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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. [ABSTRACT FROM AUTHOR]

Transpiration is a globally important component of evapotranspiration. Careful upscaling of transpiration from point measurements is thus crucial for quantifying water and energy fluxes. In spatially heterogeneous landscapes common across the boreal biome, upscaled transpiration estimates are difficult to determine due to variation in local environmental conditions (e.g., basal area, soil moisture, permafrost). Here, we sought to determine stand‐level attributes that influence transpiration scalars for a forested boreal peatland complex consisting of sparsely treed wetlands and densely treed permafrost plateaus as land cover types. The objectives were to quantify spatial and temporal variability in stand‐level transpiration, and to identify sources of uncertainty when scaling point measurements to the stand‐level. Using heat ratio method sap flow sensors, we determined sap velocity for black spruce and tamarack for 2‐week periods during peak growing season in 2013, 2017 and 2018. We found greater basal area, drier soils, and the presence of permafrost increased daily sap velocity in individual trees, suggesting that local environmental conditions are important in dictating sap velocity. When sap velocity was scaled to stand‐level transpiration using gridded 20 × 20 m resolution data across the ~10 ha Scotty Creek ForestGEO plot, we observed significant differences in daily plot transpiration among years (0.17–0.30 mm), and across land cover types. Daily transpiration was lowest in grid‐cells with sparsely treed wetlands compared to grid‐cells with well‐drained and densely treed permafrost plateaus, where daily transpiration reached 0.80 mm, or 30% of the daily evapotranspiration. When transpiration scalars (i.e., sap velocity) were not specific to the different land cover types (i.e., permafrost plateaus and wetlands), scaled stand‐level transpiration was overestimated by 42%. To quantify the relative contribution of tree transpiration to ecosystem evapotranspiration, we recommend that sampling designs stratify across local environmental conditions to accurately represent variation associated with land cover types, especially with different hydrological functioning as encountered in rapidly thawing boreal peatland complexes. [ABSTRACT FROM AUTHOR]

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 FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Time series of vegetation indices derived from satellite imagery are useful in measuring vegetation response to climate warming in remote northern regions. These indices show that productivity is generally declining in the boreal forest, but it is unclear which components of boreal vegetation are driving these trends. We aimed to compare trends in the normalized difference vegetation index (NDVI) to forest growth and demographic data taken from a 10 ha mapped plot located in a spruce‐dominated boreal peatland. We used microcores to quantify recent growth trends and tree census data to characterize mortality and recruitment rates of the three dominant tree species. We then compared spatial patterns in growth and demography to patterns in Landsat‐derived maximum NDVI trends (1984–2019) in 78 pixels that fell within the plot. We found that NDVI trends were predominantly positive (i.e., "greening") in spite of the ongoing loss of black spruce (the dominant species; 80% of stems) from the plot. The magnitude of these trends correlated positively with black spruce growth trends, but was also governed to a large extent by tree mortality and recruitment. Greening trends were weaker (lower slope) in areas with high larch mortality, and high turnover of spruce and birch, but stronger (higher slope) in areas with high larch recruitment. Larch dominance is currently low (~11% of stems), but it is increasing in abundance as permafrost thaw progresses and will likely have a substantial influence on future NDVI trends. Our results emphasize that NDVI trends in boreal peatlands can be positive even when the forest as a whole is in decline, and that the magnitude of trends can be strongly influenced by the demographics of uncommon species. [ABSTRACT FROM AUTHOR]

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Copyright of Journal of Ecology is the property of Wiley-Blackwell and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

The acceleration of permafrost thaw due to warming, wetting, and disturbance is altering circumpolar landscapes. The effect of thaw is largely determined by ground ice content in near‐surface permafrost, making the characterization and prediction of ground ice content critical. Here we evaluate the spatial and stratigraphic variation of near‐surface ground ice characteristics in the dominant forest types in the North Slave region near Yellowknife, Northwest Territories, Canada. Physical variation in the permafrost was assessed through cryostructure, soil properties, and volumetric ice content, and relationships between these parameters were determined. Near‐surface ground ice characteristics were contrasted between forest types. In black spruce forests the top of the permafrost was ice‐rich and characterized by lenticular and ataxitic cryostructures, indicating the presence of an intermediate layer. Most white spruce/birch forests showed similar patterns; however, an increase in the active layer thickness and permafrost thaw at some sites have eradicated the transition zone, and the large ice lenses encountered at depth reflect segregated ground ice developed during initial downward aggradation of permafrost. Our findings indicate that white spruce/birch terrain will be less sensitive than black spruce forests to near‐surface permafrost thaw. However, if permafrost thaws completely, white spruce/birch terrain will probably be transformed into wetland–thaw lake complexes due to high ground ice content at depth. [ABSTRACT FROM AUTHOR]

Abstract: 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. [ABSTRACT FROM AUTHOR]

Water stress has been identified as a key mechanism of the contemporary increase in tree mortality rates in northwestern North America. However, a detailed analysis of boreal tree hydrodynamics and their interspecific differences is still lacking. Here we examine the hydraulic behaviour of co-occurring larch (Larix laricina) and black spruce (Picea mariana), two characteristic boreal tree species, near the southern limit of the boreal ecozone in central Canada. Sap flux density (Js), concurrently recorded stem radius fluctuations and meteorological conditions are used to quantify tree hydraulic functioning and to scrutinize tree water-use strategies. Our analysis revealed asynchrony in the diel hydrodynamics of the two species with the initial rise in Js occurring 2 h earlier in larch than in black spruce. Interspecific differences in larch and black spruce crown architecture explained the observed asynchrony in their hydraulic functioning. Furthermore, the two species exhibited diverging stomatal regulation strategies with larch and black spruce employing relatively isohydric and anisohydric behaviour, respectively. Such asynchronous and diverging tree-level hydrodynamics provide new insights into the ecosystem-level complementarity in tree form and function, with implications for understanding boreal forests' water and carbon dynamics and their resilience to environmental stress. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

• We examined the controls of growing season (GS GPP) in boreal forests. • Temperature and green-up explain the start of GS GPP in permafrost-free stands. • The selected predictors cannot explain the start of GS GPP in permafrost stands. • Soil water content and continuous snowpack formation explain the end of GS GPP. 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 (SOS GPP). Results from commonality analysis and structural equation modeling suggest that canopy 'green-up' and air temperature directly affected SOS GPP in permafrost-free forest stands. In addition, soil temperature acted as mediator for an indirect effect of air temperature on SOS GPP. In contrast, none of the environmental variables, or their combination, explained the variation in SOS GPP in forest stands with permafrost. The explanatory power of environmental variables was more consistent regarding the end-of-season (EOS GPP). In both, forest stands with and without permafrost, EOS GPP 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 SOS GPP and EOS GPP in forest stands with and without permafrost will help to refine parameterizations of the boreal biome in Earth system models. [ABSTRACT FROM AUTHOR]

• We reconstructed time series of boreal tree growth with a biometric approach. • Aboveground tree growth was a minor and decoupled fraction of carbon input. • Partitioned estimates of tree carbon sink are valuable observational constraints. • Such observational constraints can be used for model validation and policy making. The boreal biome accounts for approximately one third of the terrestrial carbon (C) sink. However, estimates of its individual C pools remain uncertain. Here, focusing on the southern boreal forest, we quantified the magnitude and temporal dynamics of C allocation to aboveground tree growth at a mature black spruce (Picea mariana)-dominated forest stand in Saskatchewan, Canada. We reconstructed aboveground tree biomass increments (AGBi) using a biometric approach, i.e., species-specific allometry combined with forest stand characteristics and tree ring widths collected with a C-oriented sampling design. We explored the links between boreal tree growth and ecosystem C input by comparing AGBi with eddy-covariance-derived ecosystem C fluxes from 1999 to 2015 and we synthesized our findings with a refined meta-analysis of published values of boreal forest C use efficiency (CUE). Mean AGBi at the study site was decoupled from ecosystem C input and equal to 71 ± 7 g C m–2 (1999–2015), which is only a minor fraction of gross ecosystem production (GEP; i.e., AGBi / GEP ≈ 9 %). Moreover, C allocation to AGBi remained stable over time (AGBi / GEP; –0.0001 yr–1; p -value=0.775), contrary to significant trends in GEP (+5.72 g C m–2 yr–2; p -value=0.02) and CUE (–0.0041 yr–1, p -value=0.007). CUE was estimated as 0.50 ± 0.03 at the study area and 0.41 ± 0.12 across the reviewed boreal forests. These findings highlight the importance of belowground tree C investments, together with the substantial contribution of understory, ground cover and soil to the boreal forest C balance. Our quantitative insights into the dynamics of aboveground boreal tree C allocation offer additional observational constraints for terrestrial ecosystem models that are often biased in converting C input to biomass, and can guide forest-management strategies for mitigating carbon dioxide emissions. [ABSTRACT FROM AUTHOR]