Forest composition change and biophysical climate feedbacks across boreal North America.

Deciduous tree cover is expected to increase in North American boreal forests with climate warming and wildfire. This shift in composition has the potential to generate biophysical cooling via increased land surface albedo. Here we use Landsat-derived maps of continuous tree canopy cover and deciduous fractional composition to assess albedo change over recent decades. We find, on average, a small net decrease in deciduous fraction from 2000 to 2015 across boreal North America and from 1992 to 2015 across Canada, despite extensive fire disturbance that locally increased deciduous vegetation. We further find near-neutral net biophysical change in radiative forcing associated with albedo when aggregated across the domain. Thus, while there have been widespread changes in forest composition over the past several decades, the net changes in composition and associated post-fire radiative forcing have not induced systematic negative feedbacks to climate warming over the spatial and temporal scope of our study.

Material Legacies and Environmental Constraints Underlie Fire Resilience of a Dominant Boreal Forest Type.

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. Supplementary Information: The online version contains supplementary material available at 10.1007/s10021-022-00772-7.

Impacts of pre-fire conifer density and wildfire severity on ecosystem structure and function at the forest-tundra ecotone.

Wildfire frequency and extent is increasing throughout the boreal forest-tundra ecotone as climate warms. Understanding the impacts of wildfire throughout this ecotone is required to make predictions of the rate and magnitude of changes in boreal-tundra landcover, its future flammability, and associated feedbacks to the global carbon (C) cycle and climate. We studied 48 sites spanning a gradient from tundra to low-density spruce stands that were burned in an extensive 2013 wildfire on the north slope of the Alaska Range in Denali National Park and Preserve, central Alaska. We assessed wildfire severity and C emissions, and determined the impacts of severity on understory vegetation composition, conifer tree recruitment, and active layer thickness (ALT). We also assessed conifer seed rain and used a seeding experiment to determine factors controlling post-fire tree regeneration. We found that an average of 2.18 ± 1.13 Kg C m-2 was emitted from this fire, almost 95% of which came from burning of the organic soil. On average, burn depth of the organic soil was 10.6 ± 4.5 cm and both burn depth and total C combusted increased with pre-fire conifer density. Sites with higher pre-fire conifer density were also located at warmer and drier landscape positions and associated with increased ALT post-fire, greater changes in pre- and post-fire understory vegetation communities, and higher post-fire boreal tree recruitment. Our seed rain observations and seeding experiment indicate that the recruitment potential of conifer trees is limited by seed availability in this forest-tundra ecotone. We conclude that the expected climate-induced forest infilling (i.e. increased density) at the forest-tundra ecotone could increase fire severity, but this infilling is unlikely to occur without increases in the availability of viable seed.

Increasing fire and the decline of fire adapted black spruce in the boreal forest.

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.

Carbon loss from boreal forest wildfires offset by increased dominance of deciduous trees.

In boreal forests, climate warming is shifting the wildfire disturbance regime to more frequent fires that burn more deeply into organic soils, releasing sequestered carbon to the atmosphere. To understand the destabilization of carbon storage, it is necessary to consider these effects in the context of long-term ecological change. In Alaskan boreal forests, we found that shifts in dominant plant species catalyzed by severe fire compensated for greater combustion of soil carbon over decadal time scales. Severe burning of organic soils shifted tree dominance from slow-growing black spruce to fast-growing deciduous broadleaf trees, resulting in a net increase in carbon storage by a factor of 5 over the disturbance cycle. Reduced fire activity in future deciduous-dominated boreal forests could increase the tenure of this carbon on the landscape, thereby mitigating the feedback to climate warming.

Wildfire combustion and carbon stocks in the southern Canadian boreal forest: Implications for a warming world.

Boreal wildfires are increasing in intensity, extent, and frequency, potentially intensifying carbon emissions and transitioning the region from a globally significant carbon sink to a source. The productive southern boreal forests of central Canada already experience relatively high frequencies of fire, and as such may serve as an analog of future carbon dynamics for more northern forests. Fire-carbon dynamics in southern boreal systems are relatively understudied, with limited investigation into the drivers of pre-fire carbon stocks or subsequent combustion. As part of NASA's Arctic-Boreal Vulnerability Experiment, we sampled 79 stands (47 burned, 32 unburned) throughout central Saskatchewan to characterize above- and belowground carbon stocks and combustion rates in relation to historical land use, vegetation characteristics, and geophysical attributes. We found southern boreal forests emitted an average of 3.3 ± 1.1 kg C/m2 from field sites. The emissions from southern boreal stands varied as a function of stand age, fire weather conditions, ecozone, and soil moisture class. Sites affected by historical timber harvesting had greater combustion rates due to faster carbon stock recovery rates than sites recovering from wildfire events, indicating that different boreal forest land use practices can generate divergent carbon legacy effects. We estimate the 2015 fire season in Saskatchewan emitted a total of 36.3 ± 15.0 Tg C, emphasizing the importance of southern boreal fires for regional carbon budgets. Using the southern boreal as an analog, the northern boreal may undergo fundamental shifts in forest structure and carbon dynamics, becoming dominated by stands <70 years old that hold 2-7 kg C/m2 less than current mature northern boreal stands. Our latitudinal approach reinforces previous studies showing that northern boreal stands are at a high risk of holding less carbon under changing disturbance conditions.

Climate change decreases the cooling effect from postfire albedo in boreal North America.

Fire is a primary disturbance in boreal forests and generates both positive and negative climate forcings. The influence of fire on surface albedo is a predominantly negative forcing in boreal forests, and one of the strongest overall, due to increased snow exposure in the winter and spring months. Albedo forcings are spatially and temporally heterogeneous and depend on a variety of factors related to soils, topography, climate, land cover/vegetation type, successional dynamics, time since fire, season, and fire severity. However, how these variables interact to influence albedo is not well understood, and quantifying these relationships and predicting postfire albedo becomes increasingly important as the climate changes and management frameworks evolve to consider climate impacts. Here we developed a MODIS-derived 'blue sky' albedo product and a novel machine learning modeling framework to predict fire-driven changes in albedo under historical and future climate scenarios across boreal North America. Converted to radiative forcing (RF), we estimated that fires generate an annual mean cooling of -1.77 ± 1.35 W/m2 from albedo under historical climate conditions (1971-2000) integrated over 70 years postfire. Increasing postfire albedo along a south-north climatic gradient was offset by a nearly opposite gradient in solar insolation, such that large-scale spatial patterns in RF were minimal. Our models suggest that climate change will lead to decreases in mean annual postfire albedo, and hence a decreasing strength of the negative RF, a trend dominated by decreased snow cover in spring months. Considering the range of future climate scenarios and model uncertainties, we estimate that for fires burning in the current era (2016) the cooling effect from long-term postfire albedo will be reduced by 15%-28% due to climate change.

Increasing wildfires threaten historic carbon sink of boreal forest soils.

Boreal forest fires emit large amounts of carbon into the atmosphere primarily through the combustion of soil organic matter1-3. 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 fires5-7, 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.

Wildfire severity reduces richness and alters composition of soil fungal communities in boreal forests of western Canada.

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.

Cross-scale controls on carbon emissions from boreal forest megafires.

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.

Impacts of fire on non-native plant recruitment in black spruce forests of interior Alaska.

Climate change is expected to increase the extent and severity of wildfires throughout the boreal forest. Historically, black spruce (Picea mariana (Mill.) B.S.P.) forests in interior Alaska have been relatively free of non-native species, but the compounding effects of climate change and an altered fire regime could facilitate the expansion of non-native plants. We tested the effects of wildfire on non-native plant colonization by conducting a seeding experiment of non-native plants on different substrate types in a burned black spruce forest, and surveying for non-native plants in recently burned and mature black spruce forests. We found few non-native plants in burned or mature forests, despite their high roadside presence, although invasion of some burned sites by dandelion (Taraxacum officinale) indicated the potential for non-native plants to move into burned forest. Experimental germination rates were significantly higher on mineral soil compared to organic soil, indicating that severe fires that combust much of the organic layer could increase the potential for non-native plant colonization. We conclude that fire disturbances that remove the organic layer could facilitate the invasion of non-native plants providing there is a viable seed source and dispersal vector.

Stable carbon isotope analysis reveals widespread drought stress in boreal black spruce forests.

Unprecedented rates of climate warming over the past century have resulted in increased forest stress and mortality worldwide. Decreased tree growth in association with increasing temperatures is generally accepted as a signal of temperature-induced drought stress. However, variations in tree growth alone do not reveal the physiological mechanisms behind recent changes in tree growth. Examining stable carbon isotope composition of tree rings in addition to tree growth can provide a secondary line of evidence for physiological drought stress. In this study, we examined patterns of black spruce growth and carbon isotopic composition in tree rings in response to climate warming and drying in the boreal forest of interior Alaska. We examined trees at three nested scales: landscape, toposequence, and a subsample of trees within the toposequence. At each scale, we studied the potential effects of differences in microclimate and moisture availability by sampling on northern and southern aspects. We found that black spruce radial growth responded negatively to monthly metrics of temperature at all examined scales, and we examined ∆(13)C responses on a subsample of trees as representative of the wider region. The negative ∆(13)C responses to temperature reveal that black spruce trees are experiencing moisture stress on both northern and southern aspects. Contrary to our expectations, ∆(13)C from trees on the northern aspect exhibited the strongest drought signal. Our results highlight the prominence of drought stress in the boreal forest of interior Alaska. We conclude that if temperatures continue to warm, we can expect drought-induced productivity declines across large regions of the boreal forest, even for trees located in cool and moist landscape positions.