Natural stand structures, disturbance regimes and successional dynamics in the Eurasian boreal forests: a review with special reference to Russian studies

? This review summarizes early stand-scale studies of pristine forest structures, disturbance regimes and successional patterns carried out in boreal Eurasia. We attempt to reveal, characterize and classify stand dynamic types that can be used as templates for nature-based forest management.

? The studies reviewed demonstrate multiple successional pathways in stand development in all types of pristine forests. All-aged stands driven by small-scale disturbances are formed over successional development of several hundreds of years. This endogenous development can be interrupted by stand-replacing or partial disturbances leading to successions with even-aged or cohort-structured stands, respectively. In Western Europe, the most common disturbances are windthrows, surface fires and fluctuations in moisture regime; in Eastern Europe and Siberia, the most common disturbances are crown and surface fires and insect outbreaks. Type, return interval and severity of disturbances are strongly influenced by the site conditions and successional stage of a stand.

? Based on characteristics of forest stands and disturbance regime, four main types of pristine boreal forest stand dynamics can be distinguished: (1) even-aged, compositional change dynamics, (2) even-aged, mono-dominant dynamics, (3) cohort dynamics and (4) fine-scale gap dynamics. These types can be mimicked in developing scenarios of ecological sustainable forest management in Eurasian boreal forests.

     

The role of gaps and tree regeneration in the transition from dense to open black spruce stands

Black spruce forests growing on clay soils in northwestern Quebec change structure from dense even-aged stands to open uneven-aged stands such that almost all forests older than 200 years have an open canopy. These forests become unproductive over time because they are prone to paludification. The main goal of our study was to document the transition between dense and open stands in terms of gap dynamics, with a focus on tree regeneration. Our objective was to determine whether forests remain open due to a lack of regeneration, a lack of growth or both. Nine stands along a 50–250-year-old time since fire gradient were sampled with the line intersect sampling method. Gap fraction increased with stand age and reached a maximum of 77% in the oldest site. In old-growth stands, gaps were interconnected due to the low density of these forests. Most of the gap makers were found with broken stems. Regeneration was dominated by black spruce layers and was relatively abundant (1.71 stems/m²). However, the majority of gap fillers were smaller than 1 m in height in stands of all ages. Instead of a lack of regeneration, the opening of the forests is due to a lack of growth associated with cold and wet organic deposits. Partial harvesting could be implemented on the most productive sites, while management techniques including soil disturbances will be required on low productivity sites to recreate good growth conditions.     

Long-term influence of fire and harvesting on boreal forest age structure and forest composition in eastern Québec

In boreal forests, historical variations in the area disturbed by natural disturbances or harvesting have rarely been compared. We measured temporal and spatial variations in areas affected by severe fires and clearcutting throughout the 20th century in a 57, 332 km² section of the eastern Canadian boreal forest. We examined the effects of these disturbances on spatio-temporal variations in the abundance of forests >60 years. Natural variability for the abundance of forests >60 years was estimated from simulations of natural disturbance regimes. We also measured compositional and structural differences between three categories of stands originating from relatively recent disturbances (∼50 years; clearcutting, fires, and clearcutting followed by fires), and one category of stands that were undisturbed for at least 200 years. At the regional level, we observed that forests >60 years gradually became scarcer throughout the 20th century due to a gradual expansion of harvested areas, an effect most pronounced in the southern part of the region, where mature and old forest abundance was clearly outside the range of natural variability at the end of the studied period. At the stand level, forest composition and structure differed between stand-origin categories: clearcutting-origin stands contained more balsam fir (Abies balsamea), fire-origin stands more black spruce (Picea mariana), and fire/clearcutting-origin stands more hardwoods (Betula papyrifera and Populus tremuloides). Overall, we estimate that strict forest management targets based on natural disturbance regimes will be difficult to achieve in eastern North-American boreal forests, most notably because contemporary disturbance rates, including both clearcutting and fire, have gradually become higher than the fire rates observed during the preindustrial period.     

A density management diagram for Norway spruce in the temperate European montane region

Norway spruce is one of the most important conifer tree species in Europe, paramount for timber provision, habitat, recreation, and protection of mountain roads and settlements from natural hazards. Although natural Norway spruce forests exhibit diverse structures, even-aged stands can arise after disturbance or as the result of common silvicultural practice, including off-site afforestation. Many even-aged Norway spruce forests face issues such as senescence, insufficient regeneration, mechanical stability, sensitivity to biotic disturbances, and restoration. We propose the use of Density Management Diagrams (DMD), stand-scale graphical models designed to project growth and yield of even-aged forests, as a heuristic tool for assessing the structure and development of even-aged Norway spruce stands. DMDs are predicated on basic tree allometry and the assumption that self-thinning occurs predictably in forest stands. We designed a DMD for Norway spruce in temperate Europe based on wide-ranging forest inventory data. Quantitative relationships between tree- and stand-level variables that describe resistance to selected natural disturbances were superimposed on the DMD. These susceptibility zones were used to demonstrate assessment and possible management actions related to, for example, windfirmness and effectiveness of the protective function against rockfall or avalanches. The Norway spruce DMD provides forest managers and silviculturists a simple, easy-to-use, tool for evaluating stand dynamics and scheduling needed density management actions.     

Epiphytic lichen diversity in old-growth and managed Picea abies stands in Alpine spruce forests

In the last decades, a large body of literature has grown to evaluate the impact of forest management on epiphytic lichens in boreal coniferous forests. However, information is still lacking on coniferous forests of the Alps. This study compares lichen diversity between spruce forest stands of four successional stages: (1) young, (2) intermediate, (3) mature forests managed for timber production with a rotation cycle of 120–180 years, and (4) old-growth protected forests. The emphasis was placed on the occurrence of nationally rare and calicioid species (lichens and fungi traditionally referred to as Caliciales, known to be indicative of forest age and continuity). For each forest successional stage, four plots were selected. In each plot, 7 spruce individuals were surveyed for epiphytic lichens according to a standardised sampling method. Species richness increased from young to mature stands, while no difference was detected between mature and old-growth stands. This pattern was also confirmed for rare and calicioid species which are, however, more frequent in old-growth stands. Differences in species composition were also found between the different forest successional stages. Mature and old-growth plots slightly overlap, indicating that to some extent comparable lichen assemblages could be found in these stands. A nested pattern of species assemblages was found, old-growth stands hosting most of the species which were also found in stands belonging to the previous forest successional stages. Our results support the hypothesis that the management regime applied to spruce forests of the Italian Alps renders mature stands managed for timber production somewhat similar to old-growth stands as lichen habitat. However, we found a higher complexity in old-growth forests, and many species of conservation concern clearly preferred old-growth stands. In this perspective, a further prolongation of the normal cycle it is likely to be a most favourable conservation-oriented management to be recommended at least within protected areas and Natura 2000 sites, where conservation purposes should receive a high priority.     

Influence of post-harvest silviculture on understory vegetation: Implications for forage in a multi-ungulate system

Natural disturbance emulation has emerged as a key management approach to maintaining biodiversity in logged boreal forests. Forest managers’ success in emulating understory forest ecosystem functions, e.g., for the provision of habitat even for large mammals, has not been tested due, in many cases, to incomplete records of silviculture. We examined regenerating areas of previously conifer-dominated forests in northwestern Ontario, Canada, 10 and 30 years after logging and 10 and 30 years after fire to test if understory development and moose (Alces alces) forage abundance differed between the two disturbance types and artificial or natural regeneration approaches. In addition, we counted moose pellet groups as a measure of moose use of the region. Specific treatments included: (1) naturally regenerating, fire-origin forests, (2) post-harvest, regenerating forests with natural establishment of trees, and (3) post-harvest, regenerating forests with mechanical or chemical site preparation and planting and/or herbicide spraying. We hypothesized that the understory in post-harvest stands would support higher forage availability for moose compared to similarly-aged, fire-origin stands. Abundance of hardwoods, shrubs, and herbaceous plants was greater in naturally-regenerated post-harvest stands than in fire-origin and artificially regenerated post-harvest stands at both 10 and 30 years post-disturbance. However, post-harvest, naturally regenerating stands were not significantly associated with higher moose use, rather evidence of moose use increased as a function of the amount of naturally regenerating logged forest in the surrounding landscape. This study suggests that, relative to fire, the intensity of post-harvest silviculture influences habitat suitability for moose. The effect likely cascades to other ungulates, such as woodland caribou (Rangifer tarandus caribou), and vegetation management needs to be considered at scales greater than the stand level in order to achieve habitat management for large mammals.     

A simulation of the development and restoration of old-growth structural features in northern hardwoods

Thinning treatments in second-growth forest may be a practical means of accelerating the development of certain old-growth structural features in regions where old stands are presently uncommon. We used CANOPY, an individual-tree model calibrated with data from thinned and unthinned stands, to simulate effects of thinning on growth rates and development of old-growth structural features in second-growth northern hardwoods. Three simulated, moderately heavy thinnings over a period of 45 years nearly doubled the predicted mean radial increment of canopy trees, percent of stand basal area in large trees, and area of canopy gaps. Compared to untreated stands, thinned stands had fewer dead trees per ha, but the dead trees were larger in size and the overall volume of snags and logs was little affected. In a 77-year old even-aged stand, moderately heavy thinning was predicted to reduce the time needed to attain the minimum structural features of an old-growth forest from 79 to 36 years. Simulated treatments in an older, uneven-aged stand gave mixed results; the moderately heavy treatment stimulated individual tree growth, but the removal of some medium-sized canopy trees in conjunction with natural mortality delayed the development of old-growth structure. Total volume of dead wood may still be deficient under the thinning regimes investigated in this study, but predicted live-tree structure 45 years after moderately heavy thinning was typical of stands in the advanced transition and steady-state stages of old-growth development. Results suggest that thinning can substantially accelerate the development of old-growth structure in pole and mature northern hardwoods, but response in older, uneven-aged stands is more modest, and treatments in these stands may need to be more conservative to achieve restoration goals.     

Short term effects of alternative silvicultural treatments on stand attributes in irregular balsam fir-black spruce stands

The eastern Canadian boreal forest exhibits a specific disturbance regime where forest fires are less frequent than in the western part. This particularity may explain the abundance of irregular stands with distinct ecological features. To ensure sustainable forest management, these characteristics require the implementation of an adapted silviculture regime. In this context, two selection cutting methods were developed and compared with more conventional techniques, initially designed for cutting more regular stands of the boreal forest (cutting leaving small merchantable stems, careful logging preserving advance regeneration). The comparison focused on the capacity of treatments to maintain the primary attributes of irregular boreal forests, including complex vertical structure, abundant tree cover, species composition, and an abundance of dead wood. Mortality and regeneration processes were also compared.     

Presettlement and modern disturbance regimes in coast redwood forests: Implications for the conservation of old-growth stands

Coast redwood (Sequoia sempervirens), a western North American conifer of ancient lineage, has a paradoxical combination of late-successional characteristics and strong adaptations to disturbance. Despite its shade tolerance and heavy dominance of the canopy on many sites, redwood saplings are uncommon in upland old-growth stands. Information needed to ensure the conservation of old-growth redwood forests has been limited. In this review paper, we integrate evidence on redwood biology with data on the historic and modern disturbance regimes to help clarify the degree to which key attributes of redwood forests may have been dependent upon periodic disturbance. Available evidence suggests that episodes of fire, flooding, and slope failure prior to European settlement were frequent but predominantly of low to moderate severity and extent, resulting in broadly uneven-aged forests. The majority of fires prior to European settlement were apparently of human origin. Frequency and severity of the major disturbance agents have been radically changed in modern times. Fires have been largely excluded, and flooding has been altered in ways that have often been detrimental to old-growth redwoods on alluvial terraces. However, because of the apparent anthropogenic origin of most presettlement fires, the long-term evolutionary role of fire for coast redwood is ecologically ambiguous. With fire exclusion, redwood possibly could be displaced to some extent on upland sites by increasing abundance of fire-sensitive competitors. Alternatively, redwood may be able to maintain dominance by vegetative sprouting and new seedling establishment on root-wad mounds, fallen logs, and on soil exposed by slope failure. Future research priorities are suggested that will help resolve some of the current ambiguities.     

Structural and environmental characterization of old-growth temperate rainforests of northern Chiloé Island,Chile: Regional and global relevance

Old-growth forests are ecologically relevant reservoirs of biodiversity and provide valuable and unique ecosystem functions in the landscape. However, what constitutes an old-growth stand is confusing because the definition depends largely on the forest type under study. Despite the ecological importance of old-growth temperate rainforests in southern Chile in comparison to other global forests, no attempts have been made to characterize them as a way to assess their structural variability. Here, we characterized old-growth stands of Valdivian and North Patagonian rain forest types located in Chiloé Island (Chile, 42°30′S) using inventory data from 23 permanent plots (0.1 ha each) located in rural landscapes and protected areas of northern Chiloé Island. For each stand, its age (average age of the oldest trees present in each stand) and disturbance regimes (evidence of recent human impact, e.g. cuttings or fires, and tree growth rates) were used as defining old-growth criteria. We characterized the structure (tree species richness, size-density distributions, vertical stratification and presence of snags) and floristic composition of each stand. Environmental variables (i.e. temperature, distance to coastline and elevation) were related to stand structure using multivariate constrained correspondence analysis. Old-growth forests were commonly characterized by (a) tree basal areas >80 m²/ha; (b) density of shade-tolerant tree species in the emergent and dominant canopy layer >36%; (c) higher tree species richness (>7 tree species) than successional stands; (d) presence of large canopy emergents (>80 cm dbh, >25 m tall); (e) high vertical heterogeneity; and (f) minimum stand ages older than 200 years. Old-growth forests showed a distinctive structural variability and floristic diversity influenced both by stand age and disturbance history of the stands. Structural variability was also related to environmental differences among sites (e.g. air temperature, distance to coastline, soil types). Old-growth forest features described here can offer a baseline for managers interested in maintaining and restoring old-growth forest structure in southern temperate rain forests.     

Forest management conditioning ground ant community structure and composition in temperate conifer forests in the Pyrenees Mountains

The search for indicators to monitor management impact on biodiversity is a crucial question because management practices promote changes in community structure and composition of different animal groups. This study explores the effect of widely conducted management practices (forest logging and livestock) in Pinus uncinata forests in the Pyrenees range (NE Spain) on the structure and composition of ground ant communities compared to those of old-growth stands. Forest structure clearly differed in stands with different forest managements. These stands managed for different uses also showed marked differences in structure and composition of ground ant communities. There was a great dominance of a single species, Formica lugubris, which accounted for 99% of ants collected in pitfall traps. Rarefaction curves indicated that species richness was highest in old-growth stands and lowest in even-aged ones, with woodland pasture stands showing an intermediate value. Classification methods allowed us to identify two groups of species: six species related to old-growth plots and three species (including F. lugubris) associated to managed stands. Habitat structure played an important role in determining the structure of ant communities: forests with high tree density but low basal area were the most favourable forest type for F. lugubris, while the abundance of the remaining ant species was negatively affected by the abundance of F. lugubris and by tree cover.     

Age and growth of a fire prone Tasmanian temperate old-growth forest stand dominated by Eucalyptus regnans, the world's tallest angiosperm

Forests are key components of the global carbon cycle, with deforestation being an important driver of increased atmospheric carbon dioxide. Temperate old-growth forests have some of the highest above ground stores of carbon of any forest types on Earth. Unlike tropical forests, the ecology of many temperate forests is dominated by episodic disturbance, such as high intensity fire. An exemplar of a particularly carbon dense temperate forest system adapted to infrequent catastrophic fires is the Eucalyptus regnans forests of south eastern Australia. Knowledge of the growth and longevity of old-growth trees is crucial to understanding the carbon balance and fire regimes of these forest systems. In an old-growth E. regnans stand in the Styx Valley in southern Tasmania we used dendrochronological techniques and radiocarbon dating to determine the age and stem growth of E. regnans and Phyllocladus aspleniifolius, an understorey rainforest conifer. Our analysis revealed that an even-aged cohort of E. regnans and P. aspleniifolius established in 1490–1510AD, apparently after a stand-replacing fire. The stem growth rates of E. regnans in the first 100 years were very rapid compared to the co-occurring P. aspleniifolius. That the longevity of E. regnans is >500 years challenges the suggested 350–450 year timeframe proposed for the widely held model of succession from eucalypt to rainforest. These forests not only have the potential to store vast amounts of carbon, but can also maintain these high carbon densities for a long period of time. Estimates of the capacity of these forests to sequester and store carbon should explicitly consider past harvesting and fire regimes and the potential increases in the risk of fire associated with climate change.     

Stand-landscape integration in natural disturbance-based management of the southern boreal forest

Forest ecosystem management, based partly on a greater understanding of natural disturbance regimes, has many variations but is generally considered the most promising approach to accommodating biodiversity concerns in managed forested regions. Using the Lake Duparquet Forest in the southeastern Canadian boreal forest as an example, we demonstrate an approach that attempts to integrate forest and stand-level scales in biodiversity maintenance. The concept of cohorts is used to integrate stand age, composition and structure into broad successional or stand development phases. Mean forest age (MFA), because it partly incorporates historic variability of the regional fire cycle, is used as a target fire cycle. At the landscape level, forest composition and cohort objectives are derived from regional natural disturbance history, ecosystem classification, stand dynamics and a negative exponential age distribution based on a 140 year fire cycle. The resulting multi-cohort structure provides a framework for maintaining the landscape in a semi-natural age structure and composition. At the stand level, the approach relies on diversifying interventions, using both even-aged and uneven-aged silviculture to reflect natural stand dynamics, control the passage (“fluxes”) between forest types of different cohorts and maintain forest-level objectives. Partial and selective harvesting is intended to create the structural and compositional characteristics of mid- to late-successional forest types and, as such, offers an alternative to increasing rotation lengths to maintain ecosystem diversity associated with over-mature and old-growth forests. The approach does not however supplant the necessity for complementary strategies for maintaining biodiversity such as the creation of reserves to protect rare, old or simply natural ecosystems. The emphasis on maintaining the cohort structure and forest type diversity contrasts significantly with current even-aged management in the Canadian boreal forest and has implications for stand-level interventions, notably in necessitating a greater diversification of silvicultural practices including more uneven-aged harvesting regimes. The approach also presents a number of operational challenges and potentially higher risks associated with multiply stand entries, partial cutting and longer intervals between final harvests. There is a need for translating the conceptual model into a more quantitative silvicultural framework. Silvicultural trials have been established to evaluate stand-level responses to treatments and operational aspects of the approach.     

Timber management with variable retention in Nothofagus pumilio forests of Southern Patagonia

Forestry practices integrating ecological and social criteria have been replacing those based only on economic values. Traditional silviculture, such as shelterwood cuts (SC), transforms uneven-aged original stands to an even-aged managed forest. Recently, other methods have proposed to conserve some of the original heterogeneity of the old-growth forests. One proposal leaves 30% of the timber quality forest area as aggregated retention and 20% basal area as dispersed retention. The aim of this study was to analyze the feasibility of timber management with aggregated and dispersed retention in Nothofagus pumilio old-growth forests by analyzing timber and harvesting yield potential compared with traditional regeneration systems. Also, remnant tree stability of aggregated retention was analyzed.     

Influence of climate change,fire and harvest on the carbon dynamics of black spruce in Central Canada

The results of EFIMOD simulations for black spruce (Picea mariana [Miller]) forests in Central Canada show that climate warming, fire, harvesting and insects significantly influence net primary productivity (NPP), soil respiration (Rs), net ecosystem production (NEP) and pools of tree biomass and soil organic matter (SOM). The effects of six climate change scenarios demonstrated similar increasing trends of NPP and stand productivity. The disturbances led to a strong decrease in NPP, stand productivity, soil organic matter (SOM) and nitrogen (N) pools with an increase in CO₂ emission to the atmosphere. However the accumulated NEP for 150 years under harvest and fire fluctuated around zero. It becames negative only at a more frequent disturbance regime with four forest fires during the period of simulation. The results from this study show that changes in climate and disturbance regimes might substantially change the NPP as well as the C and N balance, resulting in major changes in the C pools of the vegetation and soil under black spruce forests.     

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.     

Carbon storage in old-growth and second growth fire-dependent western larch (Larix occidentalis Nutt.) forests of the Inland Northwest,USA

There is limited understanding of the carbon (C) storage capacity and overall ecological structure of old-growth forests of western Montana, leaving little ability to evaluate the role of old-growth forests in regional C cycles and ecosystem level C storage capacity. To investigate the difference in C storage between equivalent stands of contrasting age classes and management histories, we surveyed paired old-growth and second growth western larch (Larix occidentalis Nutt)–Douglas-fir (Pseudostuga menziesii var. glauca) stands in northwestern Montana. The specific objectives of this study were to: (1) estimate ecosystem C of old-growth and second growth western larch stands; (2) compare C storage of paired old-growth–second growth stands; and (3) assess differences in ecosystem function and structure between the two age classes, specifically measuring C associated with mineral soil, forest floor, coarse woody debris (CWD), understory, and overstory, as well as overall structure of vegetation. Stands were surveyed using a modified USFS FIA protocol, focusing on ecological components related to soil, forest floor, and overstory C. All downed wood, forest floor, and soil samples were then analyzed for total C and total nitrogen (N). Total ecosystem C in the old-growth forests was significantly greater than that in second growth forests, storing over 3 times the C. Average total mineral soil C was not significantly different in second growth stands compared to old-growth stands; however, total C of the forest floor was significantly greater in old-growth (23.8 Mg ha⁻¹) compared to second growth stands (4.9 Mg ha⁻¹). Overstory and coarse root biomass held the greatest differences in ecosystem C between the two stand types (old-growth, second growth), with nearly 7 times more C in old-growth trees than trees found on second growth stands (144.2 Mg ha⁻¹ vs. 23.8 Mg ha⁻¹). Total CWD on old-growth stands accounted for almost 19 times more C than CWD found in second growth stands. Soil bulk density was also significantly higher on second growth stands some 30+ years after harvest, demonstrating long-term impacts of harvest on soil. Results suggest ecological components specific to old-growth western larch forests, such as coarse root biomass, large amounts of CWD, and a thick forest floor layer are important contributors to long-term C storage within these ecosystems. This, combined with functional implications of contrasts in C distribution and dynamics, suggest that old-growth western larch/Douglas-fir forests are both functionally and structurally distinctive from their second growth counterparts.     

Influence of fire and mountain pine beetle on the dynamics of lodgepole pine stands in British Columbia,Canada

An outbreak of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB), currently affecting over 10.1 million hectares of lodgepole pine forests (Pinus contorta Dougl.) in British Columbia, Canada, is the largest in recorded history. We examined the dynamics of even-aged lodgepole pine forests in southern British Columbia, which were undergoing this MPB outbreak. Using dendroecology and forest measurements we reconstructed the stand processes of stand initiation, stand disturbances, tree mortality, and regeneration, and explained the current stand structure and the potential MPB impacts in selected stands. Our results indicate that stand-replacing fires initiated even-aged seral lodgepole pine stands in this region. In the absence of fire in the 20th century, multiple MPB disturbances, which each resulted in partial canopy removal, modified the simple one-layer structure of the fire-origin stands by the initiation of post-MPB disturbance regeneration layers, transforming the stands into complex, multi-aged stands. Despite high overstory mortality due to the current MPB outbreak, regeneration layers, which are likely to survive the current outbreak, will provide important ecological legacies and will contribute to mid-term timber supply.     

Epiphytic lichen diversity in late-successional Pinus sylvestris forests along local and regional forest utilization gradients in eastern boreal Fennoscandia

Pinus sylvestris-dominated forests have been heavily utilized across all of boreal Fennoscandia and the remaining natural forests are generally highly fragmented. However, there are considerable local and regional differences in the intensity and duration of past forest utilization. We studied the impact of human forest use on the diversity of epiphytic and epixylic lichens in late-successional Pinus sylvestris-dominated forests by assessing species richness and composition along both local and regional gradients in forest utilization. The effects of local logging intensity were analysed by comparing three types of stands: (i) near-natural, (ii) selectively logged (in the early 20th century) and (iii) managed stands. The effects of regional differences in duration and intensity of past forest use were analysed by comparing stands in two contrasting regions (Häme and Kuhmo–Viena). The species richness of selectively logged stands was as high as that of near-natural stands and significantly higher in these two stand categories than in managed stands. Species richness increased with the density of small understorey Picea, which correlated strongly with decreasing intensity of local forest use and increasing structural complexity of selectively logged and near-natural stands. Stands in the Häme region hosted a lower number of species, and were less likely to host many old-growth indicator species than the Kuhmo–Viena region, suggesting that species have been lost from stands in the Häme region due to a longer history of intensive forest use. We conclude that selectively logged stands, along with near-natural stands, are valuable lichen habitats particularly for species confined to old-growth structures such as coarse trees and deadwood. In landscapes where natural forests have become fragmented, the management or restoration of the remaining late-successional Pinus-dominated forests, e.g. through the use of fire, should be carefully planned to avoid adverse effects on lichen species richness.     

Carbon dynamics of North American boreal forest after stand replacing wildfire and clearcut logging

Boreal forest carbon (C) storage and sequestration is a critical element for global C management and is largely disturbance driven. The disturbance regime can be natural or anthropogenic with varying intensity and frequency that differ temporally and spatially the boreal forest. The objective of this review was to synthesize the literature on C dynamics of North American boreal forests after most common disturbances, stand replacing wildfire and clearcut logging. Forest ecosystem C is stored in four major pools: live biomass, dead biomass, organic soil horizons, and mineral soil. Carbon cycling among these pools is inter-related and largely determined by disturbance type and time since disturbance. Following a stand replacing disturbance, (1) live biomass increases rapidly leading to the maximal biomass stage, then stabilizes or slightly declines at old-growth or gap dynamics stage at which late-successional tree species dominate the stand; (2) dead woody material carbon generally follows a U-shaped pattern during succession; (3) forest floor carbon increases throughout stand development; and (4) mineral soil carbon appears to be more or less stable throughout stand development. Wildfire and harvesting differ in many ways, fire being more of a chemical and harvesting a mechanical disturbance. Fire consumes forest floor and small live vegetation and foliage, whereas logging removes large stems. Overall, the effects of the two disturbances on C dynamics in boreal forest are poorly understood. There is also a scarcity of literature dealing with C dynamics of plant coarse and fine roots, understory vegetation, small-sized and buried dead material, forest floor, and mineral soil.