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Abstract

Natural disturbances strongly influence forest structural dynamics, and subsequently stand structural heterogeneity, biomass, and forest functioning. The impact of disturbance legacies on current forest structure can greatly influence how we interpret drivers of forest dynamics. However, without clear insight into forest history, many studies default to coarse assumptions about forest structure, for example, whether forests are even or unevenly aged. The aim of this study was to analyze the effects of past disturbances on the current diameter distributions of Norway spruce (Picea abies (L.) Karst.)-dominated landscapes throughout the Carpathian Mountains. Our dendroecological dataset comprises tree cores from 339 plots (7,845 total tree cores), nested within 28 primary forest stands, known to vary greatly in the severity of historical disturbances. Our analyses revealed that historical disturbances had a strong and significant effect on the current diameter distribution shapes at the plot level. We demonstrated that mixed-severity disturbance regimes were more frequent and create a complex pattern of diameter distributions at the plot and stand scale. Here, we show that high severity disturbance was associated with unimodal diameter distributions, while low and moderate severity was associated with the reverse J-shaped distribution. This is a result of complex disturbance patterns, with structural biological legacies. Our results will have important management implication in the context of tree size heterogeneity, biomass storage, and productivity as influenced by natural disturbances. Lastly, these results demonstrate that structural changes may arise as consequences of changing disturbance regime associated with global change.

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... The DBH distribution of tree species exhibits variable patterns owing to site quality, regional climate, and disturbances (Chhin et al. 2008;Lin et al. 2016;Rodrigo et al. 2022). Generally, consistent with the results of previous studies fitting the similar DBH distribution of the corresponding tree species in Northeast China (Hao et al. 2022;Liu et al. 2014;Xu and Jin 2012), the overall DBH distribution of most tree species predicted in this study displayed a right-skewed unimodal pattern (Fig. 5), suggesting that at the regional scale, a continuous regeneration and relatively stable community structure occurred in most tree species in Northeast China (Fang et al. 2012). ...
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Given the global intensification of forest management and climate change, protecting and studying forests that develop free of direct human intervention-also known as primary forests-are becoming increasingly important. Yet, most countries still lack data regarding primary forest distribution. Previous studies have tested remote sensing approaches as a promising tool for identifying primary forests. However, their precision is highly dependent on data quality and resolution, which vary considerably. This has led to underestimation of primary forest abundance and distribution in some regions, such as the temperate zone of Europe. Field-based inventories of primary forests and methodologies to conduct these assessments are inconsistent; incomplete or inaccurate mapping increases the vulnerability of primary forest systems to continued loss from clearing and land-use change. We developed a comprehensive methodological approach for identifying primary forests, and tested it within one of Europe's hotspots of primary forest abundance: the Carpathian Mountains. From 2009 to 2015, we conducted the first national-scale primary forest census covering the entire 49,036 km 2 area of the Slovak Republic. We analyzed primary forest distribution patterns and the representativeness of potential vegetation types within primary forest remnants. We further evaluated the conservation status and extent of primary forest loss. Remaining primary forests are small, fragmented, and often do not represent the potential natural vegetation. We identified 261 primary forest localities. However, they represent only 0.47% of the total forested area, which is 0.21% of the country's land area. The spatial pattern of primary forests was clustered. Primary forests have tended to escape anthropogenic disturbance on sites with higher elevations, steeper slopes, rugged terrain, and greater distances from roads and settlements. Primary forest stands of montane mixed and subalpine spruce forests are more abundant compared to broadleaved forests. Notably, several habitat types are completely missing within primary forests (e.g., floodplain forests). More than 30% of the remaining primary forests are not strictly protected, and harvesting occurred at 32 primary forest localities within the study period. Almost all logging of primary forests was conducted inside of protected areas, underscoring the critical status of primary forest distribution in this part of Europe. Effective conservation strategies are urgently needed to stop the rapid loss and fragmentation of the remaining primary forests. Our approach based on precise, field-based surveys is widely applicable and transferrable to other fragmented forest landscapes.
Article
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.
Article
Theoretical and empirical studies have suggested that climate and soils are the main drivers of biodiversity, stand structure and aboveground biomass in natural forests. Yet, the direct effects of climate and soils on aboveground biomass versus the indirect effects mediated by species diversity and stand structural complexity remain unclear in forest ecosystems across large-scale ecological gradients. Here, we hypothesized that 1) climate and soils would influence aboveground biomass through strong indirect effects; 2) stand structural complexity rather than species diversity would strongly mediate the response of aboveground biomass to climate and soils; and 3) species diversity and stand structural complexity would promote each other under the niche differentiation and facilitation effects, and that stand structural complexity would have positive effects on aboveground biomass across large-scale ecological gradients. To test these hypotheses, we quantified climatic water availability, soil total exchangeable bases, species diversity, stand structural complexity including tree DBH (diameter at breast height) diversity and height diversity, and aboveground biomass across 907 plots in tropical forests of Hainan Island, Southern China. We tested 126 structural equation models to examine the direct and indirect effects of climate and soils on aboveground biomass via species diversity and stand structural complexity. Climatic water availability and soil fertility did not affect aboveground biomass directly but did affect indirectly via increasing stand structural complexity rather than species diversity. Species diversity and stand structural complexity promoted each other, and both increased with increasing climatic water availability. Stand structural complexity increased aboveground biomass directly, whereas species diversity increased it indirectly via increasing stand structural complexity. The total effects of climatic water availability, soil fertility, stand structural complexity and species diversity on aboveground biomass were significantly positive. This study shows that climatic water availability exerts a strong direct effect on stand structural complexity, indicating that any decrease in climatic water availability (i.e. increasing atmospheric drought) may directly diminish stand structural complexity and hence indirectly reduce aboveground biomass and carbon storage. This study suggests that maintaining high stand structural complexity can enhance aboveground biomass under favourable climate and soils while maintaining the benefits of species diversity on stand structural complexity for better ecosystem services such as carbon storage.
Article
Examines, in a variety of contexts, a number of theoretical and empirical relationships between disturbance (environmental fluctuations and destructive events, whether predictable and/or cyclical or not) and patch dynamics (where discrete spatial patterns possess internal characteristics and also inter-relate with surrounding patch and non-patch areas). The main sections are on: patch dynamics in nature; adaptations of plants and animals in a patch dynamic setting; and implications of patch dynamics for the organisation of communities and the functioning of ecosystems. A final chapter moves towards a general theory of disturbance. All 21 chapters are abstracted separately. -P.J.Jarvis
Book
Norway spruce (Picea abies L.) is an important tree species with a remarkable natural range throughout Europe and Asia, ranging from the Balkan Peninsula to Siberia in the north and from the French Alps in the west to the Sea of Okhotsk in the east. Wherever it occurs, it is a key component of both natural and managed forests. Norway spruce is the most economically valuable conifer in Europe, producing high-quality timber and wood products. This book presents a concise and comprehensive review of the biology, ecology, and management of Norway spruce. It integrates classic and contemporary literature (more than 2000 works cited in the text), highlighting basic research and forestry practices in central and eastern Europe. The topics include anatomy and morphology, physiology and nutrition, reproductive biology and genetics, and ecology. In addition, it examines mycorrhiza, diseases and pests as well as silviculture and wood products. In the light of increasing threats to forest health from air pollution, climate change, and insects and disease, it provides an essential information source to those concerned with the ecology, conservation, and management of the species.
Article
Mixed-severity disturbance regimes are prevalent in temperate forests worldwide, but key uncertainties remain regarding the variability of disturbance-mediated structural development pathways. This study investigates the influence of disturbance history on current structure in primary, unmanaged Norway spruce (Picea abies) forests throughout the Carpathian Mountains of central and eastern Europe, where windstorms and native bark beetle outbreaks are the dominant natural disturbances. We inventoried forest structure on 453 plots (0.1 ha) spanning a large geographical gradient (\>1,000 km), coring 15–25 canopy trees per plot for disturbance history reconstruction (tree core total n = 11,309). Our specific objectives were to: (1) classify sub-hectare-scale disturbance history based on disturbance timing and severity; (2) classify current forest structure based on tree size distributions (live, dead, standing, downed); (3) characterize structural development pathways as revealed by the association between disturbance history and current forest structural complexity. We used hierarchical cluster analysis for the first two objectives and indicator analysis for the third. The disturbance-based cluster analysis yielded six groups associated with three levels of disturbance severity (low, moderate, and high canopy loss) and two levels of timing (old, recent) over the past 200 years. The structure-based cluster analysis yielded three groups along a gradient of increasing structural complexity. A large majority of plots exhibited relatively high (53\%) or very high (26\%) structural complexity, indicated by abundant large live trees, standing and downed dead trees, and spruce regeneration. Consistent with conventional models of structural development, some disturbance history groups were associated with specific structural complexity groups, particularly low-severity/recent (very high complexity) and high-severity/recent (moderate complexity) disturbances. In other cases, however, the distribution of plots among disturbance history and structural complexity groups indicated either divergent or convergent pathways. For example, multiple disturbance history groups were significantly associated with the high complexity group, demonstrating structural convergence. These results illustrate that complex forest structure – including features nominally associated with old-growth – can be associated as much with disturbance severity as it is with conventional notions of forest age. Because wind and bark beetles are natural disturbance processes that can induce relatively high levels of tree mortality while simultaneously contributing to structural complexity and heterogeneity, we suggest that forest management plans allow for the stochastic occurrence of disturbance and variable post-disturbance development trajectories. Such applications are especially appropriate in conservation areas where biodiversity and forest resilience are management objectives, particularly given projections of increasing disturbance activity under global change.
Article
In order to gauge ongoing and future changes to disturbance regimes, it is necessary to establish a solid baseline of historic disturbance patterns against which to evaluate these changes. Further, understanding how forest structure and composition respond to variation in past disturbances may provide insight into future resilience to climate-driven alterations of disturbance regimes.
Article
Mountain forests are among the most important ecosystems in Europe as they support numerous ecological, hydrological, climatic, social, and economic functions. They are unique relatively natural ecosystems consisting of long-lived species in an otherwise densely populated human landscape. Despite this, centuries of intensive forest management in many of these forests have eclipsed evidence of natural processes, especially the role of disturbances in long-term forest dynamics. Recent trends of land abandonment and establishment of protected forests have coincided with a growing interest in managing forests in more natural states. At the same time, the importance of past disturbances highlighted in an emerging body of literature, and recent increasing disturbances due to climate change are challenging long-held views of dynamics in these ecosystems. Here, we synthesize aspects of this Special Issue on the ecology of mountain forest ecosystems in Europe in the context of broader discussions in the field, to present a new perspective on these ecosystems and their natural disturbance regimes. Most mountain forests in Europe, for which long-term data are available, show a strong and long-term effect of not only human land use but also of natural disturbances that vary by orders of magnitude in size and frequency. Although these disturbances may kill many trees, the forests themselves have not been threatened. The relative importance of natural disturbances, land use, and climate change for ecosystem dynamics varies across space and time. Across the continent, changing climate and land use are altering forest cover, forest structure, tree demography, and natural disturbances, including fires, insect outbreaks, avalanches, and wind disturbances. Projected continued increases in forest area and biomass along with continued warming are likely to further promote forest disturbances. Episodic disturbances may foster ecosystem adaptation to the effects of ongoing and future climatic change. Increasing disturbances, along with trends of less intense land use, will promote further increases in coarse woody debris, with cascading positive effects on biodiversity, edaphic conditions, biogeochemical cycles, and increased heterogeneity across a range of spatial scales. Together, this may translate to disturbance-mediated resilience of forest landscapes and increased biodiversity, as long as climate and disturbance regimes remain within the tolerance of relevant species. Understanding ecological variability, even imperfectly, is integral to anticipating vulnerabilities and promoting ecological resilience, especially under growing uncertainty. Allowing some forests to be shaped by natural processes may be congruent with multiple goals of forest management, even in densely settled and developed countries.
Article
Disturbances shape forest structure and composition, but the temporal dynamics of disturbance patterns, their influence on dynamics of forest structural complexity, and the potential impacts of ongoing climate changes are not fully understood. We addressed these issues by focusing on (1) long-term, landscape level retrospective analysis of disturbance dynamics of mountain Norway spruce (Picea abies) forest; (2) testing for the prevailing disturbance agent; and (3) the detection of disturbance drivers, particularly site conditions, using a dendrochronological approach.
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
Forest structure is both a product and driver of ecosystem processes and biological diversity. It has become apparent in recent years that changes in forest structure as a result of management for timber production have undesirable consequences for other components of forest ecosystems. The objective of this paper is to provide an overview of what we have learned about the ecological roles of forest structure in the Pacific Northwest and how forest structure changes as a result of disturbance and succession. Forests are structurally diverse, but many structures derive from the same processes of disturbance and growth. Consequently, measurements on a few structural attributes can be used to estimate many other structural conditions. Particularly important components of forest structure include live-tree sizes, vertical foliage distributions, horizontal variation in canopy density, and coarse woody debris. Knowledge of the ecological roles of these structures has increased in recent years and we now have a general understanding of how these structures change during succession. Although the ecological values of forest structures are now more widely appreciated, we still have many significant knowledge gaps including the ecological roles of below-ground structure, woody debris, and landscape pattern.
Article
Windthrow is all too often looked at as an exceptional, catastrophic phenomenon rather than a recurrent natural disturbance that falls within the spectrum of chronic and acute effects of wind on forests, and that drives ecosystem patterns and processes. This paper provides an integrative overview of the nature, contributing factors and impacts of wind-caused disturbance in forests, including its effects on trees, stands, landscapes and soils. Windthrow is examined through the integrating concepts of: the capacity of trees for acclimative growth, the limitation of acclimative growth under inter-tree competition, the recurrent nature of severe weather, how terrain and soil conditions affect local stand vulnerability and the effect of recurrent windthrow on stand dynamics and soils. Windthrow management should take place within a framework of general risk management, with evaluation of the likelihood, severity and potential impacts of wind damage considered with reference to the broad and specific aims of management. There is much to be gained from interdisciplinary communication about the nature and consequences of recurrent wind damage. There are opportunities for climatologists, engineers, ecologists, geomorphologists and others to develop integrative process models at the tree, stand and landscape scales that will improve our collective understanding, and inform management decision-making.
Article
Disturbances from wind, bark beetles, and wildfires have increased in Europe's forests throughout the 20(th) century (1). Climatic changes were identified as a main driver behind this increase (2), yet how the expected continuation of climate change will affect Europe's forest disturbance regime remains unresolved. Increasing disturbances could strongly impact the forest carbon budget (3,4), and are hypothesized to contribute to the recently observed carbon sink saturation in Europe's forests (5). Here we show that forest disturbance damage in Europe has continued to increase in the first decade of the 21(st) century. Based on an ensemble of climate change scenarios we find that damage from wind, bark beetles, and forest fires is likely to increase further in coming decades, and estimate the rate of increase to +0.91·10(6) m(3) of timber per year until 2030. We show that this intensification can offset the effect of management strategies aiming to increase the forest carbon sink, and calculate the disturbance-related reduction of the carbon storage potential in Europe's forests to be 503.4 Tg C in 2021-2030. Our results highlight the considerable carbon cycle feedbacks of changing disturbance regimes, and underline that future forest policy and management will require a stronger focus on disturbance risk and resilience.
Article
For many types of forest studies, it is essential to identify the exact years of formation of annual rings in increment cores taken from living trees. To accomplish this, dendrochronologists employ cross dating, which involves both ring counting and ring-width pattern matching, to ensure against counting error, or errors, caused by missing or false rings. To date, published accounts of the cross-dating process generally describe a graphical method for achieving cross dating, known as skeleton plotting. However, when working with cores from living trees, skeleton plotting is seldom necessary. Such cores can commonly be cross-dated more quickly and easily by listing the narrow rings that are present in each core in a laboratory notebook and then comparing core notes for shared narrow rings. The latter approach permits faster recognition of ring-width patterns because calendar-year, rather than relative-year, dates are assigned to rings in cores. It also allows cross-dating records to be stored in a more concise manner.
Article
Disturbances influence forest dynamics across a range of spatial and temporal scales. In tropical forests most studies have focused on disturbances occurring at small spatial and temporal scales (i.e., gap dynamics). This is primarily due to the difficulty of reconstructing long-term disturbance histories of forests in which most tree species lack annual growth rings. Consequently, the role of past disturbances in tropical forests is poorly understood. We used a combination of direct and indirect methods to reconstruct the his- torical disturbance regime and stand development patterns in mature and regenerating seasonal dry evergreen forest (SDEF) in the Huai Kha Khaeng Wildlife Sanctuary in western Thailand. Direct estimates of long-term establishment and growth patterns were obtained from 12 tree species that form annual growth rings as a consequence of the region's strong intra-annual rainfall seasonality. Indirect estimates of establishment patterns were obtained from analyses of stand structure and individual tree architecture and application of age- estimation models to 10 dominant canopy-tree species using demographic data from a large- scale, permanent forest-dynamics plot. The combination of direct and indirect methodologies revealed a complex disturbance history in the seasonal evergreen forest over the past 250 years. In the mid-1800s, 200-300 ha of forest were destroyed by a catastrophic disturbance, which led to the synchronous es- tablishment of many of the trees that presently dominate the forest canopy. Since then wide- spread disturbances of variable intensity have occurred at least three times (1910s, 1940s, and 1960s). These disturbances created discrete temporal pulses of establishment in small to large gaps in the forest matrix across several square kilometers. Background mortality and gap formation were evident in every decade since 1790, but these varied in intensity and frequency. The SDEF retains a distinct structural and floristic legacy from the catastrophic dis- turbance of the mid-1800s. The single-age cohort that established after the disturbance has developed a complex three-dimensional structure as a consequence of differences in in- terspecific growth patterns of the canopy-tree species and subsequent disturbances of mod- erate and low intensity. While no single methodological approach provided a complete picture of the disturbance history and stand development patterns of the seasonal evergreen forest, taken together they offered new insights into the long-term dynamics of a primary tropical forest. In particular, the study highlighted the role of disturbance at multiple spatial and temporal scales and varying intensities in determining the structure and composition of a complex, species-rich tropical forest and raises important questions about the role of rare, catastrophic events on tropical forest dynamics.
Article
The authors begin by outlining the role of the dendrochronologists both in the field and in the laboratory. The basic principles of tree-ring dating are then explained in detail, followed by a guide to the collection of archaeological and modern specimens from the field. The final section deals with the laboratory techniques used: the preliminary processing and preparation of archaeological and modern specimens; the process of dating specimens; and finally the compilation of a master chronology.
Article
The frequency of natural disturbances and their influence on the forest landscape mosaic were investigated on three large tracts of primary forest in Upper Michigan. Seventy 0.5-ha plots were randomly distributed in a total forest area of 23 000 ha dominated by sugar maple (Acer saccharum) and eastern hemlock (Tsuga canadensis). Radial increment patterns were used to estimate canopy accession dates for each of a number of randomly selected overstory trees on each plot. From these data a disturbance chronology, representing the percentage of stand area occupied by cohorts originating during each decade over the last 130 yr, was compiled for each plot. Average rates of disturbance or canopy mortality are estimated at 5.7 to 6.9% per decade. The corresponding average canopy residence time of a tree is 145-175 yr. No significant differences were detected in average disturbance rates among the three study areas, between plots near the coast of Lake Superior and inland plots, among several different aspects, and among several different slope positions. Natural rotation periods increase exponentially with increasing disturbance intensity, which is defined as the approximate percentage of the plot area converted to gaps during a disturbance episode. Estimates of rotation periods range from 69 yr for greater-than-or-equal-to 10% canopy removal to 1920 yr for greater-than-or-equal-to 60% canopy removal. Spatial autocorrelation analysis indicated that plots with light and medium disturbances (< 40%) are randomly distributed over the landscape. Plots with heavy disturbances (greater-than-or-equal-to 40%) are clustered with a patch radius of almost-equal-to 2 km, consistent with the sizes of thunderstorm downbursts. The data indicate that light and medium disturbances dominate the disturbance regime. The majority of stands on the landscape are composed of several major and many minor age classes. Even-aged stands with one predominant age class are uncommon. The age distribution of individual patches or cohorts in the two larger study areas (14 500 and 6073 ha) follows a nearly uniform distribution. None of the three study areas had more than 15% of the forest area converted to gaps in a single decade. The two larger areas meet most of the criteria that have been proposed for equilibrium landscapes.
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
The Weibull probability density function is proposed as a diameter distribution model. Its advantages include flexibility in shape and simplicity of mathematical derivations. Estimation and interpretation of parameters are discussed and illustrated with published data. Forest Sci. 19:97-104.
Article
• After storm disturbances, there is a risk for degradation of the quality of fallen trees, and for subsequent tree mortality caused by the spruce bark beetle Ips typographus (L.) (Coleoptera: Curculionidae). Models assessing the risk for bark beetle colonization of different kinds of storm gaps would be a valuable tool for management decisions. • The present study aimed to determine which gap and landscape characteristics are correlated with the probability of colonization of wind-felled Norway spruce trees by I. typographus. • The study included 36 storm gaps, varying in size from three to 1168 wind-felled spruces, created by the storm Gudrun in southern Sweden in January 2005. • In the first summer, on average, 5% of the wind-felled spruces were colonized by I. typographus. The percentage of colonized wind-felled trees per gap was negatively correlated with the total area of storm gaps within 2000 m in the surrounding forest landscape. • In the second summer, the proportion of colonized trees increased to 50%. Both gap (mean diameter of wind-felled trees and basal area of living spruce trees) and landscape variables (amount of spruce forest) were significantly correlated with colonization percentage and explained almost 50% of the variation between gaps. • There was no relationship between gap area and colonization percentage. This implies that landscapes with many large storm gaps, where logging resources will be most effectively used, should be salvaged first.
Article
1Tree-size distributions are changing in many natural forests around the world, and it is important to understand the underlying processes that are causing these changes. Here we use a classic conceptual framework – the shifting mosaic of patches model – to explore the ways in which competitive thinning and disturbance influence tree-size distributions, and to consider the effects of temporal variability in disturbance frequency on the size structure of forests.2We monitored 250 stands of Nothofagus solandri var. cliffortiodes (mountain beech), randomly distributed over 9000 hectares, for 19 years. Mountain beech is a light-demanding species that forms monospecific forests in New Zealand mountains. For the purposes of our model, we assumed that each stand functions as an even-aged population: it is initiated by a pulse of recruitment, undergoes competitive thinning as it matures, and is eventually destroyed by a disturbance event. The tree-size distribution of the whole forest is driven partly by the frequency and temporal patchiness of disturbance events and partly by competitive processes within the constituent stands.3Temporal changes in stem density and mean tree size were observed to be remarkably similar in all young stands, indicating that a consistent packing rule operates during this phase of stand development. A popular idea in the self-thinning literature is that the maintenance of constant leaf area index (LAI) provides the mechanism for this packing rule, but our analyses suggest that LAI increased by about 30% during the thinning phase. We use leaf economic theory to develop a new packing rule based on light interception, and argue that LAI increases with stand age because of changes in canopy organisation.4Smaller trees were significantly more likely to die than larger trees within the young stands. Tree-diameter distributions within young stands were left skewed but those of older populations were normally distributed. These observations are consistent with asymmetric competition winnowing out small, suppressed trees from young stands but having less effect in older stands.5Large-scale disturbances created gaps of sufficient size to allow mass recruitment of seedlings in about 0.8% of stands each year. Older stands were most susceptible to such large-scale disturbance, but the trend was weak.6The diameter-distribution of the whole Nothofagus forest was found to be approximately exponential in form. Simulation models only produced realistic diameter distributions when competitive packing rules and disturbance were included. Therefore, the shifting mosaic model provides a general framework for understand the ways in which these mortality processes determine forest size structure.7The diameter distribution of the forest was not in equilibrium over the 19-year study. Using simulation models, we show that temporal variability in disturbance frequency can generate enormous deviations in tree-diameter distributions away from the long-term mean, leading us to conclude that modern-day disequilibrium in natural forests may be the legacy of past disturbance events.
Article
The study presents an analysis of the diameter distributions of nine virgin beech forests (Fagus sylvatica) in south-eastern Europe. Data were collected from published and unpublished sources. We included predominantly full calliperings of coherent areas between 3.6 and 13.0 ha of size. The objective of the study was to compare and characterize the curve forms and to test systematically which mathematical function provides a better fit: the negative exponential function, third or seventh degree Weibull functions. The parameters were estimated applying the maximum likelihood method. To evaluate the goodness of fit, the absolute discrepancy was used. The residuals were examined with respect to systematic deviation.The nine virgin forests displayed a great variety of structures, only four out of 36 possible pairs of diameter distributions were found to be from the same population. The negative exponential model produced a good fit for four of the nine empirical distributions. However, the analysis of the residuals produced by the models exhibited systematic errors. Both the negative exponential and the third degree Weibull function underpredict to some extent the number of stems in the midsize diameter range in all nine stands. The higher number of trees frequently found in the midsize range of the empirical distributions as compared to the models indicates a common trend towards a rotated sigmoid diameter distribution. For some stands even a tendency towards two peaks in the distribution can be found. These results reveal that the reverse J-shaped curve form is not the only applicable model for describing diameter distributions in virgin beech forests. The systematic nature of the deviations from the negative exponential curve leads us to conclude that we might have found a general trend typical for virgin beech forests in south-eastern Europe.
Article
Summary • Many theoretical models have been proposed to explain the empirical self-thinning relationship given by Yoda et al. in 1963 for even-aged, monospecific stands of plants, but the models are inadequate to allow consensus on the processes driving variation in density-dependent mortality and self-thinning. • Most non-individual based models (non-IBMs), and many IBMs, employ a common representation of competition in which a finite amount of potential crown area remains completely allocated throughout self-thinning, making stand density inversely proportional to mean projected crown area. • This representation entails four assumptions regarding the competition process: the population is adequately represented by the mean plant; total stand resource utilization is constant throughout self-thinning; competition is a horizontal packing process; and differences in initial stand conditions may affect the rate of competition but not the process itself. • Reviewing published empirical data, the competition literature and the logical implications for the self-thinning process shows that all four assumptions are untenable as generalizations. Unfortunately, their application provides neither a mortality-inducing mechanism nor insight into the relationship between stand growth and mortality. • Explaining the observed variation in self-thinning relationships therefore requires improved representation of the competition process. This improvement is likely to require IBMs that explicitly represent variation in plant size or resource acquisition, two-dimensional stand distribution, dynamic rather than static stand resource utilization, and, perhaps, explicit three-dimensional stand development. Most importantly, the requirement for explicitly modelling mortality mechanisms implies that whole plant models may be insufficient for insight into the self-thinning process. • The review reinforces the need to assess mechanistic models for more than their ability to reproduce a single, high-level pattern. Such models should be assessed for their ability to simultaneously reproduce multiple features selected from the levels of both the modelled mechanisms and the high-level patterns. • Progress in understanding the observed variation in self-thinning currently requires a shift from searching for universal insight into the modelling of specific mechanisms for specific plant types, eventually leading to a broader theory explaining how variation in plants affects the competition process. Journal of Ecology (2005) doi: 10.1111/j.1365-2745.2005.00976.x
Article
Question: To what extent do tree growth, mortality, and long-term disturbance patterns affect stand structure and composition of an old-growth Picea abies forest?Location: Boreal Sweden.Methods: We linked data from three 50 m × 50 m permanent plots established in 1986 with dendrochronology data to evaluate tree growth and mortality over an 18-year period and to describe a several-hundred-year disturbance history for this forest type.Results: Averaged over all diameters, P. abies trees had an annual mortality rate of 0.60%; however, diameter had a striking effect on both growth and mortality, with trees of intermediate diameters (ca. 20–30 cm) showing faster growth and lower mortality. Their increased vigor gave rise to a diameter distribution resembling the ‘rotated sigmoid’(not reverse-J) proposed for such conditions, and it led to a deficit of snags of intermediate diameters. Slow-growing trees had an increased likelihood of dying. Although recruitment occurred in most decades over the past 400 years, two prominent recruitment peaks occurred (mid 1700s and 1800s), neither of which appeared to cause a shift in tree species composition. The lack of fire evidence suggests that fire was not responsible for these recruitment peaks.Conclusions: Taken together, these results depict a rather impassive system, where canopy trees die slowly over decades. Field observations suggest that fungal infections, mediated by wind, account for much of the mortality during these periods of relative quiescence. However, these periods are at times punctuated by moderate-severity disturbances that foster abundant recruitment.
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
We show that explicit mathematical and biological relationships exist among the scaling exponents and the allometric constants (α and β, respectively) of log - log linear tree-community size frequency distributions, plant density N T, and minimum, maximum and average stem diameters (D min, D max, and D̄, respectively). As individuals grow in size and D max increases, N T is predicted to decrease as reflected by a decrease in the numerical value of α and an increase in the value of β. Our derivations further show that N T decreases as D̄ increases even if D min or D max remain unchanged. Because D max and the age of the largest individuals in a community are correlated, albeit weakly, we argue that the interdependent relationships among the numerical values of α, β, N T, and D̄ shed light on the extent to which communities have experienced recent global disturbance. These predicted relationships receive strong statistical support using two large datasets spanning a broad spectrum of tree-dominated communities.