park (Czech Republic). Two different approaches were applied to the attacked forests: (1) a small portion of the stands in the core zone of the national park was left without intervention, relying upon natural regeneration, and (2) traditional technical measures were adopted, in which attacked trees were felled and removed. Under the dead canopy of the stands left without intervention, there was a good regeneration of spruce and rowan (Sorbus aucuparia) and, sporadically, beech (Fagus sylvatica). In clear-cut areas, the numbers of spruce and rowan were significantly lower than under the dead canopy. Pioneer species such as willow (Salix aurita), birch (Betula pubescens), and aspen (Populus tremula) appeared in the clear-cut areas. The dependence of spruce regeneration on the availability of suitable microhabitats was found: decaying wood and spruce litter was found as the most favourable. The results confirmed that the original tree species of the mountain spruce forests regenerate well under dead canopy. The bark beetle outbreak does not result in the complete loss of the forests and could even be considered as a tool for the restoration of their natural character.
Article
Disturbances play a major role in forest ecosystems, they are main constituents of forest dynamics and are often a relevant driver in forest management decisions. The European Spruce Bark Beetle (Ips typographus L. Col. Scol.) is one of the major biotic disturbances in Norway spruce (Picea abies (L.) Karst.) forests. In this contribution a sub-model of disturbances by I. typographus was developed and integrated in the existing hybrid forest patch model PICUS v1.4. The new disturbance sub-model builds on a recently developed phenology model for risk assessment of outbreaks of I. typographus (PHENIPS) and elements from an existing predisposition assessment system (PAS). Model parameterisation was based on data from 28 Norway spruce stands in Austria. In a preliminary model evaluation a comprehensive sensitivity analysis at sites along an elevation gradient in the Eastern Alps is presented. Sensitivity over the elevation gradient was found to be highest and strongly non-linear with regard to the thermal environment. Furthermore, in accordance with general expectations and observations simulated damages were high under colline and submontane conditions and strongly decreased over the elevation transect. Based on the results of the simulation experiments it is concluded that the presented model is a promising tool to analyse the dynamic interaction of disturbances by I. typographus, environmental conditions and forest structure as affected by natural forest development and management interventions. Limitations of the model and possible approaches for extensive validation against empirical damage data are discussed.