Conifer forests historically have been resilient to wildfires in part due to thick organic soil layers that regulate combustion and post-fire moisture and vegetation change. However, recent shifts in fire activity in western North America may be overwhelming these resilience mechanisms with potential impacts for energy and carbon exchange. Here, we quantify the long-term recovery of the organic soil layer and its carbon pools across 511 forested plots. Our plots span ~ 140,000 km2 across two ecozones of the Northwest Territories, Canada, and allowed us to investigate the impacts of time-after-fire, site moisture class, and dominant canopy type on soil organic layer thickness and associated carbon stocks. Despite thinner soil organic layers in xeric plots immediately after fire, these drier stands supported faster post-fire recovery of the soil organic layer than in mesic plots. Unlike xeric or mesic stands, post-fire soil carbon accumulation rates in hydric plots were negligible despite wetter forested plots having greater soil organic carbon stocks immediately post-fire compared to other stands. While permafrost and high-water tables inhibit combustion and maintain thick organic soils immediately after fire, our results suggest that these wet stands are not recovering their pre-fire carbon stocks on a century timescale. We show that canopy conversion from black spruce to jack pine or deciduous dominance could reduce organic soil carbon stocks by 60–80% depending on stand age. Our two main findings—decreasing organic soil carbon storage with increasing deciduous cover and the lack of post-fire SOL recovery in hydric sites—have implications for the turnover time of carbon stocks in the western boreal forest region and also will impact energy fluxes by controlling albedo and surface soil moisture.

Abstract Key message Black spruce ( Picea mariana (Mill.) B.S.P.) has historically self-replaced following wildfire, but recent evidence suggests that this is changing. One factor could be negative impacts of intensifying fire activity on black spruce seed rain. We investigated this by measuring black spruce seed rain and seedling establishment. Our results suggest that increases in fire activity could reduce seed rain meaning reductions in black spruce establishment. Context Black spruce is an important conifer in boreal North America that develops a semi-serotinous, aerial seedbank and releases a pulse of seeds after fire. Variation in postfire seed rain has important consequences for black spruce regeneration and stand composition. Aims We explore the possible effects of changes in fire regime on the abundance and viability of black spruce seeds following a very large wildfire season in the Northwest Territories, Canada (NWT). Methods We measured postfire seed rain over 2 years at 25 black spruce-dominated sites and evaluated drivers of stand characteristics and environmental conditions on total black spruce seed rain and viability. Results We found a positive relationship between black spruce basal area and total seed rain. However, at high basal areas, this increasing rate of seed rain was not maintained. Viable seed rain was greater in stands that were older, closer to unburned edges, and where canopy combustion was less severe. Finally, we demonstrated positive relationships between seed rain and seedling establishment, confirming our measures of seed rain were key drivers of postfire forest regeneration. Conclusion These results indicate that projected increases in fire activity will reduce levels of black spruce recruitment following fire.

Abstract. Fire is the dominant disturbance agent in Alaskan and Canadian boreal ecosystems and releases large amounts of carbon into the atmosphere. Burned area and carbon emissions have been increasing with climate change, which have the potential to alter the carbon balance and shift the region from a historic sink to a source. It is therefore critically important to track the spatiotemporal changes in burned area and fire carbon emissions over time. Here we developed a new burned area detection algorithm between 2001–2019 across Alaska and Canada at 500 meters (m) resolution that utilizes finer-scale 30 m Landsat imagery to account for land cover unsuitable for burning. This method strictly balances omission and commission errors at 500 m to derive accurate landscape- and regional-scale burned area estimates. Using this new burned area product, we developed statistical models to predict burn depth and carbon combustion for the same period within the NASA Arctic-Boreal Vulnerability Experiment (ABoVE) core and extended domain. Statistical models were constrained using a database of field observations across the domain and were related to a variety of response variables including remotely-sensed indicators of fire severity, fire weather indices, local climate, soils, and topographic indicators. The burn depth and aboveground combustion models performed best, with poorer performance for belowground combustion. We estimate 2.37 million hectares (Mha) burned annually between 2001–2019 over the ABoVE domain (2.87 Mha across all of Alaska and Canada), emitting 79.3 +/- 27.96 (+/- 1 standard deviation) Teragrams of carbon (C) per year, with a mean combustion rate of 3.13 +/- 1.17 kilograms C m-2. Mean combustion and burn depth displayed a general gradient of higher severity in the northwestern portion of the domain to lower severity in the south and east. We also found larger fire years and later season burning were generally associated with greater mean combustion. Our estimates are generally consistent with previous efforts to quantify burned area, fire carbon emissions, and their drivers in regions within boreal North America; however, we generally estimate higher burned area and carbon emissions due to our use of Landsat imagery, greater availability of field observations, and improvements in modeling. The burned area and combustion data sets described here (the ABoVE Fire Emissions Database, or ABoVE-FED) can be used for local to continental-scale applications of boreal fire science.

Abstract Foundation species have disproportionately large impacts on ecosystem structure and function. As a result, future changes to their distribution may be important determinants of ecosystem carbon (C) cycling in a warmer world. We assessed the role of a foundation tussock sedge ( Eriophorum vaginatum ) as a climatically vulnerable C stock using field data, a machine learning ecological niche model, and an ensemble of terrestrial biosphere models (TBMs). Field data indicated that tussock density has decreased by ~0.97 tussocks per m2 over the past ~38 years on Alaska’s North Slope from ~1981 to 2019. This declining trend is concerning because tussocks are a large Arctic C stock, which enhances soil organic layer C stocks by 6.9% on average and represents 745 Tg C across our study area. By 2100, we project that changes in tussock density may decrease the tussock C stock by 41% in regions where tussocks are currently abundant (e.g. -0.8 tussocks per m2 and -85 Tg C on the North Slope) and may increase the tussock C stock by 46% in regions where tussocks are currently scarce (e.g. +0.9 tussocks per m2 and +81 Tg C on Victoria Island). These climate-induced changes to the tussock C stock were comparable to, but sometimes opposite in sign, to vegetation C stock changes predicted by an ensemble of TBMs. Our results illustrate the important role of tussocks as a foundation species in determining future Arctic C stocks and highlights the need for better representation of this species in TBMs.

Resilience of plant communities to disturbance is supported by multiple mechanisms, including ecological legacies affecting propagule availability, species' environmental tolerances, and biotic interactions. Understanding the relative importance of these mechanisms for plant community resilience supports predictions of where and how resilience will be altered with disturbance. We tested mechanisms underlying resilience of forests dominated by black spruce (Picea mariana) to fire disturbance across a heterogeneous forest landscape in the Northwest Territories, Canada. We combined surveys of naturally regenerating seedlings at 219 burned plots with experimental manipulations of ecological legacies via seed addition of four tree species and vertebrate exclosures to limit granivory and herbivory at 30 plots varying in moisture and fire severity. Black spruce recovery was greatest where it dominated pre-fire, at wet sites with deep residual soil organic layers, and fire conditions of low soil or canopy combustion and longer return intervals. Experimental addition of seed indicated all species were seed-limited, emphasizing the importance of propagule legacies. Black spruce and birch (Betula papyrifera) recruitment were enhanced with vertebrate exclusion. Our combination of observational and experimental studies demonstrates black spruce is vulnerable to effects of increased fire activity that erode ecological legacies. Moreover, black spruce relies on wet areas with deep soil organic layers where other species are less competitive. However, other species can colonize these areas if enough seed is available or soil moisture is altered by climate change. Testing mechanisms underlying species' resilience to disturbance aids predictions of where vegetation will transform with effects of climate change.The online version contains supplementary material available at 10.1007/s10021-022-00772-7.

In boreal North America, much of the landscape is covered by fire-adapted forests dominated by serotinous conifers. For these forests, reductions in fire return interval could limit reproductive success, owing to insufficient time for stands to reach reproductive maturity i.e., to initiate cone production. Improved understanding of the drivers of reproductive maturity can provide important information about the capacity of these forests to self-replace following fire. Here, we assessed the drivers of reproductive maturity in two dominant and widespread conifers, semi-serotinous black spruce and serotinous jack pine. Presence or absence of female cones were recorded in approximately 15,000 individuals within old and recently burned stands in two distinct ecozones of the Northwest Territories (NWT), Canada. Our results show that reproductive maturity was triggered by a minimum tree size threshold rather than an age threshold, with trees reaching reproductive maturity at smaller sizes where environmental conditions were more stressful. The number of reproductive trees per plot increased with stem density, basal area, and at higher latitudes (colder locations). The harsh climatic conditions present at these higher latitudes, however, limited the recruitment of jack pine at the treeline ecotone. The number of reproductive black spruce trees increased with deeper soils, whereas the number of reproductive jack pine trees increased where soils were shallower. We examined the reproductive efficiency i.e., the number of seedlings recruited per reproductive tree, linking pre-fire reproductive maturity of recently burned stands and post-fire seedling recruitment (recorded up to 4 years after the fires) and found that a reproductive jack pine can recruit on average three times more seedlings than a reproductive black spruce. We suggest that the higher reproductive efficiency of jack pine can explain the greater resilience of this species to wildfire compared with black spruce. Overall, these results help link life history characteristics, such as reproductive maturity, to variation in post-fire recruitment of dominant serotinous conifers.

Intensifying wildfire activity and climate change can drive rapid forest compositional shifts. In boreal North America, black spruce shapes forest flammability and depends on fire for regeneration. This relationship has helped black spruce maintain its dominance through much of the Holocene. However, with climate change and more frequent and severe fires, shifts away from black spruce dominance to broadleaf or pine species are emerging, with implications for ecosystem functions including carbon sequestration, water and energy fluxes, and wildlife habitat. Here, we predict that such reductions in black spruce after fire may already be widespread given current trends in climate and fire. To test this, we synthesize data from 1,538 field sites across boreal North America to evaluate compositional changes in tree species following 58 recent fires (1989 to 2014). While black spruce was resilient following most fires (62%), loss of resilience was common, and spruce regeneration failed completely in 18% of 1,140 black spruce sites. In contrast, postfire regeneration never failed in forests dominated by jack pine, which also possesses an aerial seed bank, or broad-leaved trees. More complete combustion of the soil organic layer, which often occurs in better-drained landscape positions and in dryer duff, promoted compositional changes throughout boreal North America. Forests in western North America, however, were more vulnerable to change due to greater long-term climate moisture deficits. While we find considerable remaining resilience in black spruce forests, predicted increases in climate moisture deficits and fire activity will erode this resilience, pushing the system toward a tipping point that has not been crossed in several thousand years.

Fungi play key roles in carbon (C) dynamics of ecosystems: saprotrophs decompose organic material and return C in the nutrient cycle, and mycorrhizal species support plants that accumulate C through photosynthesis. The identities and functions of extremophile fungi present after fire can influence C dynamics, particularly because plant-fungal relationships are often species-specific. However, little is known about the function and distribution of fungi that survive fires. We aim to assess the distribution of heat-resistant soil fungi across burned stands of boreal forest in the Northwest Territories, Canada, and understand their functions in relation to decomposition and tree seedling growth. We cultured and identified fungi from heat-treated soils and linked sequences from known taxa with high throughput sequencing fungal data (Illumina MiSeq, ITS1) from soils collected in 47 plots. We assessed functions under controlled conditions by inoculating litter and seedlings with heat-resistant fungi to assess decomposition and effects on seedling growth, respectively, for black spruce ( Picea mariana ), birch ( Betula papyrifera ), and jack pine ( Pinus banksiana ). We also measured litter decomposition rates and seedling densities in the field without inoculation. We isolated seven taxa of heat-resistant fungi and found their relative abundances were not associated with environmental or fire characteristics. Under controlled conditions, Fayodia gracilipes and Penicillium arenicola decomposed birch, but no taxa decomposed black spruce litter significantly more than the control treatment. Seedlings showed reduced biomass and/or mortality when inoculated with at least one of the fungal taxa. Penicillium turbatum reduced growth and/or caused mortality of all three species of seedlings. In the field, birch litter decomposed faster in stands with greater pre-fire proportion of black spruce, while black spruce litter decomposed faster in stands experiencing longer fire-free intervals. Densities of seedlings that had germinated since fire were positively associated with ectomycorrhizal richness while there were fewer conifer seedlings with greater heat-resistant fungal abundance. Overall, our study suggests that extremophile fungi present after fires have multiple functions and may have unexpected negative effects on forest functioning and regeneration. In particular, heat-resistant fungi after fires may promote shifts away from conifer dominance that are observed in these boreal forests.

Increases in fire frequency, extent, and severity are expected to strongly impact the structure and function of boreal forest ecosystems. An important function of the boreal forest is its ability to sequester and store carbon (C). Increasing disturbance from wildfires, emitting large amounts of C to the atmosphere, may create a positive feedback to climate warming. Variation in ecosystem structure and function throughout the boreal forest are important for predicting the effects of climate warming and changing fire regimes on C dynamics. In this study, we compiled data on soil characteristics, stand structure, pre-fire C pools, C loss from fire, and the potential drivers of these C metrics from 527 sites distributed across six ecoregions of North America’s western boreal forests. We assessed structural and functional differences between these fire-prone ecoregions using data from 417 recently burned sites (2004-2015) and estimated ecoregion-specific relationships between soil characteristics and depth from 167 of these sites plus an additional 110 sites (27 burned, 83 unburned). We found that northern boreal ecoregions were generally older, stored and emitted proportionally more belowground than aboveground C and exhibited lower rates of C accumulation over time than southern ecoregions. We present ecoregion specific estimates of depth-wise soil characteristics that are important for predicting C combustion from fire. As climate continues to warm and disturbance from wildfires increases, the C dynamics of these fire-prone ecoregions are likely to change with significant implications for the global C cycle and its feedbacks to climate change.

© 2020 The Authors. Ecography published by John Wiley & Sons Ltd on behalf of Nordic Society Oikos Climate change is altering disturbance regimes outside historical norms, which can impact biodiversity by selecting for plants with particular traits. The relative impact of disturbance characteristics on plant traits and community structure may be mediated by environmental gradients. We aimed to understand how wildfire impacted understory plant communities and plant regeneration strategies along gradients of environmental conditions and wildfire characteristics in boreal forests. We established 207 plots (60 m2) in recently burned stands and 133 plots in mature stands with no recent fire history in comparable gradients of stand type, site moisture (drainage) and soil organic layer (SOL) depth in two ecozones in Canada's Northwest Territories. At each plot, we recorded all vascular plant taxa in the understory and measured the regeneration strategy (seeder, resprouter, survivor) in burned plots, along with seedbed conditions (mineral soil and bryophyte cover). Dispersal, longevity and growth form traits were determined for each taxon. Fire characteristics measured included proportion of pre-fire SOL combusted (fire severity), date of burn (fire seasonality) and pre-fire stand age (time following fire). Results showed understory community composition was altered by fire. However, burned and mature stands had similar plant communities in wet sites with deep SOL. In the burned plots, regeneration strategies were determined by fire severity, drainage and pre- and post-fire SOL depth. Resprouters were more common in wet sites with deeper SOL and lower fire severity, while seeders were associated with drier sites with thinner SOL and greater fire severity. This led to drier burned stands being compositionally different from their mature counterparts and seedbed conditions were important. Our study highlights the importance of environment–wildfire interactions in shaping plant regeneration strategies and patterns of understory plant community structure across landscapes, and the overriding importance of SOL depth and site drainage in mediating fire severity, plant regeneration and community structure.

Carbon (C) emissions from wildfires are a key terrestrial–atmosphere interaction that influences global atmospheric composition and climate. Positive feedbacks between climate warming and boreal wildfires are predicted based on top-down controls of fire weather and climate, but C emissions from boreal fires may also depend on bottom-up controls of fuel availability related to edaphic controls and overstory tree composition. Here we synthesized data from 417 field sites spanning six ecoregions in the northwestern North American boreal forest and assessed the network of interactions among potential bottom-up and top-down drivers of C emissions. Our results indicate that C emissions are more strongly driven by fuel availability than by fire weather, highlighting the importance of fine-scale drainage conditions, overstory tree species composition and fuel accumulation rates for predicting total C emissions. By implication, climate change-induced modification of fuels needs to be considered for accurately predicting future C emissions from boreal wildfires.

Wildfire is the dominant disturbance in boreal forests and fire activity is increasing in these regions. Soil fungal communities are important for plant growth and nutrient cycling postfire but there is little understanding of how fires impact fungal communities across landscapes, fire severity gradients, and stand types in boreal forests. Understanding relationships between fungal community composition, particularly mycorrhizas, and understory plant composition is therefore important in predicting how future fire regimes may affect vegetation. We used an extreme wildfire event in boreal forests of Canada's Northwest Territories to test drivers of fungal communities and assess relationships with plant communities. We sampled soils from 39 plots 1 year after fire and 8 unburned plots. High-throughput sequencing (MiSeq, ITS) revealed 2,034 fungal operational taxonomic units. We found soil pH and fire severity (proportion soil organic layer combusted), and interactions between these drivers were important for fungal community structure (composition, richness, diversity, functional groups). Where fire severity was low, samples with low pH had higher total fungal, mycorrhizal, and saprotroph richness compared to where severity was high. Increased fire severity caused declines in richness of total fungi, mycorrhizas, and saprotrophs, and declines in diversity of total fungi and mycorrhizas. The importance of stand age (a surrogate for fire return interval) for fungal composition suggests we could detect long-term successional patterns even after fire. Mycorrhizal and plant community composition, richness, and diversity were weakly but significantly correlated. These weak relationships and the distribution of fungi across plots suggest that the underlying driver of fungal community structure is pH, which is modified by fire severity. This study shows the importance of edaphic factors in determining fungal community structure at large scales, but suggests these patterns are mediated by interactions between fire and forest stand composition.

Boreal forest fires emit large amounts of carbon into the atmosphere primarily through the combustion of soil organic matter1,2,3. During each fire, a portion of this soil beneath the burned layer can escape combustion, leading to a net accumulation of carbon in forests over multiple fire events4. Climate warming and drying has led to more severe and frequent forest fires5,6,7, which threaten to shift the carbon balance of the boreal ecosystem from net accumulation to net loss1, resulting in a positive climate feedback8. This feedback will occur if organic-soil carbon that escaped burning in previous fires, termed ‘legacy carbon’, combusts. Here we use soil radiocarbon dating to quantitatively assess legacy carbon loss in the 2014 wildfires in the Northwest Territories of Canada2. We found no evidence for the combustion of legacy carbon in forests that were older than the historic fire-return interval of northwestern boreal forests9. In forests that were in dry landscapes and less than 60 years old at the time of the fire, legacy carbon that had escaped burning in the previous fire cycle was combusted. We estimate that 0.34 million hectares of young forests (<60 years) that burned in the 2014 fires could have experienced legacy carbon combustion. This implies a shift to a domain of carbon cycling in which these forests become a net source—instead of a sink—of carbon to the atmosphere over consecutive fires. As boreal wildfires continue to increase in size, frequency and intensity7, the area of young forests that experience legacy carbon combustion will probably increase and have a key role in shifting the boreal carbon balance.

Climate warming and drying is associated with increased wildfire disturbance and the emergence of megafires in North American boreal forests. Changes to the fire regime are expected to strongly increase combustion emissions of carbon (C) which could alter regional C balance and positively feedback to climate warming. In order to accurately estimate C emissions and thereby better predict future climate feedbacks, there is a need to understand the major sources of heterogeneity that impact C emissions at different scales. Here, we examined 211 field plots in boreal forests dominated by black spruce (Picea mariana) or jack pine (Pinus banksiana) of the Northwest Territories (NWT), Canada after an unprecedentedly large area burned in 2014. We assessed both aboveground and soil organic layer (SOL) combustion, with the goal of determining the major drivers in total C emissions, as well as to develop a high spatial resolution model to scale emissions in a relatively understudied region of the boreal forest. On average, 3.35 kg C m−2 was combusted and almost 90% of this was from SOL combustion. Our results indicate that black spruce stands located at landscape positions with intermediate drainage contribute the most to C emissions. Indices associated with fire weather and date of burn did not impact emissions, which we attribute to the extreme fire weather over a short period of time. Using these results, we estimated a total of 94.3 Tg C emitted from 2.85 Mha of burned area across the entire 2014 NWT fire complex, which offsets almost 50% of mean annual net ecosystem production in terrestrial ecosystems of Canada. Our study also highlights the need for fine-scale estimates of burned area that represent small water bodies and regionally specific calibrations of combustion that account for spatial heterogeneity in order to accurately model emissions at the continental scale.

Increased fire frequency, extent and severity are expected to strongly affect the structure and function of boreal forest ecosystems. In this study, we examined 213 plots in boreal forests dominated by black spruce (Picea mariana) or jack pine (Pinus banksiana) of the Northwest Territories, Canada, after an unprecedentedly large area burned in 2014. Large fire size is associated with high fire intensity and severity, which would manifest as areas with deep burning of the soil organic layer (SOL). Our primary objectives were to estimate burn depth in these fires and then to characterise landscapes vulnerable to deep burning throughout this region. Here we quantify burn depth in black spruce stands using the position of adventitious roots within the soil column, and in jack pine stands using measurements of burned and unburned SOL depths. Using these estimates, we then evaluate how burn depth and the proportion of SOL combusted varies among forest type, ecozone, plot-level moisture and stand density. Our results suggest that most of the SOL was combusted in jack pine stands regardless of plot moisture class, but that black spruce forests experience complete combustion of the SOL only in dry and moderately well-drained landscape positions. The models and calibrations we present in this study should allow future research to more accurately estimate burn depth in Canadian boreal forests.

Northern ecosystem processes play out across scales that are rare elsewhere on contemporary earth: large ranging predator–prey systems are still operational, invasive species are rare, and large-scale natural disturbances occur extensively. Disturbances in the far north affect huge areas of land and are difficult to control or manage. Historically, disturbance patterns and processes ranging across a number of spatio-temporal scales have played an important role in the resilience of northern ecosystems. However, due to interactions with a warming climate, these disturbances are now erasing key legacies of the last millennia of ecosystem processes. Building on the concepts of legacies and cross-scale interactions, we highlight several general conceptual issues that represent key challenges for the future of northern ecosystem science, but that also have relevance to other biomes.