Storm legacies shaping post-windthrow forest regeneration: learnings from spatial indices in unmanaged Norway spruce stands

The anticipated increase in extreme disturbance events due to climate change is likely to expose Norway spruce (Picea abies (L.) Karst.) dominated forests in northern Europe to new conditions. Empirical data on the resilience of such natural (unmanaged) forests to disturbance and the long-term patterns of regeneration in its aftermath are currently scarce. We performed a quantitative assessment of natural forest stands in north–western Latvia to identify and characterise the patterns of stand structure 44 years after a stand-replacing disturbance and investigated the effects of legacies on regeneration. The spatial distribution of tree species and their dimensions were assessed in 71 circular sample plots (500 m² each) in natural forest areas, where Norway spruce dominated prior to the windthrow and salvage logging was not carried out. Spatial indices (species mingling index, size differentiation index, and aggregation index) were used to characterise stand structure and diversity. The different initial states (age and coverage of surviving trees) of stands affected eventual tree species dominance, size differentiation, degree of mingling and aggregation. Our results demonstrate a close relationship between disturbance legacies and spatial indices. The pre-storm understory and canopy survivors decreased species mingling, whereas survivors increased size differentiation. The size differentiation increased also with a higher degree of species mingling. Leaving differential post-storm legacies untouched promotes a higher structural and species diversity and therefore supports the management approach of preserving canopy survivors.

     

Mitigating fire risk to late-successional forest reserves on the east slope of the Washington Cascade Range, USA

A fire-risk model was developed using a stand-structure approach for the forests of the eastern slopes of the Washington Cascade Range, USA. The model was used to evaluate effects of seven landscape-scale silvicultural regimes on fire risk at two spatial scales: (1) the risk to the entire landscape; and (2) the risk to three reserve stands with stand structures associated with high conservation priorities (layered canopy, large trees, multiple species). A 1000 ha landscape was projected five decades for each management regime using an individual tree, distance-independent growth model. Results suggest that a variety of silvicultural approaches will reduce landscape fire risk; however, reserve stand fire risk is minimally decreased by thinning treatments to neighboring stands. Intensive fuel reduction through prescribed burning and selection of reserve stands in favorable topographic positions provide substantial fire risk reductions.     

Modeling silviculture after natural disturbance to sustain biodiversity in the longleaf pine (Pinus palustris) ecosystem: balancing complexity and implementation

Modeling silviculture after natural disturbance to maintain biodiversity is a popular concept, yet its application remains elusive. We discuss difficulties inherent to this idea, and suggest approaches to facilitate implementation, using longleaf pine (Pinus palustris) as an example. Natural disturbance regimes are spatially and temporally variable. Variability leads to a range of structural outcomes, or results in different pathways leading to similar structures. In longleaf pine, lightning, hurricanes, surface fires, and windthrow all lead to similar structures, but at different rates. Consequently, a manager can select among various natural disturbance patterns when searching for an appropriate silvicultural model. This facilitates management by providing flexibility to meet a range of objectives. The outcomes of natural disturbances are inherently different from those of silviculture, for example, harvesting always removes boles. It is instructive to think of silvicultural disturbances along a gradient in structural outcomes, reflecting degree of disparity with natural disturbance. In longleaf pine this might involve managing for two-cohort structure, instead of multi-cohort structure characteristic of old growth stands. While two-cohort structure is a simplification over the old growth condition, it is an improvement over single-cohort management. Reducing structural disparity between managed and unmanaged forests is key to sustaining biodiversity because of linkages that exist between structural elements, forest biota, and ecosystem processes. Finally, interactions of frequency, severity, intensity, seasonality, and spatial pattern define a disturbance regime. These components may not have equal weight in affecting biodiversity. Some are easier to emulate with silviculture than are others. For instance, ecologists consider growing-season fire more reflective of the natural fire regime in longleaf pine and critical for maintenance of biodiversity. However, dormant season fire is easier to use and recent work with native plants suggests that seasonality of fire may be less critical to maintenance of species richness, as one component of biodiversity, than is generally believed. Science can advance the goal of modeling silviculture after natural disturbances by better illustrating cause and effect relationships among components of disturbance regimes and the structure and function of ecosystems. Wide application requires approaches that are adaptable to different operational situations and landowner objectives. A key point for managers to remember is that strict adherence to a silvicultural regime that closely parallels a natural disturbance regime may not always be necessary to maintain biodiversity. We outline examples of silvicultural systems for longleaf pine that demonstrates these ideas.     

Three-dimensional light structure of an upland Quercus stand post-tornado disturbance

Light is the most common limiting factor in forest plant communities,influencing species composition,stand structure,and stand productivity in closed canopy stands.Stand vertical light structure is relatively simple under a closed canopy because most light is captured by overstory trees.However,wind disturbance events create canopy openings from local to landscape scales that increase understory light intensity and vertical light structural complexity.We studied the effects of an EF-1 tornado on horizontal and vertical(i.e.three-dimensional)light structure within a Quercus stand to determine how light structure changed with increasing disturbance severity.We used a two-tiered method to collect photosynthetic photon flux density at 4.67 m and 1.37 m above the forest floor to construct three-dimensional light structure across a canopy disturbance severity gradient to see if light intensity varied with increasing tornado damage.Results indicate that increased canopy disturbance closer to the tornado track increased light penetration and light structure heterogeneity at lower forest strata.Increased light intensity correlated with increased sapling density that was more randomly distributed across the plot and had shifted light capture higher in the stand structure.Light penetration through the overstory was most strongly correlated with decreased stem density in the two most important tree species(based on relative dominance and relative density)in the stand,Quercus alba L.(r=-0.31)and Ostrya virginiana(Mill.)K.Koch(r=-0.27,p<.01),and indicated that understory light penetration was most affected by these two species.As managers are increasingly interested in patterning silvicultural entries on natural disturbances,they must understand residual stand and light structures that occur after natural disturbance events.By providing spatial light data that quantifies light structure post-disturbance,managers can use these results to improve planning required for long-term management.The study also provides comparisons with anthropogenic disturbances to the midstory that may offer useful comparisons to natural analogs for future silvicultural consideration.     

A silvicultural management tool for the Oriental beech (Fagus orientalis Lipsky) forests

Stand density management tools help forest managers and landowners to more effectively allocate growing space so that specific silvicultural objectives can be met. Due to the economic importance of Oriental beech (Fagus orientalis Lipsky) forests in Turkey, a stand density management tool was developed for this species to optimize regeneration success rate and tree growth. For the development of this tool, named stand density management diagram (SDMD), we utilized forest inventory data from the Kastamonu Regional Forest Directorate in Turkey. Previously published forest management approaches and models were employed during the development of the tool. The SDMD illustrates the relation among four forest indexes: the basal area per hectare, number of trees per hectare, forest stand volume per hectare, and quadratic mean diameter of the beech stands. The stand stocking percent (SSP) can be determined based upon any two of these four measurements. The results suggest that SSP is a better predictor of tree growth than BA in Oriental beech forests. The newly developed SDMD allows for a more effective use of the growing space to achieve specific silvicultural objectives including tree regeneration, timber production, thinning planning, and wildlife protection in Oriental beech forests.     

Fine-scale analysis reveals a potential influence of forest management on the spatial genetic structure of Eremanthus erythropappus

Forest management may have significant effects on forest connectivity and natural population sizes.Harvesting old-growth single trees may also change natural patterns of genetic variation and spatial genetic structure.This study evaluated the impacts of forest management using a silvicultural system of seed trees on the genetic diversity and spatial genetic structure of Eremanthus erythropappus(DC.)MacLeish.A complete survey of 275 trees on four plots was undertaken out to compare the genetic variation of a managed stand with an unmanaged stand.We genotyped all adult and juvenile individuals 60 months after the management and compared the genetic diversity and the spatial genetic structure parameters.Genetic diversity was considered high because of an efficient gene flow between stands.There were no genetic differences between stands and no evidence of inbreeding.Genetic clustering identified a single population(K=1),indicating no genetic differentiation between managed and unmanaged stands.Adult and juvenile individuals of the unmanaged stand were more geographically structured than individuals from the managed one.There was a tendency of coancestry among juveniles at the first class of distance of the managed stand,suggesting a drift of genetic structure possibly caused by management.Understanding early responses to management on genetic diversity and stand structure is a first step to ensuring the effectiveness of conservation practices of tree species.The sustainability of forest management of E.erythropappus on genetic diversity,and more accurately,on spatial genetic structure needs evaluation over time to promote effective conservation of the population size and genetic variability.     

Height-diameter models with random coefficients and site variables for tree species of Central Maine

Height-diameter models were developed for nine tree species common to the northeastern United States: Abies balsamea, Acer rubrum, Betula papyrifera, B. populifolia, Picea rubens, P. mariana, Pinus strobus, Populus tremuloides, and Tsuga canadensis. Stem heights and diameters were collected from 6 146 trees (between 136 and 2615 trees per species) on 50 plots within 10 structurally diverse stands that are part of a long-term silvicultural experiment in central Maine. The models were developed using both generalized nonlinear least squares (GNLS) and multi-level, mixed-effects approaches. Mixed-effects approaches were superior to GNLS, with inclusion of site covariates (tree density and basal area) accounting for some of the variability explained by the random coefficients in the full mixed-effect models. Analysis of plot-level parameter estimates suggested that differences in stand structure (even-aged vs. uneven-aged silvicultural practices) had a significant influence on the height-diameter relationships.     

Successional process of Picea crassifolia forest after logging disturbance in semiarid mountains: A case study in the Qilian Mountains,northwestern China

Selective logging is the most widely employed method of commercial timber production in Asia, and its impact on forest structure, composition, and regeneration dynamics is considerable. However, the successional processes in forest communities after logging in semiarid mountains are poorly understood. To provide more information on these processes, we used data from tree rings, direct and indirect age determinations, and field measurements of stand structure to reconstruct the historical disturbance regime, stand development patterns, and successional processes in a natural Picea crassifolia forest community in the Qilian Mountains of northwestern China. The results showed that the density of P. crassifolia forest increased significantly after logging. The densities of second growth forests 30 and 70 years after logging disturbance had increased to 2874% and 294% of primary forest's density, respectively. Logging disturbance did not alter tree species composition of logged stands. However, the diversity of understory species changed significantly among the successional phases. Logging disturbance decreased the spatial heterogeneity of second growth forest. The spatial distributions of recruitment were affected by the location of the remaining trees. There was less recruitment near the remaining trees than near forest that had been cut. In addition, logging disturbance also induced a growth release for the trees on the sites sampled. Our results imply that the succession and regeneration of P. crassifolia forest may be improved if the remaining trees could be retained relative uniform distribution pattern, thinning or selective logging could be performed to height density, exotic shrubs could be removed or the shrubs cover could be reduced during the earlier successional stages.     

Managing forest complexity through application of disturbance–recovery knowledge in development of silvicultural systems and ecological rehabilitation in natural forest systems in Africa

How can we accommodate the diversity in tree species and sizes in mixed-species/size/age stands in the sustainable management of natural forests and woodlands in Africa for diverse timber and non-timber forest products and services, and during rehabilitation of degraded forests? The evergreen moist tropical to warm-temperate forests, from the equator to 34°S, generally function through relative shade tolerance. The tropical, strongly seasonal, drier deciduous woodlands generally function through adaptation to fire and/or grazing/browsing. Silvicultural systems, when implemented, are often not aligned with the ecological characteristics of the particular forest systems or the specific targeted species. This paper presents the concept of using the basic disturbance–recovery processes, with recovery development via stand development stages, as the basis for the development of silvicultural systems suitable for maintenance of forest complexity. Grain analysis and stem diameter distributions, analysed from resource inventories, are used to determine the specific shade- or fire-tolerance characteristics of key economical and ecological tree species. The gained knowledge of the ecosystem and species characteristics (including modes of regeneration, i.e. from seed or vegetative regrowth) and processes is used to simulate the ecological disturbance–recovery processes through the development of mixed silvicultural systems, such as a single-tree selection system, a group felling system and a coppice management system in the same forest. Very similar concepts are used to develop rehabilitation strategies to recover the processes towards regrowth stands of diverse species and structure: through stands of introduced plantations and invader tree species; and through early regrowth stages in deciduous woodlands and evergreen forests, after fire, slash-and-burn traditional agriculture, charcoal production or open-cast mining.     

On the structural and species diversity effects of bark beetle disturbance in forests during initial and advanced early-seral stages at different scales

Following disturbances, early-seral stages of forests provide a variety of structures. Whether this variety is a short-term phenomenon or influences forest succession for several decades or even longer is not known. We tested the hypotheses that after spruce dieback caused by bark beetles, a high spatial heterogeneity of stand structures will persist within stands and among stands even in advanced early-seral stages and that species taxonomical and functional diversity measures will reflect this heterogeneity. We used a chronosequence of unmanaged forests in the Berchtesgaden National Park (Germany) consisting of mature undisturbed spruce stands (control), stands belonging to an initial early-seral stage (~3 years after disturbance) and stands in an advanced early-seral stage (~20 years after disturbance). We analysed diversity and heterogeneity of these forest stands including stand structure, species density, species composition and functional–phylogenetic diversity of vascular plants, wood-inhabiting fungi and saproxylic beetles within plots, among plots of the same successional stage and among stages. Stands of the advanced early-seral stage were characterized by a high spatial heterogeneity of structural attributes, such as crown cover, regeneration density and spatial distribution of trees. Among-plot taxonomic beta diversity was highest in the advanced early-seral stage for beetles, but lowest for fungi, while beta diversity of plants among plots remained unchanged during succession. The mosaic of successional stages initiated by bark beetles increased the gamma diversity of the study area, especially for fungi and beetles. Our findings support the hypothesis that structural heterogeneity continues for at least two decades at stand and landscape scales and that species turnover among successional stages is a major mechanism for gamma diversity in forests after bark beetle disturbance.     

Influence of fire intensity on structure and composition of jack pine stands in the boreal forest of Quebec: Live trees,understory vegetation and dead wood dynamics

North American jack pine (Pinus banksiana Lamb.) stands are generally characterized by an even-aged structure resulting from high intensity fires (HIF). However, non-lethal fires of moderate intensity (MIF), which leave behind surviving trees, have also been reported. The objectives of this study were two-fold: (1) assess the concurrent dynamics of live trees, understory vegetation and different types of coarse woody debris (CWD) during succession after HIF; and (2) document how MIF affects stand structure component dynamics compared to HIF. Stands affected by both HIF and MIF were selected. Tree characteristics and age structure, understory biomass, and CWD volume were assessed. Our results suggest that the structural succession of jack pine stands following HIF comprises three stages: young stands (<48 years), premature and mature stands (58–100 years) and old stands (>118 years). Canopy openness and jack pine density significantly decreased with time since HIF, while black spruce density and CWD volume significantly increased. The highest structural diversity was measured in the premature and mature stands. Compared to HIF, MIF increased mean jack pine basal area, decreased average stand density, delayed the replacement of jack pine by black spruce replacement in the canopy, decreased CWD volume, and significantly increased bryophytes mass. MIF increased the diversity of live trees and generally decreased CWD structural diversity. The study confirms the diversity of natural disturbance magnitude and successional processes thereby initiated. Thereafter, it appeared to be relevant for adjustment of disturbance emulating forest-management systems.     

Singular and interactive effects of blowdown, salvage logging, and wildfire in sub-boreal pine systems

The role of disturbance in structuring vegetation is widely recognized; however, we are only beginning to understand the effects of multiple interacting disturbances on ecosystem recovery and development. Of particular interest is the impact of post-disturbance management interventions, particularly in light of the global controversy surrounding the effects of salvage logging on forest ecosystem recovery. Studies of salvage logging impacts have focused on the effects of post-disturbance salvage logging within the context of a single natural disturbance event. There have been no formal evaluations of how these effects may differ when followed in short sequence by a second, high severity natural disturbance. To evaluate the impact of this management practice within the context of multiple disturbances, we examined the structural and woody plant community responses of sub-boreal Pinus banksiana systems to a rapid sequence of disturbances. Specifically, we compared responses to Blowdown (B), Fire (F), Blowdown-Fire, and Blowdown-Salvage-Fire (BSF) and compared these to undisturbed control (C) stands. Comparisons between BF and BSF indicated that the primary effect of salvage logging was a decrease in the abundance of structural legacies, such as downed woody debris and snags. Both of these compound disturbance sequences (BF and BSF), resulted in similar woody plant communities, largely dominated by Populus tremuloides; however, there was greater homogeneity in community composition in salvage logged areas. Areas experiencing solely fire (F stands) were dominated by P. banksiana regeneration, and blowdown areas (B stands) were largely characterized by regeneration from shade tolerant conifer species. Our results suggest that salvage logging impacts on woody plant communities are diminished when followed by a second high severity disturbance; however, impacts on structural legacies persist. Provisions for the retention of snags, downed logs, and surviving trees as part of salvage logging operations will minimize these structural impacts and may allow for greater ecosystem recovery following these disturbance combinations.     

Spatial structure of deciduous forest stands with contrasting human influence in northwest Spain

Five contrasting deciduous forest stands were studied to characterize the spatial structural variability in human-influenced forests. These stands are representative of cultural forest types widely represented in western Europe: one plantation, two coppices, one wood-pasture forest and one high forest stand. All stems with DBH > 5 cm were measured and mapped, and stem DBH distributions, spatial structure of DBH, spatial point patterns and spatial associations were analysed. Spatial autocorrelation for DBH was calculated with Moran’s I correlograms and semivariograms. Complete spatial randomness hypothesis for spatial point patterns, and both independence and random labelling hypotheses for spatial associations were analysed using Ripley’s K function. The results showed that tree sizes were conditioned by particular former management systems, which determined unimodal symmetric, positively skewed or compound DBH distributions. Spatial structure was more complex when human influence became reduced. Coppice stands showed clumped spatial patterns and independence among size classes, as a consequence of sexual and vegetative establishment of new stems in open areas. The largest clumping intensity was observed in the wood-pasture with an intermediate disturbance frequency and low inter-tree competition. The high forest stand displayed spatial traits consistent with the gap-dynamics paradigm, such as clumping of smaller trees, random arrangement of larger trees, negative association between juveniles and adults, and high structural heterogeneity. It can be expected that after cessation of human interference, coppices and wood-pastures would evolve to a more heterogeneous structure, probably with a higher habitat and species diversity.     

Structure of a Central-European mountain spruce old-growth forest with respect to historical development

This study examines the structural characteristics of the tree layer, dead wood, canopy openings, and regeneration patterns of a spruce old-growth forest in the Bohemian Forest, Czech Republic. An old-growth stand with minor human influence and a stand that was presumably logged about 200 years ago were analyzed and compared, as some forest managers considered the presumable human impact as a reason for salvage logging. Even though the stands differed in tree density, height and DBH structure, it was not possible to conclude whether it was due to management history or the environmental differences. The volume of dead wood also differed between the stands. There was about 142 and 83 m³ ha⁻¹ of dead wood in the old-growth stand and presumably logged stand, respectively. The amount of dead wood found in the old-growth stand was comparable with values reported from spruce old-growth stands across Central Europe. In both stands, many canopy trees were arranged in linear patterns, which was a result of spruce regeneration on nurse logs. This suggests that the origin and development of the stands were characterized by natural processes and during the past 200 years typical old-growth structural characteristics have already evolved.     

Silviculture for old-growth attributes

Silviculture to maintain old-growth forest attributes appears to be an oxymoron since the late developmental phases of forest dynamics, described by the term old-growth, represent forests that have not experienced human intervention or timber removal for a long time. In the past, silvicultural systems applied to old-growth aimed to convert it into simplified, more productive regrowth forests substantially different in structure and composition. Now it is recognised that the maintenance of biodiversity associated with structural and functional complexity of late forest development successional stages cannot rely solely on old-growth forests in reserves. Therefore, in managed forests, silvicultural systems able to develop or maintain old-growth forest attributes are being sought. The degree to which old-growth attributes are maintained or developed is called “old-growthness”. In this paper, we discuss silvicultural approaches that promote or maintain structural attributes of old-growth forests at the forest stand level in (a) current old-growth forests managed for timber production to retain structural elements, (b) current old-growth forests requiring regular, minor disturbances to maintain their structure, and (c) regrowth and secondary forests to restore old-growth structural attributes. While the functions of different elements of forest structure, such as coarse woody debris, large veteran trees, etc., have been described in principle, our knowledge about the quantity and distribution, in time and space, of these elements required to meet certain management objectives is rather limited for most ecosystems. The risks and operational constraints associated with managing for structural attributes create further complexity, which cannot be addressed adequately through the use of traditional silvicultural approaches. Silvicultural systems used in the retention and restoration of old-growthness can, and need, to employ a variety of approaches for managing spatial and temporal structural complexity. We present examples of silvicultural options that have been applied in creative experiments and forestry practice over the last two decades. However, these largely comprise only short-term responses, which are often accompanied by increased risks and disturbance. Much research and monitoring is required still to develop and optimise new silvicultural systems for old-growthness for a wide variety of forest ecosystem types.     

林木生长和收获预估模型的研究动态

从近年来林木生长和收获模型的研究中,可以看出有如下的发展动向:（1）由林分水平模型向径级水平模型、单木水平模型转变;（2）从林木的自然生长模型发展来考虑人为育林措施的经营模型;（3）由简单的人工同龄纯林模型深化到复杂的天然异龄混交林;（4）关注点从测树学调查因子的经验性模型转向生理生态的机理性模型。还就今后我国的林木生长模型研究提出了几点建议。     

Topographic and compositional influences on disturbance patterns in a northern Maine old-growth landscape

Disturbance patterns are strongly coupled with forest composition and structure, and patterns change through time in response to shifts in climate, anthropogenic impacts and other factors. Knowledge of the natural disturbance patterns for establishing baseline conditions for a forest type or ecosystem facilitates change detection for other elements of the biophysical system important to management and conservation. Dendrochronological reconstructions from old-growth forest remnants throughout northeastern North America document average decadal rates of disturbance of 5%–<10% over the last 150–300 years. Relatively frequent, low severity disturbance characterized by small gaps representing canopy openings made by 1–3 trees prevail in these forests dominated by varying mixtures of late-successional tree species. Few studies, though, have explicitly characterized differences related to composition or topographic setting in old-growth landscapes. We addressed this by comparing the temporal and spatial disturbance patterns reconstructed from tree rings at two spatial resolutions (0.5 ha and 200 m²). Sites were selected to assess the influence of topography (slope) and cover type in stands where red spruce (Picea rubens Sarg) and balsam fir (Abies balsamea L. Mill) were key components. Low rates of disturbance (average <10% per decade) and small gap sizes (≤30 m²) prevailed in all stands during the decades from 1850–1980. Episodic pulses of disturbance, of nearly moderate intensity in some stands, opened ca. 20–30% of the canopy area and were associated with wind events and/or insect outbreaks that differentially affected stands. We found no significant difference in the average temporal rates of disturbance related to cover type or topography in 0.5-ha plots. However, the influence of these factors was evident in comparisons of gap areas estimated for 200-m² plot sections. At this resolution, the largest canopy openings (≥100 m²) occurred most frequently in slope sites, enabling pulses of canopy accession for Betula alleghaniensis (Britton). Whereas the smallest canopy openings (≤30 m²) dominated softwood stands, favoring red spruce, balsam fir and other shade-tolerant species throughout the forest. The variable effects of common disturbance agents, regardless of topographic position and/or cover types, points to the important role of biological legacies determining stand structure and composition on subsequent disturbance events and long-term patterns.