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.     

Thirty-Year Repeat Measures of Mangrove Above- and Below-Ground Biomass Reveals Unexpectedly High Carbon Sequestration

Ecosystems - Mangrove ecosystems store large quantities of organic carbon for long periods of time. This study explores organic carbon stock change through the first comparative study of...     

Above- and Below-Ground Carbon Stocks in an Indigenous Tree (Mytilaria laosensis) Plantation Chronosequence in Subtropical China

More than 60% of the total area of tree plantations in China is in subtropical, and over 70% of subtropical plantations consist of pure stands of coniferous species. Because of the poor ecosystem services provided by pure coniferous plantations and the ecological instability of these stands, a movement is under way to promote indigenous broadleaf plantation cultivation as a promising alternative. However, little is known about the carbon (C) stocks in indigenous broadleaf plantations and their dependence on stand age. Thus, we studied above- and below-ground biomass and C stocks in a chronosequence of Mytilaria laosensis plantations in subtropical China; stands were 7, 10, 18, 23, 29 and 33 years old. Our assessments included tree, shrub, herb and litter layers. We used plot-level inventories and destructive tree sampling to determine vegetation C stocks. We also measured soil C stocks by analyses of soil profiles to 100 cm depth. C stocks in the tree layer dominated the above-ground ecosystem C pool across the chronosequence. C stocks increased with age from 7 to 29 years and plateaued thereafter due to a reduction in tree growth rates. Minor C stocks were found in the shrub and herb layers of all six plantations and their temporal fluctuations were relatively small. C stocks in the litter and soil layers increased with stand age. Total above-ground ecosystem C also increased with stand age. Most increases in C stocks in below-ground and total ecosystems were attributable to increases in soil C content and tree biomass. Therefore, considerations of C sequestration potential in indigenous broadleaf plantations must take stand age into account.     

Changes in the Above- and Below-ground Biomass and Nutrient Pools of Ground Vegetation After Clear-cutting of a Mixed Boreal Forest

Ground vegetation may act as a sink for nutrients after clear-cutting and thus decrease leaching losses. Biomass and nutrient (N, P, K, Ca) pools of ground vegetation (mosses, roots and above-ground parts of field layer) were determined one year before and five years after clear-cutting of a Norway spruce (Picea abies (L.) H. Karst.) dominated boreal mixed forest stand in eastern Finland (63°51′ N, 28°58′ E). Before clear-cutting the average biomass of ground vegetation was 5307 kg ha⁻¹, with nutrient contents of 46.9 kg N ha⁻¹1, 4.1 kg P ha⁻¹1, 16.2 kg K ha⁻¹1 and 13.9 kg Ca ha⁻¹1. The biomass and nutrient pools decreased after clear-cutting being lowest in the second year, the biomass decreasing by 46–65% in the cut plots. The nutrient pools decreased as follows: N 54–72%, P 36–68%, K 51–71% and Ca 57–74%. The decrease in ground vegetation nutrient uptake, and the observed reduced depth of rooting may decrease nutrient retention after clear-cutting and decomposing dead ground vegetation is a potential source of leached nutrients. These negative effects of clear-cutting on the nutrient binding capacity of ground vegetation was short-lived since the total biomass and nutrient pools returned to pre-cutting levels or were even greater by the end of the 5-year study period.     

Understory Plant Community Composition Is Associated with Fine-Scale Above- and Below-Ground Resource Heterogeneity in Mature Lodgepole Pine (Pinus contorta) Forests

Understory plant communities play critical ecological roles in forest ecosystems. Both above- and below-ground ecosystem properties and processes influence these communities but relatively little is known about such effects at fine (i.e., one to several meters within-stand) scales, particularly for forests in which the canopy is dominated by a single species. An improved understanding of these effects is critical for understanding how understory biodiversity is regulated in such forests and for anticipating impacts of changing disturbance regimes. Our primary objective was to examine the patterns of fine-scale variation in understory plant communities and their relationships to above- and below-ground resource and environmental heterogeneity within mature lodgepole pine forests. We assessed composition and diversity of understory vegetation in relation to heterogeneity of both the above-ground (canopy tree density, canopy and tall shrub basal area and cover, downed wood biomass, litter cover) and below-ground (soil nutrient availability, decomposition, forest floor thickness, pH, and phospholipid fatty acids (PLFAs) and multiple carbon-source substrate-induced respiration (MSIR) of the forest floor microbial community) environment. There was notable variation in fine-scale plant community composition; cluster and indicator species analyses of the 24 most commonly occurring understory species distinguished four assemblages, one for which a pioneer forb species had the highest cover levels, and three others that were characterized by different bryophyte species having the highest cover. Constrained ordination (distance-based redundancy analysis) showed that two above-ground (mean tree diameter, litter cover) and eight below-ground (forest floor pH, plant available boron, microbial community composition and function as indicated by MSIR and PLFAs) properties were associated with variation in understory plant community composition. These results provide novel insights into the important ecological associations between understory plant community composition and heterogeneity in ecosystem properties and processes within forests dominated by a single canopy species.     

Carbon Storage Declines in Old Boreal Forests Irrespective of Succession Pathway

The boreal forest plays a critical role in regulating global atmospheric carbon dioxide and is highly influenced by wildfire. However, the long-term recovery of forest carbon (C) storage following wildfire remains unclear, especially during late succession. Uncertainty surrounding C storage in old forests largely stems from both a lack of repeated measurements in forest stands older than the longevity of the pioneer cohort and a lack of consideration of multiple succession pathways. In this study, we constructed a replicated chronosequence, which covered a wide range of forest stand age classes (up to 210 years old) following fire in the boreal forest of central Canada. We selected stands of different overstorey types (that is, broadleaf, conifer, or mixedwood) and age classes to account for multiple succession pathways known to occur in our study area. Our results show a strong relationship between total ecosystem C storage and stand age following fire. Broadleaf stands had on average higher total ecosystem C; however, the inferred temporal dynamics of total ecosystem C were similar among all three overstorey types. Importantly, we found that total ecosystem C storage declined from canopy transition to late-succession stages, irrespective of succession pathway, contradicting views that old forests continually accumulate C as they age. Our findings emphasize that further study of stands older than the longevity of the pioneer cohort is critical to better understand the contribution of old forests to the global C cycle.     

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.     

Carbon Pools and Emissions from Deforestation in Extra-Tropical Forests of Northern Argentina Between 1900 and 2005

We estimated carbon pools and emissions from deforestation in northern Argentine forests between 1900 and 2005, based on forest inventories, deforestation estimates from satellite images and historical data on forests and agriculture. Carbon fluxes were calculated using a book-keeping model. We ran 1000 simulations for a 105-year period with different combinations of values of carbon stocks (Mg C ha⁻¹), soil carbon in the top 0.2 m, and annual deforestation series. The 1000 combinations of parameters were performed as a sensitivity analysis that for each run, randomly selected the values of each variable within a predefined range of values and probability distributions. Using the simulation outputs, we calculated the accumulated C emissions due to deforestation from 1900 to 2005 and the annual emission as the average of the 1000 simulations, and uncertainties of our estimates as the standard deviation. We found that northern Argentine forests contain an estimated 4.54 Pg C (2.312 Pg C in biomass and 2.233 Pg C in soil). Between 1900 and 2005 approximately 30% of the forests were deforested, yielding carbon emissions of 0.945 (SD = 0.270) Pg C. Estimated average annual carbon emissions between 1996 and 2005, mostly from deforestation of the Chaco dry forests, were 20,875 (SD = 6,156) Gg C y⁻¹ (1 Gg = 10⁻⁶ Pg). These values represent the largest source of carbon from land-cover change in the extra-tropical southern hemisphere, between 0.9 and 2.7% of the global carbon emissions from deforestation, and approximately 10% of carbon emissions from the Brazilian Amazon. Deforestation, which has accelerated during the last decades as a result of modern agriculture expansion, represents a major national source of greenhouse gases and the second emission source, after fossil fuel consumption by fixed sources. We conclude that Argentine forests are an important carbon pool and emission source that need more attention for accurate global estimates, and seasonally dry forest deforestation is a key component of the Argentine carbon cycle. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Iron Reduction and Soil Phosphorus Solubilization in Humid Tropical Forests Soils: The Roles of Labile Carbon Pools and an Electron Shuttle Compound

The affinity of iron oxides and hydroxides for phosphorus is thought to contribute to phosphorus limitation to net primary productivity in humid tropical forests on acidic, highly weathered soils. Perennially warm, humid conditions and high biological activity in these soils can result in fluctuating redox potential that in turn leads to considerable iron reduction in the presence of labile carbon and humic substances. We investigated the effects of reducing conditions in combination with the addition of labile carbon substrates (glucose and acetate) and an electron shuttle compound on iron reduction and phosphorus release in a humid tropical forest soil. Glucose or acetate was added to soils as a single dose at the beginning of the experiment, and as pulsed inputs over time, which more closely mimics patterns in labile carbon availability. Iron reduction and phosphorus mobilization were weakly stimulated by a single low level addition of carbon, and the addition of the electron shuttle compound with or without added carbon. Pulsed labile carbon additions produced a significant increase in soil pH, soluble iron, and phosphorus concentrations. Pulsed labile carbon inputs also promoted the precipitation of ferrous hydroxide complexes which could increase the capacity for P sorption, although our results suggest that rates of P solubilization exceeded re-adsorption. Plant and microbial P demand are also likely to serve as an important sinks for released P, limiting the role of P re-adsorption. Our results suggest that reducing conditions coupled with periodic carbon inputs can stimulate iron reduction and a corresponding increase in soil phosphorus mobilization, which may provide a source of phosphorus to plants and microorganisms previously undocumented in these ecosystems.     

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...     

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.     

Above- and Below-ground Growth of Forest Stands: a Carbon Budget Model

The utilization and allocation of carbon by a forest stand areexamined through a simple dynamic, mechanistic model which incorporatesradiation interception, photosynthesis, respiration, assimilatepartitioning, litterfall, root mortality and turnover. Qualitativemathematical analysis of the balance between carbon gains andlosses provides an intuitive insight into the determinants ofabove- and below-ground growth. The patterns of dry matter accumulationproduced by the model are compared with observed trends in standdevelopment. Reported differences between biomass distributionon sites of high and low productivity are reproduced by adjustingthe coefficients of assimilate partitioning. Forest stand growth, model forest growth, carbon budget, photosynthesis, assimilate partitioning     

Impacts of an Invasive Non-Native Annual Weed,Impatiens glandulifera,on Above- and Below-Ground Invertebrate Communities in the United Kingdom

Vegetation community composition and the above- and below-ground invertebrate communities are linked intrinsically, though few studies have assessed the impact of non-native plants on both these parts of the community together. We evaluated the differences in the above- (foliage- and ground-dwelling) and below-ground invertebrate communities in nine uninvaded plots and nine plots invaded by the annual invasive species Impatiens glandulifera, in the UK during 2007 and 2008. Over 139,000 invertebrates were identified into distinct taxa and categorised into functional feeding groups. The impact of I. glandulifera on the vegetation and invertebrate community composition was evaluated using multivariate statistics including principal response curves (PRC) and redundancy analysis (RDA). In the foliage-dwelling community, all functional feeding groups were less abundant in the invaded plots, and the species richness of Coleoptera and Heteroptera was significantly reduced. In the ground-dwelling community, herbivores, detritivores, and predators were all significantly less abundant in the invaded plots. In contrast, these functional groups in the below-ground community appeared to be largely unaffected, and even positively associated with the presence of I. glandulifera. Although the cover of I. glandulifera decreased in the invaded plots in the second year of the study, only the below-ground invertebrate community showed a significant response. These results indicate that the above- and below-ground invertebrate communities respond differently to the presence of I. glandulifera, and these community shifts can potentially lead to a habitat less biologically diverse than surrounding native communities; which could have negative impacts on higher trophic levels and ecosystem functioning.     

Vertical Redistribution of Soil Organic Carbon Pools After Twenty Years of Nitrogen Addition in Two Temperate Coniferous Forests

Nitrogen (N) inputs from atmospheric deposition can increase soil organic carbon (SOC) storage in temperate and boreal forests, thereby mitigating the adverse effects of anthropogenic CO₂ emissions on global climate. However, direct evidence of N-induced SOC sequestration from low-dose, long-term N addition experiments (that is, addition of < 50 kg N ha⁻¹ y⁻¹ for > 10 years) is scarce worldwide and virtually absent for European temperate forests. Here, we examine how tree growth, fine roots, physicochemical soil properties as well as pools of SOC and soil total N responded to 20 years of regular, low-dose N addition in two European coniferous forests in Switzerland and Denmark. At the Swiss site, the addition of 22 kg N ha⁻¹ y⁻¹ (or 1.3 times throughfall deposition) stimulated tree growth, but decreased soil pH and exchangeable calcium. At the Danish site, the addition of 35 kg N ha⁻¹ y⁻¹ (1.5 times throughfall deposition) impaired tree growth, increased fine root biomass and led to an accumulation of N in several belowground pools. At both sites, elevated N inputs increased SOC pools in the moderately decomposed organic horizons, but decreased them in the mineral topsoil. Hence, long-term N addition led to a vertical redistribution of SOC pools, whereas overall SOC storage within 30 cm depth was unaffected. Our results imply that an N-induced shift of SOC from older, mineral-associated pools to younger, unprotected pools might foster the vulnerability of SOC in temperate coniferous forest soils.

     

Correction to: Vertical Redistribution of Soil Organic Carbon Pools After Twenty Years of Nitrogen Addition in Two Temperate Coniferous Forests

Ecosystems - In the article by Forstner et al. (2018), the surnames of co-authors Katharina M. Keiblinger and Patrick Schleppi were misspelled. We apologize and ask readers to cite the corrected...     

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.     

Modelling Variable Fire Severity in Boreal Forests: Effects of Fire Intensity and Stand Structure

It is becoming clear that fires in boreal forests are not uniformly stand-replacing. On the contrary, marked variation in fire severity, measured as tree mortality, has been found both within and among individual fires. It is important to understand the conditions under which this variation can arise. We integrated forest sample plot data, tree allometries and historical forest fire records within a diameter class-structured model of 1.0 ha patches of mono-specific black spruce and jack pine stands in northern Québec, Canada. The model accounts for crown fire initiation and vertical spread into the canopy. It uses empirical relations between fire intensity, scorch height, the percent of crown scorched and tree mortality to simulate fire severity, specifically the percent reduction in patch basal area due to fire-caused mortality. A random forest and a regression tree analysis of a large random sample of simulated fires were used to test for an effect of fireline intensity, stand structure, species composition and pyrogeographic regions on resultant severity. Severity increased with intensity and was lower for jack pine stands. The proportion of simulated fires that burned at high severity (e.g. >75% reduction in patch basal area) was 0.80 for black spruce and 0.11 for jack pine. We identified thresholds in intensity below which there was a marked sensitivity of simulated fire severity to stand structure, and to interactions between intensity and structure. We found no evidence for a residual effect of pyrogeographic region on simulated severity, after the effects of stand structure and species composition were accounted for. The model presented here was able to produce variation in fire severity under a range of fire intensity conditions. This suggests that variation in stand structure is one of the factors causing the observed variation in boreal fire severity.