Contribution of Old Growth Forests to the Carbon Budget of the Boreal Zone in Central Siberia

Old growth (mature and overmature) forests of Central Siberia accumulate carbon not only in the biomass but also in the phytodetritus, including in coarse woody debris (CWD), due to the slow seasonally depressed decomposition of organic matter. Slowly decomposing and accumulating on the soil surface stock of CWD, as a soil humus, serves as a long-term carbon pool in boreal forest ecosystems.     

A Canopy Shift in Interior Alaskan Boreal Forests: Consequences for Above- and Belowground Carbon and Nitrogen Pools during Post-fire Succession

Global change models predict that high-latitude boreal forests will become increasingly susceptible to fire activity as climate warms, possibly causing a positive feedback to warming through fire-driven emissions of CO₂ into the atmosphere. However, fire-climate feedbacks depend on forest regrowth and carbon (C) accumulation over the post-fire successional interval, which is influenced by nitrogen (N) availability. To improve our understanding of post-fire C and N accumulation patterns in boreal forests, we evaluated above- and belowground C and N pools within 70 stands throughout interior Alaska, a region predicted to undergo a shift in canopy dominance as fire severity increases. Stands represented gradients in age and successional trajectory, from black spruce (Picea mariana) self-replacement to species replacement by deciduous species of trembling aspen (Populus tremuloides) and Alaska paper birch (Betula neoalaskana). Stands undergoing deciduous trajectories stored proportionally more of their C and N in aboveground stemwood and had 5–7 times faster rates of aboveground net primary productivity of trees compared to stands undergoing a black spruce trajectory, which stored more of their C and N in the soil organic layer (SOL), a thick layer of mostly undecomposed mosses. Thus, as successional trajectories shift, total C and N pool sizes will remain relatively unchanged, but there will be a trade-off in pool location and a potential increase in C and N longevity due to decreased flammability and decomposition rates of deciduous stemwood. Despite often warmer, drier conditions in deciduous compared to black spruce stands, deciduous stemwood has a C:N around 10 times higher than the black spruce SOL and often remains standing for many years with reduced exposure to fungal decomposers. Thus, a fire-driven shift in successional trajectories could cause a negative feedback to climate warming because of increased pool longevity in deciduous trajectories.     

Variation in Carbon Storage and Its Distribution by Stand Age and Forest Type in Boreal and Temperate Forests in Northeastern China

The northeastern forest region of China is an important component of total temperate and boreal forests in the northern hemisphere. But how carbon (C) pool size and distribution varies among tree, understory, forest floor and soil components, and across stand ages remains unclear. To address this knowledge gap, we selected three major temperate and two major boreal forest types in northeastern (NE) China. Within both forest zones, we focused on four stand age classes (young, mid-aged, mature and over-mature). Results showed that total C storage was greater in temperate than in boreal forests, and greater in older than in younger stands. Tree biomass C was the main C component, and its contribution to the total forest C storage increased with increasing stand age. It ranged from 27.7% in young to 62.8% in over-mature stands in boreal forests and from 26.5% in young to 72.8% in over-mature stands in temperate forests. Results from both forest zones thus confirm the large biomass C storage capacity of old-growth forests. Tree biomass C was influenced by forest zone, stand age, and forest type. Soil C contribution to total forest C storage ranged from 62.5% in young to 30.1% in over-mature stands in boreal and from 70.1% in young to 26.0% in over-mature in temperate forests. Thus soil C storage is a major C pool in forests of NE China. On the other hand, understory and forest floor C jointly contained less than 13% and <5%, in boreal and temperate forests respectively, and thus play a minor role in total forest C storage in NE China.     

Impact of Agricultural Land-use Change on Carbon Storage in Boreal Alaska

Climate warming is most pronounced at high latitudes, which could result in the intensification of the extensively cultivated areas in the boreal zone and could further enhance rates of forest clearing in the coming decades. Using paired forest‐field sampling and a chronosequence approach, we investigated the effect of conversion of boreal forest to agriculture on carbon (C) and nitrogen (N) dynamics in interior Alaska. Chronosequences showed large soil C losses during the first two decades following deforestation, with mean C stocks in agricultural soils being 44% or 8.3 kg m^?2 lower than C stocks in original forest soils. This suggests that soil C losses from land‐use change in the boreal region may be greater than those in other biomes. Analyses of changes in stable C isotopes and in quality of soil organic matter showed that organic C was lost from soils by combustion of cleared forest material, decomposition of organic matter and possibly erosion. Chronosequences indicated an increase in C storage during later decades after forest clearing, with 60‐year‐old grassland showing net ecosystem C gain of 2.1 kg m^?2 over the original forest. This increase in C stock resulted probably from a combination of large C inputs from belowground biomass and low C losses due to a small original forest soil C stock and low tillage frequency. Reductions in soil N stocks caused by land‐use change were smaller than reductions in C stocks (34% or 0.31 kg m^?2), resulting in lower C/N ratios in field compared with forest mineral soils, despite the occasional incorporation of high‐C forest‐floor material into field soils. Carbon mineralization per unit of mineralized N was considerably higher in forests than in fields, which could indicate that decomposition rates are more sensitive in forest soils than in field soils to inorganic N addition (e.g. by increased N deposition from the atmosphere). If forest conversion to agriculture becomes more widespread in the boreal region, the resulting C losses (51% or 11.2 kg m^?2 at the ecosystem level in this study) will induce a positive feedback to climatic warming and additional land‐use change. However, by selecting relatively C‐poor soils and by implementing management practices that preserve C, losses of C from soils can be reduced.     

Mapping Above- and Below-Ground Carbon Pools in Boreal Forests: The Case for Airborne Lidar

A large and growing body of evidence has demonstrated that airborne scanning light detection and ranging (lidar) systems can be an effective tool in measuring and monitoring above-ground forest tree biomass. However, the potential of lidar as an all-round tool for assisting in assessment of carbon (C) stocks in soil and non-tree vegetation components of the forest ecosystem has been given much less attention. Here we combine the use airborne small footprint scanning lidar with fine-scale spatial C data relating to vegetation and the soil surface to describe and contrast the size and spatial distribution of C pools within and among multilayered Norway spruce (Picea abies) stands. Predictor variables from lidar derived metrics delivered precise models of above- and below-ground tree C, which comprised the largest C pool in our study stands. We also found evidence that lidar canopy data correlated well with the variation in field layer C stock, consisting mainly of ericaceous dwarf shrubs and herbaceous plants. However, lidar metrics derived directly from understory echoes did not yield significant models. Furthermore, our results indicate that the variation in both the mosses and soil organic layer C stock plots appears less influenced by differences in stand structure properties than topographical gradients. By using topographical models from lidar ground returns we were able to establish a strong correlation between lidar data and the organic layer C stock at a stand level. Increasing the topographical resolution from plot averages (~2000 m²) towards individual grid cells (1 m²) did not yield consistent models. Our study demonstrates a connection between the size and distribution of different forest C pools and models derived from airborne lidar data, providing a foundation for future research concerning the use of lidar for assessing and monitoring boreal forest C.     

Recovery of Ecosystem Carbon Stocks in Young Boreal Forests: A Comparison of Harvesting and Wildfire Disturbance

Corresponding with the increasing global resource demand, harvesting now affects millions of hectares of boreal forest each year, and yet our understanding of harvesting impacts on boreal carbon (C) dynamics relative to wildfire remains unclear. We provide a direct comparison of C stocks following clearcut harvesting and fire over a 27-year chronosequence in the boreal forest of central Canada. Whereas many past studies have lacked measurement of all major C pools, we attempt to provide complete C pool coverage, including live biomass, deadwood, forest floor, and mineral soil C pools. The relative contribution of each C pool to total ecosystem C varied considerably between disturbance types. Live biomass C was significantly higher following harvesting compared with fire because of residual live trees and advanced regeneration. Conversely, most live biomass was killed following fire, and thus post-fire stands contained higher stocks of deadwood C. Snag and stump C mass peaked immediately following fire, but dramatically decreased 8 years after fire as dead trees began to fall over, contributing to the downed woody debris C pool. Forest floor C mass was substantially lower shortly after fire than harvesting, but this pool converged 8 years after fire and harvesting. When total ecosystem C stocks were examined, we found no significant difference during early stand development between harvesting and fire. Maximum total ecosystem C occurred at age 27 years, 185.1 ± 18.2 and 163.6 ± 8.0 Mg C ha^?1 for harvesting and fire, respectively. Our results indicate strong differences in individual C pools, but similar total ecosystem C after fire and clearcutting in boreal forests, and shall help improve modeling terrestrial C flux after stand-replacing disturbances.     

Stand Structural Dynamics of North American Boreal Forests

Stand structure, the arrangement and interrelationships of live and dead trees, has been linked to forest regeneration, nutrient cycling, wildlife habitat, and climate regulation. The objective of this review was to synthesize literature on stand structural dynamics of North American boreal forests, addressing both live tree and coarse woody debris (CWD) characteristics under different disturbance mechanisms (fire, clearcut, wind, and spruce budworm), while identifying regional differences based on climate and surficial deposit variability. In fire origin stands, both live tree and CWD attributes are influenced initially largely by the characteristics of the stand replacing fire and later increasingly by autogenic processes. Differences in stand structure have also been observed between various stand cover types. Blowdown and insect outbreaks are two significant non-stand replacing disturbances that can alter forest stand structure through removing canopy trees, freeing up available growing space, and creating microsites for new trees to establish. Climate and surficial deposits are highly variable in the boreal forest due to its extensive geographic range, influencing stand and landscape structure by affecting tree colonization, stand composition, successional trajectories, CWD dynamics, and disturbance regimes including regional fire cycles. Further, predicted climate change scenarios are likely to cause regional-specific alterations in stand and landscape structure, with the implications on ecosystem components including wildlife, biodiversity, and carbon balance still unclear. Some stand structural attributes are found to be similar between clearcut and fire origin stands, but others appear to be quite different. Future research shall focus on examining structural variability under both disturbance regimes and management alternatives emulating both stand replacing and non-stand replacing natural disturbances.

     

Understory Vegetation Dynamics of North American Boreal Forests

Understory vegetation is the most diverse and least understood component of North American boreal forests. Understory communities are important as they act as drivers of overstory succession and nutrient cycling. The objective of this review was to examine how understory vegetation abundance, composition, and diversity change with stand development after a major stand replacing disturbance. Understory vegetation abundance and diversity increase rapidly after fire, in response to abundant resources and an influx of disturbance adapted species. The highest diversity occurs within the first 40 years following fire, and declines indefinitely thereafter as a result of decreasing productivity and increased dominance of a small number of late successional feather mosses and woody plant species. Vascular plant and bryophyte/lichen communities undergo very different successional changes. Vascular plant communities are dynamic and change more dramatically with time after fire, whereas bryophyte and lichen communities are much slower to establish and change over time. Considerable variations in these processes exist depending on canopy composition, site condition, regional climate, and frequently occurring non-stand-replacing disturbances. Forest management practices represent a unique disturbance process and can result in different understory vegetation communities from those observed for natural processes, with potential implications for overstory succession and long-term productivity. Because of the importance of understory vegetation on nutrient cycling and overstory composition, post-harvest treatments emulating stand-replacing fire are required to maintain understory diversity, composition, and promote stand productivity in boreal forests.     

Carbon Accumulation Patterns During Post-Fire Succession in Cajander Larch (Larix cajanderi) Forests of Siberia

Increased fire activity within boreal forests could affect global terrestrial carbon (C) stocks by decreasing stand age or altering tree recruitment, leading to patterns of forest regrowth that differ from those of pre-fire stands. To improve our understanding of post-fire C accumulation patterns within boreal forests, we evaluated above- and belowground C pools within 17 Cajander larch (Larix cajanderi) stands of northeastern Siberia that varied in both years since fire and stand density. Early-successional stands (<20-year old) exhibited low larch recruitment, and consequently, low density, aboveground larch biomass, and aboveground net primary productivity (ANPP_tree). Mid-successional stands (21- to 70-year old) were even-aged with considerable variability in stand density. High-density mid-successional stands had 21 times faster rates of ANPP_tree than low-density stands (252 vs. 12?g?C?m^?2?y^?1) and 26 times more C in aboveground larch biomass (2,186 vs. 85?g?C?m^?2). Density had little effect on total soil C pools. During late-succession (>70-year old), aboveground larch biomass, ANPP_tree, and soil organic layer C pools increased with stand age. These stands were low density and multi-aged, containing both mature trees and new recruits. The rapid accumulation of aboveground larch biomass in high-density, mid-successional stands allowed them to obtain C stocks similar to those in much older low-density stands (~8,000?g?C?m^?2). If fire frequency increases without altering stand density, landscape-level C storage could decline, but if larch density also increases, large aboveground C pools within high-density stands could compensate for a shorter successional cycle.     

Refugium of the Boreal Forests of the Circumpolar Urals

The results of studies of the vegetative cover and ontogenetic composition of the cenopopulations of Picea obovata Ledeb. and Larix sibirica Ledeb. in the Gorno-Khadytinskii Forest Reserve, which is the refugium of the northern taiga forests of Western Siberia, have been summarized. According to the Brown–Blanke system, two associations of forests were identified and described, as well as two tundra associations, for comparison. The unique association of the Aconito septentrionalis–Piceetum obovatae taiga forests Zaugolnova et Morozova 2009 were analyzed in detail, and a comparison of these associations was carried out in the Gorno-Khadytinskii and Pechoro-Ilychskii reserves, where these forests are the reference standards of the zonal-type forests. An estimate of the ontogenetic composition of the cenopopulations of P. obovata Ledeb. and L. sibirica Ledeb. is given, as well as the absolute age of the trees of these species.     

Simulation and Analysis of the Effect of a Spruce Budworm Outbreak on Carbon Dynamics in Boreal Forests of Quebec

Ecosystems - In a climate change context, the sequestration of atmospheric carbon (C) in forests is key for achieving emission targets. It is thus critical to understand how large-scale...     

Hemlock Declines Rapidly with Hemlock Woolly Adelgid Infestation: Impacts on the Carbon Cycle of Southern Appalachian Forests

The recent infestation of southern Appalachian eastern hemlock stands by hemlock woolly adelgid (HWA) is expected to have dramatic and lasting effects on forest structure and function. We studied the short-term changes to the carbon cycle in a mixed stand of hemlock and hardwoods, where hemlock was declining due to either girdling or HWA infestation. We expected that hemlock would decline more rapidly from girdling than from HWA infestation. Unexpectedly, in response to both girdling and HWA infestation, hemlock basal area increment (BAI) reduced substantially compared to reference hardwoods in 3 years. This decline was concurrent with moderate increases in the BAI of co-occurring hardwoods. Although the girdling treatment resulted in an initial pulse of hemlock needle inputs, cumulative litter inputs and O horizon mass did not differ between treatments over the study period. Following girdling and HWA infestation, very fine root biomass declined by 20–40% in 2 years, which suggests hemlock root mortality in the girdling treatment, and a reduction in hemlock root production in the HWA treatment. Soil CO₂ efflux (E _soil) declined by approximately 20% in 1 year after both girdling and HWA infestation, even after accounting for the intra-annual variability of soil temperature and moisture. The reduction in E _soil and the concurrent declines in BAI and standing very fine root biomass suggest rapid declines in hemlock productivity from HWA infestation. The accelerated inputs of detritus resulting from hemlock mortality are likely to influence carbon and nutrient fluxes, and dictate future patterns of species regeneration in these forest ecosystems. AEN performed research and analyzed data; NW performed research, analyzed data, and wrote the article; CRF contributed new methods, analyzed data, and wrote the article; RLH designed the study; JMV conceived of and designed the study; and BDK performed research.     

Effect of Climate Change on Soil Temperature in Swedish Boreal Forests

Complex non-linear relationships exist between air and soil temperature responses to climate change. Despite its influence on hydrological and biogeochemical processes, soil temperature has received less attention in climate impact studies. Here we present and apply an empirical soil temperature model to four forest sites along a climatic gradient of Sweden. Future air and soil temperature were projected using an ensemble of regional climate models. Annual average air and soil temperatures were projected to increase, but complex dynamics were projected on a seasonal scale. Future changes in winter soil temperature were strongly dependent on projected snow cover. At the northernmost site, winter soil temperatures changed very little due to insulating effects of snow cover but southern sites with little or no snow cover showed the largest projected winter soil warming. Projected soil warming was greatest in the spring (up to 4°C) in the north, suggesting earlier snowmelt, extension of growing season length and possible northward shifts in the boreal biome. This showed that the projected effects of climate change on soil temperature in snow dominated regions are complex and general assumptions of future soil temperature responses to climate change based on air temperature alone are inadequate and should be avoided in boreal regions.     

Plant Community Composition as a Predictor of Regional Soil Carbon Storage in Alaskan Boreal Black Spruce Ecosystems

The boreal forest is the largest terrestrial biome in North America and holds a large portion of the world’s reactive soil carbon. Therefore, understanding soil carbon accumulation on a landscape or regional scale across the boreal forest is useful for predicting future soil carbon storage. Here, we examined the relationship between floristic composition and ecosystem parameters, such as soil carbon pools, the carbon-to-nitrogen (C/N) ratio of live black spruce needles, and normalized basal area increment (NBAI) of trees in black spruce communities, the most widespread forest type in the boreal forest of Alaska. Variability in ecosystem properties among black spruce stands was as large as that which had previously been documented among all forest types in the central interior of Alaska; we found an eightfold range in NBAI and fivefold range in mineral soil carbon and nitrogen pools. Acidic black spruce communities had significantly more carbon in the organic soil horizon than did nonacidic black spruce communities, but did not differ in any other measured ecosystem parameter. We explained 48% of the variation in total soil carbon with a combination of plant community indices and abiotic and biotic factors. Plant community composition was at least as effective as any single environmental factor or stand characteristic in predicting soil C pools in Alaskan black spruce ecosystems. We conclude that among the community properties analyzed, the presence of key groups of species, overall species composition, and diversity of certain functional types, especially Sphagnum moss species, are important predictors of soil carbon sequestration in the black spruce forest type.     

Fire Interval Effects on Successional Trajectory in Boreal Forests of Northwest Canada

Although succession may follow multiple pathways in a given environment, the causes of such variation are often elusive. This paper describes how changes in fire interval mediate successional trajectory in conifer-dominated boreal forests of northwestern Canada. Tree densities were measured 5 and 19 years after fire in permanent plots and related to pre-fire vegetation, site and fire characteristics. In stands that were greater than 75 years of age when they burned, recruitment density of conifers was significantly correlated with pre-fire species basal area, supporting the expectation of stand self-replacement as the most common successional pathway in these forests. In contrast, stands that were under 25 years of age at the time of burning had significantly reduced conifer recruitment, but showed no change in recruitment of trembling aspen (Populus tremuloides). As a result, young-burned stands had a much higher probability of regenerating to deciduous dominance than mature-burned stands, despite the dominance of both groups by spruce (Picea mariana and Picea glauca) and pine (Pinus contorta) before the fire. Once initiated, deciduous-dominated stands may be maintained across subsequent fire cycles through mechanisms such as low on-site availability of conifer seed, competition with the aspen canopy, and rapid asexual regeneration of aspen after fire. We suggest that climate-related increases in fire frequency could trigger more frequent shifts from conifer to deciduous-dominated successional trajectories in the future, with consequent effects on multiple ecosystem processes.     

Relationships between Stand Composition and Ectomycorrhizal Community Structure in Boreal Mixed-Wood Forests

We investigated the community structure of ectomycorrhizal fungi under varying overstory tree compositions in the southern mixed-wood boreal forest of Quebec. Sampling took place at two locations of differing postfire ages and nine 100-m² plots were sampled per location. The dominant overstory tree species in the plots were trembling aspen (Populus tremuloides Michx.), white birch (Betula papyrifera Marsh.) or white spruce [Picea glauca (Moench) Voss], and balsam fir [Abies balsamea (L.) Mill.]. Mycorrhizae were analyzed using morphological as well as molecular methods, employing fungal-specific primers to amplify ribosomal DNA for subsequent cloning and sequencing. A total of 1800 mycorrhizal root tips collected from the 18 plots were morphologically classified into 26 morphotypes, with Cenococcum geophilum dominating (36% of root tips). A second set of root tips, selected from the same 18 samples on which the morphological analysis was based, were analyzed using molecular methods. From this analysis, 576 cloned polymerase chain reaction products were screened by restriction fragment length polymorphism analysis and a total of 207 unique types were found. No one type dominated the system and 159 occurred only once. Sequence analysis of the types that occurred more than once revealed that Piloderma sp., Russula sp., Cortinarius sp., and Lactarius sp. were the most common mycorrhizae. The ectomycorrhizal fungal community structure revealed by the rDNA analysis differed from that observed using morphological methods. Canonical correspondence analyses of the sequenced restriction types and % overstory composition indicate that the distributions of ectomycorrhizal fungi are influenced by the relative proportions of host tree species. The distinct fungal assemblages found in the different plots supported by the different combinations of host tree species provides further support for the need to conserve stand diversity in the southern boreal forest.     

Effects of Retention Felling on Epixylic Species in Boreal Spruce Forests in Southern Finland

We studied the effects of patch retention felling and soil scarification by harrowing on the coverage and species richness of epixylic species in boreal Norway spruce (Picea abies) forests in Southern Finland. The epixylics were investigated from both the retention tree groups (RTGs) and the surrounding felling areas before and after fellings and after scarification on consecutive years. The cover percentage of all included species groups was shown to decrease after the felling, especially in the felling areas (vascular plants, ?0.4%; mosses, ?27.8%; liverworts, ?4.0%; and lichens,?2.1%). The decrease was considerable also in the RTGs. The amount of dead moss increased in both the areas indicating microclimatic changes. Species richness also declined rapidly after the first year, especially in the felling areas (vascular plants, ?2.2%; mosses, ?27.3%; liverworts, ?30.3%; and lichens, ?22.9%). Scarification also decreases covers and species richness of bryophytes. After the second year, the covers of the species groups generally started to regain, especially in the untreated RTGs. The size of RTG was in positive correlation with the total species number. Another main result indicates that it is possible to maintain much higher initial vegetation abundance and diversity in the RTGs than in the felling areas. Coarse woody debris formed by the frequent tree uprootings may also enhance the long‐term survival of epixylics over forest regeneration period. RTGs should be at least 10 times larger than the size used in current Finnish forestry, so that they could function as species refugia.