Boreal Black Spruce Forest Research Articles

What explains the year-to-year variation in growing season timing of boreal black spruce forests?

Amplified climate warming in high latitudes is expected to affect growing season timing of the vast boreal biome. It is unclear whether the presence of permafrost (perennially frozen ground) might have an influence on changes in growing season timing. This study examined how different environmental variables explained, either directly or indirectly, the variation in growing season timing of boreal forest stands with and without permafrost. We expected that environmental variables explaining the variation in growing season timing differed or had different explanatory power depending on permafrost presence or absence. The growing season was delineated from daily gross primary productivity (GPP) time series derived from 40 site-year data of net ecosystem carbon dioxide exchange measured with eddy covariance techniques over five black spruce (Picea mariana [Mill.])-dominated boreal forest stands in North America. In permafrost-free forest stands, a combination of start in canopy ‘green-up’ in spring and the timing of air and soil temperature increasing above freezing explained the start-of-season (SOSGPP). Results from commonality analysis and structural equation modeling suggest that canopy ‘green-up’ and air temperature directly affected SOSGPP in permafrost-free forest stands. In addition, soil temperature acted as mediator for an indirect effect of air temperature on SOSGPP. In contrast, none of the environmental variables, or their combination, explained the variation in SOSGPP in forest stands with permafrost. The explanatory power of environmental variables was more consistent regarding the end-of-season (EOSGPP). In both, forest stands with and without permafrost, EOSGPP was directly explained by mean soil water content in the fall and the first day of continuous snowpack formation. A better understanding how environmental variables control SOSGPP and EOSGPP in forest stands with and without permafrost will help to refine parameterizations of the boreal biome in Earth system models.

The effects of clearcut harvesting on moss chloroplast lipidome and adaptation to light stress during boreal forest regeneration

Moss plays an important role in boreal forest ecosystems as an understory bryophyte species. Clearcut harvesting is a common boreal forest regeneration method that can expose understory vegetation to abiotic stressors impeding their recovery following post-harvest conditions. Very little is known concerning how moss remodel their chloroplast lipidome to enhance photosynthetic performance for successful acclimation to light and water stress during boreal forest regeneration following clearcut harvesting. The chloroplast lipidome and photosynthetic performance of Sphagnum sp. and three feathermoss species (Pleurozium schreberi, Hylocomium splendens, and Ptilium crista-castrensis) from a boreal black spruce (Picea mariana) forest were assessed using liquid chromatography-mass spectrometry (LC-MS), photospectrometry, and light response curves. We observed an overall increase in monogalactosyldiacylglycerol (MGDG) and sulfoquinovosyldiacylglycerol (SQDG) and decrease in digalactosyldiacylglycerol (DGDG) and phosphatidylglycerol (PG). In addition, unsaturation of the chloroplast lipidome occurred concomitant with photoprotection by carotenoid pigments to enhance the efficiency and photosynthetic capacity in moss exposed to light and water stress following clearcut harvesting. This appears to be a successful acclimation strategy used by moss to circumvent light stress during boreal forest regeneration following clearcut harvesting. These findings could be of significance in the development of boreal forest management strategies following resource harvesting.

Bryophyte Conservation in Managed Boreal Landscapes: Fourteen-Year Impacts of Partial Cuts on Epixylic Bryophytes

Clear cut harvest simplifies and eliminates old growth forest structure, negatively impacting biodiversity. Partial cut harvest has been hypothesized (1) to have less impact on biodiversity than clear cut harvest, and (2) to encourage old growth forest structures. Long-term studies are required to test this hypothesis as most studies are conducted soon after harvest. Using epixylic bryophytes as indicators, this study addresses this knowledge gap. Fourteen years after harvest, we examined changes in epixylic bryophyte community composition richness and traits, and their microhabitats (coarse woody debris characteristics and microclimate) along an unharvested, partial cuts and clear cuts harvest treatment in 30 permanent plots established in the boreal black spruce (Picea mariana) forests of northwestern Quebec, Canada. Our results were compared to those of an initial post-harvest study (year 5) and to a chronosequence of old growth forests to examine species changes over time and the similarity of bryophyte communities in partial cut and old growth forests. Coarse woody debris (CWD) volume by decay class varied among harvest treatments with partial cuts and clear cuts recording lower volumes of early decay CWD. The epixylic community was richer in partial cuts than in mature unharvested forests and clear cuts. In addition, species richness and overall abundance doubled in partial and clear cuts between years 5 and 14. Species composition also differed among treatments between years 5 and 14. Furthermore, conditions in partial cut stands supported small, drought sensitive, and old growth confined species that are threatened by conditions in clear cut stands. Lastly, over time, species composition in partial cuts became more similar to old growth forests. Partial cuts reduced harvest impacts by continuing to provide favorable microhabitat conditions that support epixylic bryophytes. Also, partial cut harvest has the potential to encourage old growth species assemblages, which has been a major concern for biodiversity conservation in managed forest landscapes. Our findings support the promotion of partial cut harvest as an effective strategy to achieve species and habitat conservation goals.

Annual aboveground carbon uptake enhancements from assisted gene flow in boreal black spruce forests are not long-lasting

Assisted gene flow between populations has been proposed as an adaptive forest management strategy that could contribute to the sequestration of carbon. Here we provide an assessment of the mitigation potential of assisted gene flow in 46 populations of the widespread boreal conifer Picea mariana, grown in two 42-year-old common garden experiments and established in contrasting Canadian boreal regions. We use a dendroecological approach taking into account phylogeographic structure to retrospectively analyse population phenotypic variability in annual aboveground net primary productivity (NPP). We compare population NPP phenotypes to detect signals of adaptive variation and/or the presence of phenotypic clines across tree lifespans, and assess genotype‐by‐environment interactions by evaluating climate and NPP relationships. Our results show a positive effect of assisted gene flow for a period of approximately 15 years following planting, after which there was little to no effect. Although not long lasting, well-informed assisted gene flow could accelerate the transition from carbon source to carbon sink after disturbance.

Study of Fuel-Smoke Dynamics in a Prescribed Fire of Boreal Black Spruce Forest through Field-Deployable Micro Sensor Systems

Understanding the combustion dynamics of fuels, and the generation and propagation of smoke in a wildland fire, can inform short-range and long-range pollutant transport models, and help address and mitigate air quality concerns in communities. Smoldering smoke can cause health issues in nearby valley bottoms, and can create hazardous road conditions due to low-visibility. We studied near-field smoke dynamics in a prescribed fire of 3.4 hectares of land in a boreal black spruce forest in central Alberta. Smoke generated from the fire was monitored through a network of five field-deployable micro sensor systems. Sensors were placed within 500–1000 m of the fire area at various angles in downwind. Smoke generated from flaming and smoldering combustions showed distinct characteristics. The propagation rates of flaming and smoldering smoke, based on the fine particulate (PM2.5) component, were 0.8 and 0.2 m/s, respectively. The flaming smoke was characterized by sharp rise of PM2.5 in air with concentrations of up to 940 µg/m3, followed by an exponential decay with a half-life of ~10 min. Smoldering combustion related smoke contributed to PM2.5 concentrations above 1000 µg/m3 with slower decay half-life of ~18 min. PM2.5 emissions from the burn area during flaming and smoldering phases, integrated over the combustion duration of 2.5 h, were ~15 and ~16 kilograms, respectively, as estimated by our mass balance model.

Recent Crown Thinning in a Boreal Black Spruce Forest Does Not Reduce Spread Rate nor Total Fuel Consumption: Results from an Experimental Crown Fire in Alberta, Canada

A 3.6 ha experimental fire was conducted in a black spruce peatland forest that had undergone thinning the year prior. After 50 m of spread in a natural stand at 35–60 m min−1, the crown fire (43,000 kW m−1 intensity using Byram’s method) encountered the 50% stem removal treatment; spread rates in the treatment were 50–60 m min−1. Fuel consumption in the control (2.75 kg m−2) was comparable to the treatment (2.35 kg m−2). Proxy measurements of fire intensity using in-stand heat flux sensors as well as photogrammetric flame heights had detected intensity reductions to 30–40% of the control. Crown fuel load reductions (compensated by higher surface fuel load) appear to be the most significant contributor to the decline in intensity, despite drier surface fuels in the treatment. The burn depth of 5 cm in moss and organic soil did not differ between control and treatment. These observations point to the limited effectiveness (likely reductions in crown fire intensity but not spread rate) of stem removal in boreal black spruce fuel types with high stem density, low crown base height and high surface fuel load. The observed fire behaviour impacts differ from drier conifer forests across North America.

Changes in forest structure along a chronosequence in the black spruce boreal forest: Identifying structures to be reproduced through silvicultural practices

In managed boreal forests, partial harvesting has been proposed to promote forest structural complexity and to therefore maintain associated biodiversity. However, there have been few studies identifying forest structures that should be maintained within the forest matrix, and fewer still on changes in these structures during succession. Consequently, there is no tool to identify these different structures in the field or their sequence along natural succession. This study proposes a key that can be used in the field and allows for the identification of different forest diameter structures along a successional sequence in the black spruce boreal forest. The specific objectives of this study were (1) to classify the types of forest structures encountered in natural black spruce boreal forest based on their diameter distribution, and (2) to link this classification to time since last fire and its spatial homogeneity at the stand level. This study shows that the forest stand structure, in black spruce forests, is varied and that this structural variety is mainly controlled by time since last fire. It also shows that the timing of stand structural maturation varies with severity of the last fire and surficial deposit. The identification key suggests that based on tree diameter distribution it is possible to discriminate among young, mature and old forest structures, which could help forest managers select stands to be harvested according to different objectives and hence maintain the variety of black spruce forest structures at the landscape scale.

Sensitivity of Boreal Carbon Stocks to Fire Return Interval, Fire Severity and Fire Season: A Simulation Study of Black Spruce Forests

Boreal forests store substantial amounts of carbon in vegetation and soil pools. The magnitude of these pools is related to fire regime attributes. Climate change is expected to alter boreal fire regimes, leading to changes in the amount of stored carbon. Quantifying these changes is of importance to understanding and managing global carbon budgets. We investigate how fire return interval (FRI) interacts with seasonal variation in fire intensity and severity to affect carbon stocks and fluxes in the boreal black spruce forests of Quebec, Canada. A size-class structured population model of stand dynamics was coupled with an established model of boreal carbon dynamics and linked to a simplified representation of fire regime that simulates the occurrence of fires and their direct effects on canopy tree mortality, surface fuels combustion and regeneration. We simulated carbon stocks and fluxes under seven levels of FRI and two fire seasons: spring and summer. We tested for an effect of these fire regime parameters on equilibrium mean C stocks. All dead organic matter and biomass carbon stocks were sensitive to FRI between 60 and 300 years. These C stocks were lower for summer fires that occurred under shorter FRIs. Net primary production was highest at FRIs between 150 and 300 years. Total C stocks were highest for FRIs from 150 to 700 years, varying little over this range. There was a small but significant difference of C pool sizes between stands with even and uneven tree-diameter distributions. This difference was greatest for FRIs of 150 years or less. Reductions in equilibrium C storage are forecasted for nearly 27% of the study region under expected end-of-century climates.

Range Extensions of 35 Bryophyte Species in the Black Spruce–Feather Moss Forest of Western Quebec, Canada

Although the North American bryophyte flora are relatively well known, bryophytes of the Black Spruce–feather moss forest in the Nord-du-Québec administrative region, especially its southern portion (49–51°N, 74–79°W), remain under-sampled. Here, we report 169 bryophyte taxa for this region, of which 35 (14 true mosses, 20 liverworts, one sphagnum) represent noteworthy records, including 20 taxa new for the region. These new occurrences close several gaps in distribution in the study area and, more broadly, in the boreal Black Spruce (Picea mariana) forest of adjacent ontario. Microhabitat preferences of the species are also documented. This work represents a substantial contribution to knowledge of the bryophyte flora, which will help refine protection priority ranks of species of Quebec and Labrador.

Are post-fire residual forest patches refugia for boreal bryophyte species? Implications for ecosystem based management and conservation

Residual patches of forest remaining after natural or anthropogenic disturbance may facilitate regeneration of fragmented forest. However, residual patch function remains unclear, especially after natural wildfire. We investigate the role of residual boreal forest patches as refugia for bryophytes and ask the question, do they house bryophyte communities similar to those encountered in undisturbed forests? Bryophytes were sampled in three habitat types in black spruce boreal forests illustrating a gradient of disturbance severity: undisturbed forests, residual patches and burned matrices. Temporal, disturbance severity, spatial and structural variables of habitats were also recorded. Bryophyte community composition differed among habitat types with residual patches characterized by higher species richness, the loss of forest specialists and the addition of disturbance-prone species. The bryophyte community found in residual patches is at the interface between the communities of undisturbed forests and burned matrices. As residual patches did not conserve all species, particularly forest specialists, they were not refugia. However, we identify temporal, spatial and structural characteristics that can maintain bryophyte communities most similar to undisturbed forests and enhance residual patch “refugia potential”. Residual patches enhance bryophyte diversity of the landscape housing species that cannot survive in the burned matrix. As conclusion we discuss the use of retention patches in harvested stands, together with the preservation of undisturbed stands that house singular bryophyte communities and especially sensitive forest specialists.

Carbon neutral or a sink? Uncertainty caused by gap-filling long-term flux measurements for an old-growth boreal black spruce forest

Old-growth (>100years old) boreal forests are recognized as having accumulated large inventories of terrestrial carbon, but the magnitude of annual net exchange is not well established. Eddy covariance measurements of carbon dioxide exchange were made at a flux tower at the Northern Old Black Spruce forest in Manitoba, Canada, from 1994 to 2008. We evaluated the uncertainty in estimates of annual net ecosystem production by comparing four methods of filling missing measurement periods. Three of the methods were previously developed by groups at Harvard University, the Fluxnet Canada Research Network, and the Max Planck Institute of Biogeochemistry; and the fourth method used the mean of measurements from the dataset selected for the same time and day. Two methods showed carbon losses near the beginning of the period but all four methods estimated carbon gains in the later years. Individual years and methods ranged ±90gCm−2 y−1, demonstrating a wide range of estimates if data had been available from only a single year and a single method. When averaged over multiple years, the mean and standard error among methods gave a net forest carbon sink of 29±10gCm−2 y−1, ranging from 4 to 48gCm−2 y−1. Only the upper (Mean Data and Max Planck Institute of Biogeochemistry methods) and lower (Harvard method) estimates were significantly different, with the Fluxnet Canada Research Network method being the same as all other methods. There was no difference among methods for the January to August period, or for daytime periods. All methods were statistically different from a carbon-neutral forest suggesting that the Northern Old Black Spruce forest was likely a small carbon sink. Methods that estimated high ecosystem respiration also estimated high gross ecosystem production, and the small net differences could not be easily attributed to differences in flux partitioning. For forests that are close to carbon neutral, the use of several gap-filling methods: increases confidence in conclusions regarding the ecosystem carbon balance.

Can Retention Harvest Maintain Natural Structural Complexity? A Comparison of Post-Harvest and Post-Fire Residual Patches in Boreal Forest

Variable retention harvest promotes biodiversity conservation in managed boreal forests by ensuring forest continuity and structural complexity. However, do post-harvest and post-fire patches maintain the same structural complexity? This study compares post-harvest and post-fire residual patches and proposes retention modalities that can maintain the same structural complexity as in natural forests, here considering both continuous forest stands and post-fire residual patches. In boreal black spruce forests, 41 post-fire residual patches, and 45 post-harvest retention patches of varying size and ages (exposure time to disturbed matrix) and 37 continuous forest stands were classified into six diameter structure types. Types 1 (inverted-J) and 2 (trunked-unimodal) characterized stands dominated by small trees. The abundance of small trees decreased and the abundance of large trees increased from Type 1 to Type 6. Type 6 had the most irregular structure with a wide range of diameters. This study indicates that: (1) old post-harvest residual retentions maintained the range of structural complexity found in natural stands; (2) Types 1 and 2 were generally associated with young post-fire patches and post-harvest retention clumps; (3) the structure of residual patches containing only small trees was usually younger (in terms of the age of the original forest from which residual patches were formed) than those with larger trees. To avoid the risk of simplifying the structure, retention patches should be intentionally oriented towards Types 3–6, dominated by intermediate and large trees.

More frequent burning increases vulnerability of Alaskan boreal black spruce forests

Much recent research has investigated the effects of burning on mature black spruce (Picea mariana) forests in interior Alaska, however little research has focused on how frequent reburning affects soil organic layer (SOL) vulnerability in these ecosystems. We compared organic soil layer characteristics in black spruce stands that burned after two fire-free intervals (FFI), including an intermediate-interval (37–52 years) and a more typical long-interval (70–120 years). We found that depth of burn varied significantly between intermediate-interval and long-interval sites, and as there was less material available to burn in intermediate-interval stands, percent depth reduction was greater in these stands (78.9% ± 2.6%) than in long-interval stands (62.9% ± 1.5%). As a result, less residual organic soil carbon remained post-fire in intermediate-interval than long-interval stands. Post-fire organic soil carbon stocks averaged 0.51 ± 0.08 kg C m−2 in the intermediate-interval sites, which is less than estimates of soil carbon stock for long-interval fire events (ranging from 2.07 to 5.74 kg C m−2). In addition to altering soil carbon storage, a depletion of the SOL during more frequent fire events will likely delay the recovery of permafrost and could trigger a change in the possible successional trajectory of a site, from black spruce dominated to deciduous or even shrub dominated ecosystems in the future.

Prescribed burning of harvested boreal black spruce forests in eastern Canada: effect on understory vegetation

Ecosystem-based management advocates that forestry disturbances should aim to emulate natural disturbances to mitigate the landscape-level impact of forest management. This study compares the impact of clear-cuts followed by a prescribed burn (CCPB) with clear-cuts alone (CC) and current careful logging practices (CLAAG: “careful logging around advanced growth”) on understory composition within black spruce (Picea mariana Miller (BSP)) paludified forest stands at the plot, site, and treatment levels using a functional-type approach. Vascular and nonvascular taxa showed significant differences in composition at the plot level among treatments. We found that pioneer taxa occurred mainly in CCPB sites, while late-successional taxa characterized CC sites. CLAAG sites had higher taxa richness than CCPB and CC sites, and we found that CCPB treatments were most likely to promote vascular taxa compositions that are more similar to those observed after natural disturbances. Additionally, the relative abundance of Sphagnum spp., responsible for paludification, was significantly reduced in sites treated by prescribed burning. This study therefore presents results suggesting that prescribed burning might represent a sustainable alternative to current harvesting techniques in the Clay Belt of eastern Canada that could help in preserving biodiversity (in terms of understory species assemblage) while maintaining or even enhancing forest productivity.

Effect of Organic Layer Thickness on Black Spruce Aging Mistakes in Canadian Boreal Forests

Boreal black spruce (Picea mariana) forests are prone to developing thick organic layers (paludification). Black spruce is adapted to this environment by the continuous development of adventitious roots, masking the root collar and making it difficult to age trees. Ring counts above the root collar underestimate age of trees, but the magnitude of age underestimation of trees in relation to organic layer thickness (OLT) is unknown. This age underestimation is required to produce appropriate age-correction tools to be used in land resource management. The goal of this study was to assess aging errors that are done with standard ring counts of trees growing in sites with different degrees of paludification (OLT; 0–25 cm, 26–65 cm, >65 cm). Age of 81 trees sampled at three geographical locations was determined by ring counts at ground level and at 1 m height, and real age of trees was determined by cross-dating growth rings down to the root collar (root/shoot interface). Ring counts at 1 m height underestimated age of trees by a mean of 22 years (range 13–49) and 52 years (range 14–112) in null to low vs. moderately to highly paludified stands, respectively. The percentage of aging-error explained by our linear model was relatively high (R2adj = 0.71) and showed that OLT class and age at 0-m could be used to predict total aging-error while neither DBH nor geographic location could. The resulting model has important implications for forest management to accurately estimate productivity of these forests.

Deadwood abundance in post-harvest and post-fire residual patches: An evaluation of patch temporal dynamics in black spruce boreal forest

In managed boreal forests, variable retention harvest is considered by forest managers as a means of mitigating harvest impacts on biodiversity. Variable retention harvest consists of maintaining within a cutblock structural attributes of the original forest stand in intact forest patches that could provide quality habitat (i.e., with large trees and deadwood) for many forest species during forest regeneration. However, retention patch modalities (size, shape, age of the forest) allowing both persistence and sustainable recruitment of deadwood over time remains unknown. The objective of this study is to evaluate the abundance of recent deadwood in post-harvest and post-fire residual patches and to compare their temporal dynamics in black spruce dominated stands located in northwestern Quebec. Abundance of the recent deadwood, estimated as the sum of recent standing deadwood volume and recently fallen deadwood volume was analyzed in 41 post-fire residual patches, and in 45 post-harvest retention patches of varying ages (i.e. exposure time to the disturbed matrix) and in 37 continuous black spruce forest stands (controls). This study shows that post-fire residual patches appear in general more durable than post-harvest retention patches after disturbance. In a management context, our results indicate that: (1) large island patches and large linear separators oriented to escape windthrow usually have deadwood recruitment dynamics similar to that of post-fire patches; (2) retention patches with an initial stand volume greater than 60m3/ha will generate more deadwood volume over time. This suggests that the selection of large retention patches in the shape of an island or a separator, with high volume (between 60 and 300m3/ha) should help increase the persistence of post-harvest retention patches in black spruce forest, and simultaneously ensure quality habitat for several forest species while the adjacent managed forest regenerates.

Negligible structural development and edge influence on the understorey at 16–17‐yr‐old clear‐cut edges in black spruce forest

AbstractQuestionsWhat is the distance of edge influence on the structure and understorey composition at 16–17‐yr‐old cut edges in black spruce boreal forest? How do these edges compare with more recent 2–5‐yr‐old cut edges in the same region?LocationNorthwestern Quebec, Canada.MethodsForest structure and understorey composition were sampled along transects perpendicular to ten 16–17‐yr‐old clear‐cut edges, and compared to published results from 2–5‐yr‐old cut edges. We used randomization tests to assess the magnitude and distance of edge influence, and to compare edge influence between different edge ages.ResultsBlack spruce forest next to the 16–17‐yr‐old cut edges was structurally and compositionally very similar to interior forest, with little edge influence from harvesting beyond 5 m into the forest. Edge influence on the understorey was weak (low magnitude) and not very extensive (short distance) at these edges, with no significant edge influence on the abundance of individual species. Logs peaked in abundance on the forest side of the edge, with values higher than in either adjacent ecosystem.ConclusionsOverall, 16–17‐yr‐old cut edges in black spruce forest showed little evidence of further structural change compared to the 2–5‐yr‐old cut edges. Structural development of these edges as well as regeneration of the disturbed areas also resulted in reduced edge influence on the understorey. Instead, clear‐cut edges in black spruce forest may experience more forest influence on the regenerating disturbed area.

Prescribed burning after clearcut limits paludification in black spruce boreal forest

Paludification, the accumulation over the mineral soil of poorly decomposed organic matter mainly originating from Sphagnum spp., transforms black spruce (Picea mariana) boreal forests into forested peatlands in the prolonged absence of fire, which diminishes forest productivity. High-severity wildfires reset this process by burning the soil organic layer (SOL) and reinitiating forest succession. In contrast, low severity wildfires impact mainly the soil surface and tree layer and do not significantly reduce SOL depth. In the Clay Belt region of eastern Canada, an area prone to paludification, the current forest harvest practice (careful logging around advanced growth [CLAAG]) removes trees but has little impact on the SOL and the understorey vegetation. This is thought to further promote paludification, which consequently reduces forest productivity. Conversely, clearcut (CC) disturbs the SOL and the understorey vegetation, and is thought to favor tree growth. Furthermore, prescribed burning after clearcut (CCPB) is used as a site preparation technique, but may also be used to control paludification as it can burn part of the organic soil layer. Using a retrospective approach, our study examines three hypotheses: compared to CLAAG, CC and CCPB: (1) have positive effects on soil conditions (e.g. decomposition level and pH), (2) reduce Sphagnum spp. and ericaceous shrub cover and (3) result in enhanced black spruce growth. We sampled 22 sites in which we measured SOL characteristics (e.g. depth, decomposition state), understorey vegetation cover and black spruce growth. Compared to CLAAG, CCPB resulted in increased soil decomposition level and higher pH. CCPB also reduced Sphagnum spp. cover but not ericaceous shrub cover. Black spruce growth rate was higher following CCPB than CC, and mean dominant tree height was marginally higher following CCPB than CLAAG and CC. Our results demonstrate that CCPB is beneficial to black spruce growth, presumably through its effects on forest understorey and SOL chemistry. While not similar to a high severity fire, prescribed burning after clearcut in paludified stands on the Clay Belt emulates some wildfire effects such as increasing soil pH. We suggest that unlike CLAAG, prescribed burning after clearcut can restore black spruce stand productivity and should be considered in the context of forest ecosystem management.

Decoupled stoichiometric, isotopic, and fungal responses of an ectomycorrhizal black spruce forest to nitrogen and phosphorus additions

Many northern forests are limited by nitrogen (N) availability, slight changes in which can have profound effects on ecosystem function and the activity of ectomycorrhizal (EcM) fungi. Increasing N and phosphorus (P) availability, an analog to accelerated soil organic matter decomposition in a warming climate, could decrease plant dependency on EcM fungi and increase plant productivity as a result of greater carbon use efficiency. However, the impact of altered N and P availability on the growth and activity of EcM fungi in boreal forests remains poorly understood despite recognition of their importance to host plant nutrition and soil carbon sequestration. To address such uncertainty we examined above and belowground ecosystem properties in a boreal black spruce forest following five years of factorial N and P additions. By combining detailed soil, fungal, and plant δ15N measurements with in situ metrics of fungal biomass, growth, and activity, we found both expected and unexpected patterns. Soil nitrate isotope values became 15N enriched in response to both N and P additions; fungal biomass was repressed by N yet both biomass and growth were stimulated by P; and, black spruce dependency on EcM derived N increased slightly when N and P were added alone yet significantly declined when added in combination. These findings contradict predictions that N fertilization would increase plant P demands and P fertilization would further exacerbate plant N demands. As a result, the prediction that EcM fungi predictably respond to plant N limitation was not supported. These findings highlight P as an under appreciated mediator of the activity of denitrifying bacteria, EcM fungi, and the dynamics of N cycles in boreal forests. Further, use of δ15N values from bulk soils, plants, and fungi to understand how EcM systems respond to changing nutrient availabilities will often require additional ecological information.

Early‐season fires in boreal black spruce forests produce pyrogenic carbon with low intrinsic recalcitrance

Pyrogenic carbon (PyC), a major by‐product of wildfires in boreal forests, plays several critical roles in soil biogeochemical processes. However, PyC properties, including its potential recalcitrance, may vary depending on its formation conditions. Our study aimed to characterize the chemical and physical properties of PyC formed under variable fire severity in Eastern Canada boreal forests; these latter represent an important fraction of fire‐affected circumboreal ecosystems. A total of 267 PyC samples, produced by early‐season wildfires in 2005–2007, were collected ≤5 years after fire from the forest floors of 14 black spruce sites distributed across Quebec, to cover the range of fire severity encountered in these forests. Early‐season fires occur frequently in Eastern Canada, and are predicted to increase in regional and global scenarios of future fire regimes associated with climate change. Selected PyC samples were analyzed using elemental analysis, solid‐state 13C nuclear magnetic resonance (NMR) spectroscopy, scanning electron microscopy, and surface area analysis. The NMR spectra of the PyC collected on low‐fire‐severity sites were dominated by peaks indicative of cellulose, while those for PyC from higher‐severity sites were dominated by a broad peak assigned to aromatic carbons. Atomic H/C and O/C ratios decreased with increasing fire severity. By comparing field samples to samples produced in the laboratory under controlled formation conditions, we were able to infer that the temperature of formation in the field was low (75°–250°C). In addition, for all PyC samples, the aromatic carbon : total carbon ratio was small, suggesting that PyC produced by early‐season fires in these boreal forests may be susceptible to relatively rapid degradation. Taken together, our data suggest that boreal PyC may not be as recalcitrant as previously assumed, and that its influence on soil biogeochemical processes may be short lived.