Jennifer L. Baltzer

2024

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Low-severity fires in the boreal region: reproductive implications for black spruce stands in between stand-replacing fire events
Raquel Alfaro‐Sánchez, Jill F. Johnstone, Jennifer L. Baltzer
Annals of Botany

Stand-replacing crown fires are the most prevalent type of fire regime in boreal forests in North America. However, a substantial proportion of low-severity fires are found within fire perimeters. Here we aimed to investigate the effects of low-severity fires on the reproductive potential and seedling recruitment in boreal forests stands in between stand-replacing fire events.

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Response of Boreal Plant Communities and Forest Floor Carbon Fluxes to Experimental Nutrient Additions
Katherine M. Standen, Anastasia E. Sniderhan, Oliver Sonnentag, Carolina Voigt, Jennifer L. Baltzer
Ecosystems, Volume 27, Issue 3

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Radiation, Air Temperature, and Soil Water Availability Drive Tree Water Deficit Across Temporal Scales in Canada's Western Boreal Forest
Nia Perron, Jennifer L. Baltzer, Matteo Detto, Magali F. Nehemy, Christopher Spence, Gabriel Hould‐Gosselin, Haley Alcock, Bram Hadiwijaya, Colin P. Laroque, Oliver Sonnentag
Geophysical Research Letters, Volume 51, Issue 8

Abstract Changes are projected for the boreal biome with complex and variable effects on forest vegetation including drought‐induced tree mortality and forest loss. With soil and atmospheric conditions governing drought intensity, specific drivers of trees water stress can be difficult to disentangle across temporal scales. We used wavelet analysis and causality detection to identify potential environmental controls (evapotranspiration, soil moisture, rainfall, vapor pressure deficit, air temperature and photosynthetically active radiation) on daily tree water deficit and on longer periods of tree dehydration in black spruce and tamarack. Daily tree water deficit was controlled by photosynthetically active radiation, vapor pressure deficit, and air temperature, causing greater stand evapotranspiration. Prolonged periods of tree water deficit (multi‐day) were regulated by photosynthetically active radiation and soil moisture. We provide empirical evidence that continued warming and drying will cause short‐term increases in black spruce and tamarack transpiration, but greater drought stress with reduced soil water availability.

2023

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Post-fire Recovery of Soil Organic Layer Carbon in Canadian Boreal Forests
Kristen Bill, Catherine M. Dieleman, Jennifer L. Baltzer, Geneviève É. Degré‐Timmons, Michelle C. Mack, Nicola J. Day, Steve G. Cumming, Xanthe J. Walker, M. R. Turetsky, Kristen Bill, Catherine M. Dieleman, Jennifer L. Baltzer, Geneviève É. Degré‐Timmons, Michelle C. Mack, Nicola J. Day, Steve G. Cumming, Xanthe J. Walker, M. R. Turetsky
Ecosystems

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.

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The influence of postfire recovery and environmental conditions on boreal vegetation
Alexis Gardiner Jorgensen, Raquel Alfaro‐Sánchez, Steven G. Cumming, Alison L. White, Geneviève É. Degré‐Timmons, Nicola J. Day, M. R. Turetsky, Jill F. Johnstone, Xanthe J. Walker, Jennifer L. Baltzer, Alexis Gardiner Jorgensen, Raquel Alfaro‐Sánchez, Steven G. Cumming, Alison L. White, Geneviève É. Degré‐Timmons, Nicola J. Day, M. R. Turetsky, Jill F. Johnstone, Xanthe J. Walker, Jennifer L. Baltzer
Ecosphere, Volume 14, Issue 7

Abstract Climate change is increasing the frequency and extent of fires in the boreal biome of North America. These changes can alter the recovery of both canopy and understory vegetation. There is uncertainty about plant and lichen recovery patterns following fire, and how they are mediated by environmental conditions. Here, we aim to address these knowledge gaps by studying patterns of postfire vegetation recovery at the community and individual species level over the first 100+ years following fire. Data from vegetation surveys collected from 581 plots in the Northwest Territories, Canada, ranging from 1 to 275 years postfire, were used to assess the influence of time after fire and local environmental conditions on plant community composition and to model trends in the relative abundance of several common plant and lichen species. Time after fire significantly influenced vegetation community composition and interacted with local environmental conditions, particularly soil moisture. Soil moisture individually (in the absence of interactions) was the most commonly significant variable in plant and lichen recovery models. Patterns of postfire recovery varied greatly among species. Our results provide novel information on plant community recovery after fire and highlight the importance of soil moisture to local vegetation patterns. They will aid northern communities and land managers to anticipate the impacts of increased fire activity on both local vegetation and the wildlife that relies on it.

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Spatial and temporal variation in forest transpiration across a forested boreal peatland complex
Nia Perron, Jennifer L. Baltzer, Oliver Sonnentag, Nia Perron, Jennifer L. Baltzer, Oliver Sonnentag
Hydrological Processes, Volume 37, Issue 2

Abstract 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.

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Black spruce (Picea mariana) seed availability and viability in boreal forests after large wildfires
Kirsten A. Reid, Nicola J. Day, Raquel Alfaro‐Sánchez, Jill F. Johnstone, Steven G. Cumming, Michelle C. Mack, M. R. Turetsky, Xanthe J. Walker, Jennifer L. Baltzer, Kirsten A. Reid, Nicola J. Day, Raquel Alfaro‐Sánchez, Jill F. Johnstone, Steven G. Cumming, Michelle C. Mack, M. R. Turetsky, Xanthe J. Walker, Jennifer L. Baltzer
Annals of Forest Science, Volume 80, Issue 1

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.

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Permafrost thaw induces short‐term increase in vegetation productivity in northwestern Canada
Emily L. Ogden, Steven G. Cumming, Sharon L. Smith, M. R. Turetsky, Jennifer L. Baltzer, Emily L. Ogden, Steven G. Cumming, Sharon L. Smith, M. R. Turetsky, Jennifer L. Baltzer
Global Change Biology, Volume 29, Issue 18

Abstract Over the past several decades, various trends in vegetation productivity, from increases to decreases, have been observed throughout Arctic–Boreal ecosystems. While some of this variation can be explained by recent climate warming and increased disturbance, very little is known about the impacts of permafrost thaw on productivity across diverse vegetation communities. Active layer thickness data from 135 permafrost monitoring sites along a 10° latitudinal transect of the Northwest Territories, Canada, paired with a Landsat time series of normalized difference vegetation index from 1984 to 2019, were used to quantify the impacts of changing permafrost conditions on vegetation productivity. We found that active layer thickness contributed to the observed variation in vegetation productivity in recent decades in the northwestern Arctic–Boreal, with the highest rates of greening occurring at sites where the near‐surface permafrost recently had thawed. However, the greening associated with permafrost thaw was not sustained after prolonged periods of thaw and appeared to diminish after the thaw front extended outside the plants' rooting zone. Highest rates of greening were found at the mid‐transect sites, between 62.4° N and 65.2° N, suggesting that more southernly sites may have already surpassed the period of beneficial permafrost thaw, while more northern sites may have yet to reach a level of thaw that supports enhanced vegetation productivity. These results indicate that the response of vegetation productivity to permafrost thaw is highly dependent on the extent of active layer thickening and that increases in productivity may not continue in the coming decades.

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The Northwest Territories Thermokarst Mapping Collective: A northern-driven mapping collaborative toward understanding the effects of permafrost thaw
Steven V. Kokelj, Tristan Gingras‐Hill, Seamus V. Daly, P D Morse, S A Wolfe, Ashley Rudy, Jurjen van der Sluijs, Niels Weiss, H B O'Neill, Jennifer L. Baltzer, Trevor C. Lantz, Carolyn Gibson, Dieter Cazon, Robert Fraser, Duane G. Froese, Garfield Giff, Charles Klengenberg, Scott F. Lamoureux, W. L. Quinton, M. R. Turetsky, Alexandre Chiasson, C.C. Ferguson, Mike Newton, Mike Pope, Jason Paul, A E Wilson, Joseph M. Young, Steven V. Kokelj, Tristan Gingras‐Hill, Seamus V. Daly, P D Morse, S A Wolfe, Ashley Rudy, Jurjen van der Sluijs, Niels Weiss, H B O'Neill, Jennifer L. Baltzer, Trevor C. Lantz, Carolyn Gibson, Dieter Cazon, Robert Fraser, Duane G. Froese, Garfield Giff, Charles Klengenberg, Scott F. Lamoureux, W. L. Quinton, M. R. Turetsky, Alexandre Chiasson, C.C. Ferguson, Mike Newton, Mike Pope, Jason Paul, A E Wilson, Joseph M. Young, Steven V. Kokelj, Tristan Gingras‐Hill, Seamus V. Daly, P D Morse, S A Wolfe, Ashley Rudy, Jurjen van der Sluijs, Niels Weiss, H B O'Neill, Jennifer L. Baltzer, Trevor C. Lantz, Carolyn Gibson, Dieter Cazon, Robert Fraser, Duane G. Froese, Garfield Giff, Charles Klengenberg, Scott F. Lamoureux, W. L. Quinton, M. R. Turetsky, Alexandre Chiasson, C.C. Ferguson, Mike Newton, Mike Pope, Jason Paul, A E Wilson, Joseph M. Young, Steven V. Kokelj, Tristan Gingras‐Hill, Seamus V. Daly, P D Morse, S A Wolfe, Ashley Rudy, Jurjen van der Sluijs, Niels Weiss, H B O'Neill, Jennifer L. Baltzer, Trevor C. Lantz, Carolyn Gibson, Dieter Cazon, Robert Fraser, Duane G. Froese, Garfield Giff, Charles Klengenberg, Scott F. Lamoureux, W. L. Quinton, M. R. Turetsky, Alexandre Chiasson, C.C. Ferguson, Mike Newton, Mike Pope, Jason Paul, A E Wilson, Joseph M. Young
Arctic Science

This paper documents the first comprehensive inventory of thermokarst and thaw-sensitive terrain indicators for a 2 million km2 region of northwestern Canada. This is accomplished through the Thermokarst Mapping Collective (TMC), a research collaborative to systematically inventory indicators of permafrost thaw sensitivity by mapping and aerial assessments across the Northwest Territories (NT), Canada. The increase in NT-based permafrost capacity has fostered science leadership and collaboration with government, academic, and community researchers to enable project implementation. Ongoing communications and outreach have informed study design and strengthened Indigenous and stakeholder relationships. Documentation of theme-based methods supported mapper training, and flexible data infrastructure facilitated progress by Canada-wide researchers throughout the COVID-19 pandemic. The TMC inventory of thermokarst and thaw-sensitive landforms agree well with fine-scale empirical mapping (69% to 84% accuracy) and aerial inventory (74% to 96% accuracy) datasets. National- and circumpolar-scale modelling of sensitive permafrost terrain contrasts significantly with TMC outputs, highlighting their limitations and the value of empirically-based mapping approaches. We demonstrate that the multi-parameter TMC outputs support a holistic understanding and refined depictions of permafrost terrain sensitivity, provide novel opportunities for syntheses, and inform future modelling approaches, which are urgently required to comprehend better what permafrost thaw means for Canada’s North.

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Reviews and syntheses: Recent advances in microwave remote sensing in support of terrestrial carbon cycle science in Arctic–boreal regions
Alex Mavrovic, Oliver Sonnentag, Juha Lemmetyinen, Jennifer L. Baltzer, Christophe Kinnard, Alexandre Roy, Alex Mavrovic, Oliver Sonnentag, Juha Lemmetyinen, Jennifer L. Baltzer, Christophe Kinnard, Alexandre Roy
Biogeosciences, Volume 20, Issue 14

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Burned area and carbon emissions across northwestern boreal North America from 2001–2019
Stefano Potter, Sol Cooperdock, Sander Veraverbeke, Xanthe J. Walker, Michelle C. Mack, S. J. Goetz, Jennifer L. Baltzer, Laura Bourgeau‐Chavez, Arden Burrell, Catherine M. Dieleman, Nancy H. F. French, Stijn Hantson, Elizabeth Hoy, Liza K. Jenkins, Jill F. Johnstone, Evan S. Kane, Susan M. Natali, James T. Randerson, M. R. Turetsky, Ellen Whitman, Elizabeth B. Wiggins, Brendan M. Rogers
Biogeosciences, Volume 20, Issue 13

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 m (meters) 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 2.37×106 ha (2.37 Mha) burned annually between 2001–2019 over the ABoVE domain (2.87 Mha across all of Alaska and Canada), emitting 79.3 ± 27.96 Tg (±1 standard deviation) of carbon (C) per year, with a mean combustion rate of 3.13 ± 1.17 kg 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 datasets described here (the ABoVE Fire Emissions Database, or ABoVE-FED) can be used for local- to continental-scale applications of boreal fire science.

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Drivers of legacy soil organic matter decomposition after fire in boreal forests
Brian Izbicki, Xanthe J. Walker, Jennifer L. Baltzer, Nicola J. Day, Christopher Ebert, Jill F. Johnstone, Elaine Pegoraro, Edward A. G. Schuur, M. R. Turetsky, Michelle C. Mack
Ecosphere, Volume 14, Issue 11

Abstract Boreal forests harbor as much carbon (C) as the atmosphere and significant amounts of organic nitrogen (N), the nutrient most likely to limit plant productivity in high‐latitude ecosystems. In the boreal biome, the primary disturbance is wildfire, which consumes plant biomass and soil material, emits greenhouse gasses, and influences long‐term C and N cycling. Climate warming and drying is increasing wildfire severity and frequency and is combusting more soil organic matter (SOM). Combustion of surface SOM exposes deeper older layers of accumulated soil material that previously escaped combustion during past fires, here termed legacy SOM. Postfire SOM decomposition and nutrient availability are determined by these layers, but the drivers of legacy SOM decomposition are unknown. We collected soils from plots after the largest fire year on record in the Northwest Territories, Canada, in 2014. We used radiocarbon dating to measure Δ 14 C (soil age index), soil extractions to quantify N pools and microbial biomass, and a 90‐day laboratory incubation to measure the potential rate of element mineralization and understand patterns and drivers of legacy SOM C decomposition and N availability. We discovered that bulk soil C age predicted C decomposition, where cumulatively, older soil (approximately −450.0‰) produced 230% less C during the incubation than younger soil (~0.0‰). Soil age also predicted C turnover times, with old soil turnover 10 times slower than young soil. We found respired C was younger than bulk soil C, indicating most C enters and leaves relatively quickly, while the older portion remains a stable C sink. Soil age and other indices were unrelated to N availability, but microbial biomass influenced N availability, with more microbial biomass immobilizing soil N pools. Our results stress the importance of legacy SOM as a stable C sink and highlight that soil age drives the pace and magnitude of soil C contributions to the atmosphere between wildfires.

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Greater variation in boreal plant community composition and community‐level traits at local‐ than regional‐scale
Katherine M. Standen, Jennifer L. Baltzer
Journal of Vegetation Science, Volume 34, Issue 5

Abstract Questions Rapid climate change in northern latitudes is expected to influence plant functional traits of the whole community (community‐level traits) through species compositional changes and/or trait plasticity, limiting our ability to anticipate climate warming impacts on northern plant communities. We explored differences in plant community composition and community‐level traits within and among four boreal peatland sites and determined whether intra‐ or interspecific variation drives community‐level traits. Location Boreal biome of western North America. Methods We collected plant community composition and functional trait data along dominant topoedaphic and/or hydrologic gradients at four peatland sites spanning the latitudinal extent of the boreal biome of western North America. We characterized variability in community composition and community‐level traits of understorey vascular and moss species both within (local‐scale) and among sites (regional‐scale). Results Against expectations, community‐level traits of vascular plant and moss species were generally consistent among sites. Furthermore, interspecific variation was more important in explaining community‐level trait variation than intraspecific variation. Within‐site variation in both community‐level traits and community composition was greater than among‐site variation, suggesting that local environmental gradients (canopy density, organic layer thickness, etc.) may be more influential in determining plant community processes than regional‐scale gradients. Conclusions Given the importance of interspecific variation to within‐site shifts in community‐level traits and greater variation of community composition within than among sites, we conclude that climate‐induced shifts in understorey community composition may not have a strong influence on community‐level traits in boreal peatlands unless local‐scale environmental gradients are substantially altered.

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Evidence for unexpected net permafrost aggradation driven by local hydrology and climatic triggers
Anastasia E. Sniderhan, Christopher Spence, Steven V. Kokelj, Jennifer L. Baltzer
Environmental Research Letters, Volume 18, Issue 11

Abstract Rapid rates of high latitude warming over the past century have led to widespread research on permafrost thaw and its consequences. Studies from lowland plains environments in the discontinuous permafrost zone have highlighted extensive areal loss of permafrost, largely through observations of the collapse of forested permafrost plateaus into wetland features. These low-relief environments tend to have poor drainage, which initiates runaway thaw as increased soil moisture amplifies permafrost degradation. In contrast to lowland plains, the Taiga Shield landscape features a network of lakes, wetlands, soil-filled lowlands, and forests interspersed with bedrock outcrops. With the exposed (or near-surface) bedrock in this landscape, this region may have greater terrain stability under a warming climate than the lowland plains. The hydrological complexity of the Taiga Shield may also contribute to more varied trajectories for permafrost in this landscape. We investigated land cover change and implications for permafrost in an area that typifies the Taiga Shield. We took intensive ground-based measurements of soil organic layer (SOL) thickness and frost table depth to characterize different land cover types. Archival aerial photographs and recent satellite imagery from the area allowed us to assess land cover change between 1972 and 2017. Associations between permafrost, SOL, and land cover allowed us to use land cover as a proxy for change in permafrost extent. Our results suggest that both aggradation and degradation of permafrost has occurred within the Taiga Shield landscape over this 45 year period, but interestingly we found evidence for a net increase in permafrost extent. Permafrost aggradation in this landscape seems to be driven by a combination of local hydrology and climatic triggers that lead to colder, drier soil conditions that are favourable for the development of permafrost. This study highlights the importance of considering diverse and heterogenous landscapes in the study of changing permafrost ecosystems.

2022

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Burned Area and Carbon Emissions Across Northwestern Boreal North America from 2001–2019
Stefano Potter, Sol Cooperdock, Sander Veraverbeke, Xanthe J. Walker, Michelle C. Mack, S. J. Goetz, Jennifer L. Baltzer, Laura Bourgeau‐Chavez, Arden Burrell, Catherine M. Dieleman, Nancy H. F. French, Stijn Hantson, Elizabeth Hoy, Liza K. Jenkins, Jill F. Johnstone, Evan S. Kane, Susan M. Natali, James T. Randerson, M. R. Turetsky, Ellen Whitman, Elizabeth B. Wiggins, Brendan M. Rogers

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.

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Understanding among-lake variability of mercury concentrations in Northern Pike (Esox lucius): A whole-ecosystem study in subarctic lakes
Mehdi Moslemi-Aqdam, Leanne F. Baker, Jennifer L. Baltzer, Brian A. Branfireun, Marlene S. Evans, Brian Laird, George C. Low, Mike Low, Heidi K. Swanson
Science of The Total Environment, Volume 822

Mercury concentrations ([Hg]) in fish reflect complex biogeochemical and ecological interactions that occur at a range of spatial and biological scales. Elucidating these interactions is crucial to understanding and predicting fish [Hg], particularly at northern latitudes, where environmental perturbations are having profound effects on land-water-animal interactions, and where fish are a critical subsistence food source. Using data from eleven subarctic lakes that span an area of ~60,000 km2 in the Dehcho Region of Northwest Territories (Canada), we investigated how trophic ecology and growth rates of fish, lake water chemistry, and catchment characteristics interact to affect [Hg] in Northern Pike (Esox lucius), a predatory fish of widespread subsistence and commercial importance. Results from linear regression and piecewise structural equation models showed that 83% of among-lake variability in Northern Pike [Hg] was explained by fish growth rates (negative) and concentrations of methyl Hg ([MeHg]) in benthic invertebrates (positive). These variables were in turn influenced by concentrations of dissolved organic carbon, MeHg (water), and total Hg (sediment) in lakes, which were ultimately driven by catchment characteristics. Lakes in relatively larger catchments and with more temperate/subpolar needleleaf and mixed forests had higher [Hg] in Northern Pike. Our results provide a plausible mechanistic understanding of how interacting processes at scales ranging from whole catchments to individual organisms influence fish [Hg], and give insight into factors that could be considered for prioritizing lakes for monitoring in subarctic regions.

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Snowmelt Water Use at Transpiration Onset: Phenology, Isotope Tracing, and Tree Water Transit Time
Magali F. Nehemy, Jason Maillet, Nia Perron, Christoforos Pappas, Oliver Sonnentag, Jennifer L. Baltzer, Colin P. Laroque, Jeffrey J. McDonnell
Water Resources Research, Volume 58, Issue 9

Studies of tree water source partitioning have primarily focused on the growing season. However, little is yet known about the source of transpiration before, during, and after snowmelt when trees rehydrate and recommence transpiration in the spring. This study investigates tree water use during spring snowmelt following tree's winter stem shrinkage. We document the source of transpiration of three boreal forest tree species—Pinus banksiana, Picea mariana, and Larix laricina—by combining observations of weekly isotopic signatures (δ18O and δ2H) of xylem, soil water, rainfall and snowmelt with measurements of soil moisture dynamics, snow depth and high-resolution temporal measurements of stem radius changes and sap flow. Our data shows that the onset of stem rehydration and transpiration overlaps with snowmelt for evergreens. During rehydration and transpiration onset, xylem water at the canopy reflected a constant pre-melt isotopic signature likely showing late fall conditions. As snowmelt infiltrates the soil and recharges the soil matrix, soil water shows a rapid isotopic shift to depleted-snowmelt water values. While there was an overlap between snowmelt and transpiration timing, xylem and soil water isotopic values did not overlap during transpiration onset. Our data showed 1–2-week delay in the shift in xylem water from pre-melt to clear snowmelt-depleted water signatures in evergreen species. This delay appears to be controlled by tree water transit time that was in the order of 9–18 days. Our study shows that snowmelt is a key source for stem rehydration and transpiration in the boreal forest during spring onset.

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Scale‐dependent responses of understory vegetation to the physical structure of undisturbed tundra shrub patches
Cory A. Wallace, Jennifer L. Baltzer
Ecosphere, Volume 13, Issue 9

Much of the Arctic is experiencing rapid change in the productivity and recruitment of tall, deciduous shrubs. It is well established that shrub expansion can alter tundra ecosystem composition and function; however, less is known about the degree to which variability in the physical structure of shrub patches might mediate these changes. There is also limited information as to how different physical attributes of shrub patches may covary and how they differ with topography. Here, we address these knowledge gaps by measuring the physical structure, abiotic conditions, and understory plant community composition at sampling plots within undisturbed green alder patches at a taiga–tundra ecotone site in the Northwest Territories, Canada. We found surprisingly few associations between most structural variables and abiotic conditions at the plot scale, with the notable exceptions of canopy complexity and snow depth. Importantly, neither patch structure nor abiotic conditions were associated with the vegetation community at the plot scale when among-patch variation was accounted for. However, among-patch variation in plant community composition was significant and represented a gradient in the richness of tundra specialists and Sphagnum moss abundance. This gradient was strongly associated with mean patch snow depth, which was likely controlled at least in part by mean patch canopy complexity. Overall, natural variability in green alder patch structure had less of an association with abiotic conditions than expected, suggesting future changes in physical structure at undisturbed sites may have limited environmental impact at the plot scale. However, at the patch scale, increases in snow depth, likely related to canopy complexity, were negatively associated with tundra specialist richness, potentially due to phenological limitations associated with shortened growing seasons. In summary, our data suggest emergent properties exist at the patch scale that are not apparent at the plot scale such that plot-scale measurements do not represent variation in understory community composition across the landscape. The results presented here will inform future work addressing spatial variability in shrub impacts on ecosystem function and increase our understanding of understory community variation within alder patch habitats at the taiga–tundra ecotone.

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Material Legacies and Environmental Constraints Underlie Fire Resilience of a Dominant Boreal Forest Type
Nicola J. Day, Jill F. Johnstone, Kirsten A. Reid, Steven G. Cumming, Michelle C. Mack, M. R. Turetsky, Xanthe J. Walker, Jennifer L. Baltzer
Ecosystems, Volume 26, Issue 3

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.

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What explains the year-to-year variation in growing season timing of boreal black spruce forests?
Mariam El-Amine, Alexandre Roy, Franziska Koebsch, Jennifer L. Baltzer, Alan Barr, T. Andrew Black, Hiroki Ikawa, Hiroki Iwata, Hideki Kobayashi, Masahito Ueyama, Oliver Sonnentag
Agricultural and Forest Meteorology, Volume 324

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.

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Carbon and nitrogen cycling dynamics following permafrost thaw in the Northwest Territories, Canada
Catherine M. Dieleman, Nicola J. Day, Jean Holloway, Jennifer L. Baltzer, Thomas A. Douglas, M. R. Turetsky
Science of The Total Environment, Volume 845

Rapid climate warming across northern high latitudes is leading to permafrost thaw and ecosystem carbon release while simultaneously impacting other biogeochemical cycles including nitrogen. We used a two-year laboratory incubation study to quantify concomitant changes in carbon and nitrogen pool quantity and quality as drivers of potential CO2 production in thawed permafrost soils from eight soil cores collected across the southern Northwest Territories (NWT), Canada. These data were contextualized via in situ annual thaw depth measurements from 2015 to 2019 at 40 study sites that varied in burn history. We found with increasing time since experimental thaw the dissolved carbon and nitrogen pool quality significantly declined, indicating sustained microbial processing and selective immobilization across both pools. Piecewise structural equation modeling revealed CO2 trends were predominantly predicted by initial soil carbon content with minimal influence of dissolved phase carbon. Using these results, we provide a first-order estimate of potential near-surface permafrost soil losses of up to 80 g C m−2 over one year in southern NWT, exceeding regional historic mean primary productivity rates in some areas. Taken together, this research provides mechanistic knowledge needed to further constrain the permafrost‑carbon feedback and parameterize Earth system models, while building on empirical evidence that permafrost soils are at high risk of becoming weaker carbon sinks or even significant carbon sources under a changing climate.

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What Drives Reproductive Maturity and Efficiency in Serotinous Boreal Conifers?
Raquel Alfaro‐Sánchez, Jill F. Johnstone, Steve G. Cumming, Nicola J. Day, Michelle C. Mack, Xanthe J. Walker, Jennifer L. Baltzer
Frontiers in Ecology and Evolution, Volume 10

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.

2021

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Joint effects of climate, tree size, and year on annual tree growth derived from tree‐ring records of ten globally distributed forests
Kristina J. Anderson‐Teixeira, Valentine Herrmann, Christine R. Rollinson, Bianca Gonzalez, Erika Gonzalez‐Akre, Neil Pederson, M. Ross Alexander, Craig D. Allen, Raquel Alfaro‐Sánchez, Tala Awada, Jennifer L. Baltzer, Patrick J. Baker, Joseph D. Birch, Sarayudh Bunyavejchewin, Paolo Cherubini, Stuart J. Davies, Cameron Dow, Ryan Helcoski, Jakub Kašpar, James A. Lutz, Ellis Q. Margolis, Justin T. Maxwell, Sean M. McMahon, Camille Piponiot, Sabrina E. Russo, Pavel Šamonil, Anastasia E. Sniderhan, Alan J. Tepley, Ivana Vašíčková, Mart Vlam, Pieter A. Zuidema, Kristina J. Anderson‐Teixeira, Valentine Herrmann, Christine R. Rollinson, Bianca Gonzalez, Erika Gonzalez‐Akre, Neil Pederson, M. Ross Alexander, Craig D. Allen, Raquel Alfaro‐Sánchez, Tala Awada, Jennifer L. Baltzer, Patrick J. Baker, Joseph D. Birch, Sarayudh Bunyavejchewin, Paolo Cherubini, Stuart J. Davies, Cameron Dow, Ryan Helcoski, Jakub Kašpar, James A. Lutz, Ellis Q. Margolis, Justin T. Maxwell, Sean M. McMahon, Camille Piponiot, Sabrina E. Russo, Pavel Šamonil, Anastasia E. Sniderhan, Alan J. Tepley, Ivana Vašíčková, Mart Vlam, Pieter A. Zuidema
Global Change Biology, Volume 28, Issue 1

Tree rings provide an invaluable long-term record for understanding how climate and other drivers shape tree growth and forest productivity. However, conventional tree-ring analysis methods were not designed to simultaneously test effects of climate, tree size, and other drivers on individual growth. This has limited the potential to test ecologically relevant hypotheses on tree growth sensitivity to environmental drivers and their interactions with tree size. Here, we develop and apply a new method to simultaneously model nonlinear effects of primary climate drivers, reconstructed tree diameter at breast height (DBH), and calendar year in generalized least squares models that account for the temporal autocorrelation inherent to each individual tree's growth. We analyze data from 3811 trees representing 40 species at 10 globally distributed sites, showing that precipitation, temperature, DBH, and calendar year have additively, and often interactively, influenced annual growth over the past 120 years. Growth responses were predominantly positive to precipitation (usually over ≥3-month seasonal windows) and negative to temperature (usually maximum temperature, over ≤3-month seasonal windows), with concave-down responses in 63% of relationships. Climate sensitivity commonly varied with DBH (45% of cases tested), with larger trees usually more sensitive. Trends in ring width at small DBH were linked to the light environment under which trees established, but basal area or biomass increments consistently reached maxima at intermediate DBH. Accounting for climate and DBH, growth rate declined over time for 92% of species in secondary or disturbed stands, whereas growth trends were mixed in older forests. These trends were largely attributable to stand dynamics as cohorts and stands age, which remain challenging to disentangle from global change drivers. By providing a parsimonious approach for characterizing multiple interacting drivers of tree growth, our method reveals a more complete picture of the factors influencing growth than has previously been possible.

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Increasing fire and the decline of fire adapted black spruce in the boreal forest
Jennifer L. Baltzer, Nicola J. Day, Xanthe J. Walker, David F. Greene, Michelle C. Mack, Heather D. Alexander, Dominique Arseneault, Jennifer L. Barnes, Yves Bergeron, Yan Boucher, Laura Bourgeau‐Chavez, Carissa D. Brown, Suzanne Carrière, Brian K. Howard, Sylvie Gauthier, Marc‐André Parisien, Kirsten A. Reid, Brendan M. Rogers, Carl A. Roland, Luc Sirois, Sarah E. Stehn, Dan K. Thompson, M. R. Turetsky, Sander Veraverbeke, Ellen Whitman, Jian Yang, Jill F. Johnstone, Jennifer L. Baltzer, Nicola J. Day, Xanthe J. Walker, David F. Greene, Michelle C. Mack, Heather D. Alexander, Dominique Arseneault, Jennifer L. Barnes, Yves Bergeron, Yan Boucher, Laura Bourgeau‐Chavez, Carissa D. Brown, Suzanne Carrière, Brian K. Howard, Sylvie Gauthier, Marc‐André Parisien, Kirsten A. Reid, Brendan M. Rogers, Carl A. Roland, Luc Sirois, Sarah E. Stehn, Dan K. Thompson, M. R. Turetsky, Sander Veraverbeke, Ellen Whitman, Jian Yang, Jill F. Johnstone
Proceedings of the National Academy of Sciences, Volume 118, Issue 45

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.

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Seasonal thaw and landscape position determine foliar functional traits and whole‐plant water use in tall shrubs on the low arctic tundra
Katherine L. Black, Cory A. Wallace, Jennifer L. Baltzer, Katherine L. Black, Cory A. Wallace, Jennifer L. Baltzer
New Phytologist, Volume 231, Issue 1

Climate warming is driving tundra shrub expansion with implications for ecosystem function and regional climate. Understanding associations between shrub ecophysiological function, distribution and environment is necessary for predicting consequences of expansion. We evaluated the role of topographic gradients on upland shrub productivity to understand potential constraints on shrub expansion. At a low arctic tundra site near Inuvik, Northwest Territories, Canada, we measured sap flow, stem water potential and productivity-related functional traits in green alder, and environmental predictors (water and nutrient availability and seasonal thaw depth) across a toposequence in alder patches. Seasonal thaw reduced stem sap flow whereas topographic position predicted stem water potential and productivity-related functional traits. Upslope shrubs were more water-limited than those downslope. Shrubs in drainage channels had traits associated with greater productivity than those on the tops of slopes. The effect of thaw depth on sap flow has implications for seasonal water-use patterns and warming impacts on tundra ecohydrology. Topographic variation in functional traits corresponds with observed spatial patterns of tundra shrub expansion along floodplains and concave hillslopes rather than in upland areas. Green alder is expanding rapidly across the low arctic tundra in northwestern North America; thus, anticipating the implications of its expansion is essential for predicting tundra function.

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Unexpected greening in a boreal permafrost peatland undergoing forest loss is partially attributable to tree species turnover
Katherine D. Dearborn, Jennifer L. Baltzer, Katherine D. Dearborn, Jennifer L. Baltzer
Global Change Biology, Volume 27, Issue 12

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.

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Mapping and understanding the vulnerability of northern peatlands to permafrost thaw at scales relevant to community adaptation planning
Carolyn Gibson, Karl Cottenie, Tristan Gingras‐Hill, Steve V. Kokelj, Jennifer L. Baltzer, L. Chasmer, M. R. Turetsky, Carolyn Gibson, Karl Cottenie, Tristan Gingras‐Hill, Steve V. Kokelj, Jennifer L. Baltzer, L. Chasmer, M. R. Turetsky
Environmental Research Letters, Volume 16, Issue 5

Abstract Developing spatially explicit permafrost datasets and climate assessments at scales relevant to northern communities is increasingly important as land users and decision makers incorporate changing permafrost conditions in community and adaptation planning. This need is particularly strong within the discontinuous permafrost zone of the Northwest Territories (NWT) Canada where permafrost peatlands are undergoing rapid thaw due to a warming climate. Current data products for predicting landscapes at risk of thaw are generally built at circumpolar scales and do not lend themselves well to fine-scale regional interpretations. Here, we present a new permafrost vulnerability dataset that assesses the degree of permafrost thaw within peatlands across a 750 km latitudinal gradient in the NWT. This updated dataset provides spatially explicit estimates of where peatland thermokarst potential exists, thus making it much more suitable for local, regional or community usage. Within southern peatland complexes, we show that permafrost thaw affects up to 70% of the peatland area and that thaw is strongly mediated by both latitude and elevation, with widespread thaw occuring particularly at low elevations. At the northern end of our latitudinal gradient, peatland permafrost remains climate-protected with relatively little thaw. Collectively these results demonstrate the importance of scale in permafrost analyses and mapping if research is to support northern communities and decision makers in a changing climate. This study offers a more scale-appropriate approach to support community adaptative planning under scenarios of continued warming and widespread permafrost thaw.

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Predicting patterns of terrestrial lichen biomass recovery following boreal wildfires
Ruth J. Greuel, Geneviève É. Degré‐Timmons, Jennifer L. Baltzer, Jill F. Johnstone, Eliot J. B. McIntire, Nicola J. Day, Sarah J. Hart, Philip D. McLoughlin, Fiona K. A. Schmiegelow, M. R. Turetsky, Alexandre Truchon‐Savard, Mario D. van Telgen, Steven G. Cumming, Ruth J. Greuel, Geneviève É. Degré‐Timmons, Jennifer L. Baltzer, Jill F. Johnstone, Eliot J. B. McIntire, Nicola J. Day, Sarah J. Hart, Philip D. McLoughlin, Fiona K. A. Schmiegelow, M. R. Turetsky, Alexandre Truchon‐Savard, Mario D. van Telgen, Steven G. Cumming
Ecosphere, Volume 12, Issue 4

Increased fire activity due to climate change may impact the successional dynamics of boreal forests, with important consequences for caribou habitat. Early successional forests have been shown to support lower quantities of caribou forage lichens, but geographic variation in, and controls on, the rates of lichen recovery has been largely unexplored. In this study, we sampled across a broad region in northwestern Canada to compare lichen biomass accumulation in ecoprovinces, including the Saskatchewan Boreal Shield, the Northwest Territories Taiga Shield, and Northwest Territories Taiga Plains, divided into North and South. We focused on the most valuable Cladonia species for boreal and barren-ground caribou: Cladonia mitis and C. arbuscula, C. rangiferina and C. stygia, and C. stellaris and C. uncialis. We developed new allometric equations to estimate lichen biomass from field measurements of lichen cover and height; allometries were consistent among ecoprovinces, suggesting generalizability. We then used estimates of lichen biomass to quantify patterns of lichen recovery in different stand types, ecoprovinces, and with time following stand-replacing fire. We used a hurdle model to account both for the heterogeneous nature of lichen presence (zero inflation) and for the range of abundance in stands where lichen was present. The first component of the hurdle model, a generalized linear model, identified stand age, stand type, and ecoprovince as significant predictors of lichen presence. With a logistic growth model, a measure of lichen recovery (time to 50% asymptotic value) varied from 28 to 73 yr, dependent on stand type and ecoprovince. The combined predictions of the hurdle model suggest the most rapid recovery of lichen biomass across our study region occurred in jack pine in the Boreal Shield (30 yr), while stands located in the Taiga Plains (North and South) required a longer recovery period (approximately 75 yr). These results provide a basis for estimating future caribou habitat that encompasses some of the large variation in fire effects on lichen abundance and vegetation types across the range of boreal and barren-ground caribou in North America.

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Permafrost condition determines plant community composition and community‐level foliar functional traits in a boreal peatland
Katherine M. Standen, Jennifer L. Baltzer, Katherine M. Standen, Jennifer L. Baltzer
Ecology and Evolution, Volume 11, Issue 15

Boreal peatlands are critical ecosystems globally because they house 30%–40% of terrestrial carbon (C), much of which is stored in permafrost soil vulnerable to climate warming-induced thaw. Permafrost thaw leads to thickening of the active (seasonally thawed) layer and alters nutrient and light availability. These physical changes may influence community-level plant functional traits through intraspecific trait variation and/or species turnover. As permafrost thaw is expected to cause an efflux of carbon dioxide (CO2) and methane (CH4) from the soil to the atmosphere, it is important to understand thaw-induced changes in plant community productivity to evaluate whether these changes may offset some of the anticipated increases in C emissions. To this end, we collected vascular plant community composition and foliar functional trait data along gradients in aboveground tree biomass and active layer thickness (ALT) in a rapidly thawing boreal peatland, with the expectation that changes in above- and belowground conditions are indicative of altered resource availability. We aimed to determine whether community-level traits vary across these gradients, and whether these changes are dominated by intraspecific trait variation, species turnover, or both. Our results highlight that variability in community-level traits was largely attributable to species turnover and that both community composition and traits were predominantly driven by ALT. Specifically, thicker active layers associated with permafrost-free peatlands (i.e., bogs and fens) shifted community composition from slower-growing evergreen shrubs to faster-growing graminoids and forbs with a corresponding shift toward more productive trait values. The results from this rapidly thawing peatland suggest that continued warming-induced permafrost thaw and thermokarst development alter plant community composition and community-level traits and thus ecosystem productivity. Increased productivity may help to mitigate anticipated CO2 efflux from thawing permafrost, at least in the short term, though this response may be swamped by increase CH4 release.

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Non-uniform growth dynamics of a dominant boreal tree species (Picea mariana) in the face of rapid climate change
Anastasia E. Sniderhan, Steven D. Mamet, Jennifer L. Baltzer, Anastasia E. Sniderhan, Steven D. Mamet, Jennifer L. Baltzer
Canadian Journal of Forest Research, Volume 51, Issue 4

Northwestern Canada’s boreal forest has experienced rapid warming, drying, and changes to permafrost, yet the growth responses and mechanisms driving productivity have been under-studied at broad scales. Forest responses are largely driven by black spruce (Picea mariana (Mill.) B.S.P.) — the region’s most widespread and dominant tree. We collected tree ring samples from four black spruce-dominated sites across 15° of latitude, spanning gradients in climate and permafrost. We investigated (i) differences in growth patterns, (ii) variations in climatic drivers of growth, and (iii) trends in water use efficiency (WUE) through 13 C isotope analysis from 1945 to 2006. We found positive growth trends at all sites except those at mid-latitude, where rapid permafrost thaw drove declines. Annual growth was lowest at the tree limit site and highest at the tree line. Climatic drivers of these growth patterns varied; positive growth responses at the northerly sites were associated with warmer winters, whereas Δ 13 C trends and climate-growth responses at mid-latitude sites indicated that growth was limited by moisture availability. Δ 13 C signatures indicated increased WUE at the southernmost site, with no significant trends at northern sites. These results suggest that warming will increase the growth of trees at the northern extent of black spruce, but southerly areas may face drought stress if precipitation does not balance evapotranspiration.

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Summary and synthesis of Changing Cold Regions Network (CCRN) research in the interior of western Canada – Part 2: Future change in cryosphere, vegetation, and hydrology
C. M. DeBeer, H. S. Wheater, John W. Pomeroy, Alan Barr, Jennifer L. Baltzer, Jill F. Johnstone, M. R. Turetsky, Ronald E. Stewart, Masaki Hayashi, Garth van der Kamp, Shawn J. Marshall, Elizabeth M. Campbell, Philip Marsh, Sean K. Carey, W. L. Quinton, Yanping Li, Saman Razavi, Aaron Berg, Jeffrey J. McDonnell, Christopher Spence, Warren Helgason, Andrew Ireson, T. Andrew Black, Mohamed Elshamy, Fuad Yassin, Bruce Davison, Allan Howard, Julie M. Thériault, Kevin Shook, Michael N. Demuth, Alain Pietroniro, C. M. DeBeer, H. S. Wheater, John W. Pomeroy, Alan Barr, Jennifer L. Baltzer, Jill F. Johnstone, M. R. Turetsky, Ronald E. Stewart, Masaki Hayashi, Garth van der Kamp, Shawn J. Marshall, Elizabeth M. Campbell, Philip Marsh, Sean K. Carey, W. L. Quinton, Yanping Li, Saman Razavi, Aaron Berg, Jeffrey J. McDonnell, Christopher Spence, Warren Helgason, Andrew Ireson, T. Andrew Black, Mohamed Elshamy, Fuad Yassin, Bruce Davison, Allan Howard, Julie M. Thériault, Kevin Shook, Michael N. Demuth, Alain Pietroniro
Hydrology and Earth System Sciences, Volume 25, Issue 4

Abstract. The interior of western Canada, like many similar cold mid- to high-latitude regions worldwide, is undergoing extensive and rapid climate and environmental change, which may accelerate in the coming decades. Understanding and predicting changes in coupled climate–land–hydrological systems are crucial to society yet limited by lack of understanding of changes in cold-region process responses and interactions, along with their representation in most current-generation land-surface and hydrological models. It is essential to consider the underlying processes and base predictive models on the proper physics, especially under conditions of non-stationarity where the past is no longer a reliable guide to the future and system trajectories can be unexpected. These challenges were forefront in the recently completed Changing Cold Regions Network (CCRN), which assembled and focused a wide range of multi-disciplinary expertise to improve the understanding, diagnosis, and prediction of change over the cold interior of western Canada. CCRN advanced knowledge of fundamental cold-region ecological and hydrological processes through observation and experimentation across a network of highly instrumented research basins and other sites. Significant efforts were made to improve the functionality and process representation, based on this improved understanding, within the fine-scale Cold Regions Hydrological Modelling (CRHM) platform and the large-scale Modélisation Environmentale Communautaire (MEC) – Surface and Hydrology (MESH) model. These models were, and continue to be, applied under past and projected future climates and under current and expected future land and vegetation cover configurations to diagnose historical change and predict possible future hydrological responses. This second of two articles synthesizes the nature and understanding of cold-region processes and Earth system responses to future climate, as advanced by CCRN. These include changing precipitation and moisture feedbacks to the atmosphere; altered snow regimes, changing balance of snowfall and rainfall, and glacier loss; vegetation responses to climate and the loss of ecosystem resilience to wildfire and disturbance; thawing permafrost and its influence on landscapes and hydrology; groundwater storage and cycling and its connections to surface water; and stream and river discharge as influenced by the various drivers of hydrological change. Collective insights, expert elicitation, and model application are used to provide a synthesis of this change over the CCRN region for the late 21st century.

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allodb : An R package for biomass estimation at globally distributed extratropical forest plots
Erika Gonzalez‐Akre, Camille Piponiot, Mauro Lepore, Valentine Herrmann, James A. Lutz, Jennifer L. Baltzer, Christopher W. Dick, Gregory S. Gilbert, Fangliang He, Michael Heym, Alejandra I. Huerta, Patrick A. Jansen, Daniel J. Johnson, Nikolai Knapp, Kamil Král, Dunmei Lin, Yadvinder Malhi, Sean M. McMahon, Jonathan A. Myers, David A. Orwig, Diego I. Rodríguez‐Hernández, Sabrina E. Russo, Jessica Shue, Xugao Wang, Amy Wolf, Yang Tong-hui, Stuart J. Davies, Kristina J. Anderson‐Teixeira, Erika Gonzalez‐Akre, Camille Piponiot, Mauro Lepore, Valentine Herrmann, James A. Lutz, Jennifer L. Baltzer, Christopher W. Dick, Gregory S. Gilbert, Fangliang He, Michael Heym, Alejandra I. Huerta, Patrick A. Jansen, Daniel J. Johnson, Nikolai Knapp, Kamil Král, Dunmei Lin, Yadvinder Malhi, Sean M. McMahon, Jonathan A. Myers, David A. Orwig, Diego I. Rodríguez‐Hernández, Sabrina E. Russo, Jessica Shue, Xugao Wang, Amy Wolf, Yang Tong-hui, Stuart J. Davies, Kristina J. Anderson‐Teixeira
Methods in Ecology and Evolution, Volume 13, Issue 2

Allometric equations for calculation of tree above-ground biomass (AGB) form the basis for estimates of forest carbon storage and exchange with the atmosphere. While standard models exist to calculate forest biomass across the tropics, we lack a standardized tool for computing AGB across boreal and temperate regions that comprise the global extratropics. Here we present an integrated R package, allodb, containing systematically selected published allometric equations and proposed functions to compute AGB. The data component of the package is based on 701 woody species identified at 24 large Forest Global Earth Observatory (ForestGEO) forest dynamics plots representing a wide diversity of extratropical forests. A total of 570 parsed allometric equations to estimate individual tree biomass were retrieved, checked and combined using a weighting function designed to ensure optimal equation selection over the full tree size range with smooth transitions across equations. The equation dataset can be customized with built-in functions that subset the original dataset and add new equations. Although equations were curated based on a limited set of forest communities and number of species, this resource is appropriate for large portions of the global extratropics and can easily be expanded to cover novel forest types.

2020

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Permafrost thaw in boreal peatlands is rapidly altering forest community composition
Katherine D. Dearborn, Cory A. Wallace, Rajit Patankar, Jennifer L. Baltzer
Journal of Ecology, Volume 109, Issue 3

Boreal peatlands are frequently underlain by permafrost, which is thawing rapidly. A common ecological response to thaw is the conversion of raised forested plateaus to treeless wetlands, but unexplained spatial variation in responses, combined with a lack of stand‐level data, make it difficult to predict future trajectories of boreal forest composition and structure. We sought to characterize patterns and identify drivers of forest structure, composition, mortality and recruitment in a boreal peatland experiencing permafrost thaw. To do this, we established a large (10 ha) permanent forest plot (completed in 2014), located in the Northwest Territories, Canada, that includes 40,584 mapped and measured trees. In 2018, we conducted a comprehensive mortality and recruitment recensus. We also measured frost table depth, soil moisture, soil humification and organic layer thickness within the plot between 2012 and 2018, and used habitat association tests to link these variables to forest characteristics and dynamics. Forest composition and structure varied markedly throughout the plot and were strongly governed by patterns in permafrost presence and organic layer thickness. Overall, there was a net loss of trees from the plot at a rate of 0.7% year−1. Mortality of black spruce, the dominant tree species, was more than double that of recruitment and was strongly associated with permafrost thaw. In contrast, recruitment of larch was over four times greater than mortality, and occurred primarily in low‐lying, permafrost‐free wetlands with mineral soil near the surface. Synthesis. The trends in tree demography and underlying drivers suggest that spruce‐dominated permafrost plateaus will be converted into larch‐dominated wetlands as permafrost thaw progresses in boreal peatlands, particularly in areas where mineral soil is near the surface. In the longer term, thaw could increase the hydrologic connectivity of the landscape, resulting in widespread drainage and re‐vegetation by spruce, but we did not find evidence that this is occurring yet. Given the increasing rates of permafrost thaw, and positive feedbacks between thaw and forest change, we predict that larch abundance will continue to increase in boreal peatlands over the coming decades, leading to shifts in ecosystem function, wildlife habitat, albedo and snow dynamics.

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No beating around the bush: the impact of projected high‐latitude vegetation transitions on soil and ecosystem respiration
Jennifer L. Baltzer, Oliver Sonnentag
New Phytologist, Volume 227, Issue 6

Globally, ecosystem respiration of carbon dioxide (CO2) is the second largest terrestrial carbon (C) flux after photosynthesis (Mahecha et al., 2010). Soil respiration is the main contributor to ecosystem respiration (e.g. c. 70% in temperate forests; reviewed in Ryan & Law, 2005). Plants shunt tremendous quantities of newly photosynthesized C belowground for storage in their roots but also to support rootmetabolism, root exudate production, and resource trading with root symbionts, most notably mycorrhizas (Raich & Nadelhoffer, 1989). These latter C end-points result in newly-fixed C being respired by roots or their symbionts or becoming substrate for use by free-living soil microorganisms. The respiration of this new photosynthetic C can occur within a few days to a month or two after fixation and can contribute to > 50% of the soil respiration (H€ogberg et al., 2001). Plants allocate photosynthetic C differentially aboveground and belowground depending on resource limitation and the demands of the mutualists with whom they collaborate, suggesting that this contribution to soil respiration may vary. As such, both belowground and aboveground vegetation composition, structure, function, and mutualistic partnerships are quite important for determining soil and thus ecosystem respiration. A new paper by Parker et al. (2020; pp. 1818–1830), in this issue of New Phytologist advances our understanding of the contributions of canopy-forming species to soil respiration at the boreal forest–tundra ecotone (FTE), the world’s largest vegetation transition zone spanning rapidly warming high-latitude regions.

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Identifying Functional Impacts of Heat-Resistant Fungi on Boreal Forest Recovery After Wildfire
Nicola J. Day, Steven G. Cumming, Kari E. Dunfield, Jill F. Johnstone, Michelle C. Mack, Kirsten A. Reid, M. R. Turetsky, Xanthe J. Walker, Jennifer L. Baltzer
Frontiers in Forests and Global Change, Volume 3

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.

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Patterns of Ecosystem Structure and Wildfire Carbon Combustion Across Six Ecoregions of the North American Boreal Forest
Xanthe J. Walker, Jennifer L. Baltzer, Laura Bourgeau‐Chavez, Nicola J. Day, Catherine M. Dieleman, Jill F. Johnstone, Evan S. Kane, Brendan M. Rogers, M. R. Turetsky, Sander Veraverbeke, Michelle C. Mack
Frontiers in Forests and Global Change, Volume 3

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.

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Fire characteristics and environmental conditions shape plant communities via regeneration strategy
Nicola J. Day, Alison L. White, Jill F. Johnstone, Geneviève É. Degré‐Timmons, Steven G. Cumming, Michelle C. Mack, M. R. Turetsky, Xanthe J. Walker, Jennifer L. Baltzer
Ecography, Volume 43, Issue 10

© 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.

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Impact of wildfire on permafrost landscapes: A review of recent advances and future prospects
Jean Holloway, Antoni G. Lewkowicz, Thomas A. Douglas, Xiaoying Li, M. R. Turetsky, Jennifer L. Baltzer, Huijun Jin
Permafrost and Periglacial Processes, Volume 31, Issue 3

Changes in the frequency and extent of wildfires are expected to lead to substantial and irreversible alterations to permafrost landscapes under a warming climate. Here we review recent publications (2010–2019) that advance our understanding of the effects of wildfire on surface and ground temperatures, on active layer thickness and, where permafrost is ice‐rich, on ground subsidence and the development of thermokarst features. These thermal and geomorphic changes are initiated immediately following wildfire and alter the hydrology and biogeochemistry of permafrost landscapes, including the release of previously frozen carbon. In many locations, permafrost has been resilient, with key characteristics such as active layer thickness returning to pre‐fire conditions after several decades. However, permafrost near its southern limit is losing this resiliency as a result of ongoing climate warming and increasingly common vegetation state changes. Shifts in fire return intervals, severity and extent are expected to alter the trajectories of wildfire impacts on permafrost, and to enlarge spatial impacts to more regularly include the burning of tundra areas. Modeling indicates some lowland boreal forest and tundra environments will remain resilient while uplands and areas with thin organic layers and dry soils will experience rapid and irreversible permafrost degradation. More work is needed to relate modeling to empirical studies, particularly incorporating dynamic variables such as soil moisture, snow and thermokarst development, and to identify post‐fire permafrost responses for different landscape types and regions. Future progress requires further collaboration among geocryologists, ecologists, hydrologists, biogeochemists, modelers and remote sensing specialists.

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Aboveground tree growth is a minor and decoupled fraction of boreal forest carbon input
Christoforos Pappas, Jason Maillet, Sharon Rakowski, Jennifer L. Baltzer, Alan Barr, T. Andrew Black, Simone Fatichi, Colin P. Laroque, Ashley M. Matheny, Alexandre Roy, Oliver Sonnentag, Tianshan Zha
Agricultural and Forest Meteorology, Volume 290

• 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.

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Spatial and stratigraphic variation of near‐surface ground ice in discontinuous permafrost of the taiga shield
Jason R. Paul, Steven V. Kokelj, Jennifer L. Baltzer
Permafrost and Periglacial Processes, Volume 32, Issue 1

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.

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Climate‐change refugia in boreal North America: what, where, and for how long?
Diana Stralberg, Dominique Arseneault, Jennifer L. Baltzer, Quinn E. Barber, Erin M. Bayne, Yan Boulanger, Carissa D. Brown, Hilary A. Cooke, K. J. Devito, Jason Edwards, César A. Estevo, Nadele Flynn, Lee E. Frelich, Edward H. Hogg, Mark Johnston, Travis Logan, Steven M. Matsuoka, Paul Moore, Toni Lyn Morelli, Julienne Morissette, Elizabeth A. Nelson, Hedvig K. Nenzén, Scott E. Nielsen, Marc‐André Parisien, John Pedlar, David T. Price, Fiona KA Schmiegelow, Stuart M. Slattery, Oliver Sonnentag, Daniel K. Thompson, Ellen Whitman
Frontiers in Ecology and the Environment, Volume 18, Issue 5

H latitude regions around the world are experiencing particularly rapid climate change. These regions include the 625 million ha North American boreal region, which contains 16% of the world’s forests and plays a major role in the global carbon cycle (Brandt et al. 2013). Boreal ecosystems are particularly susceptible to rapid climatedriven vegetation change initiated by standreplacing natural disturbances (notably fires), which have increased in number, extent, and frequency (Kasischke and Turetsky 2006; Hanes et al. 2018) and are expected to continue under future climate change (Boulanger et al. 2014). Such disturbances will increasingly complicate species persistence, and it will therefore be critical to identify locations of possible climatechange refugia (areas “relatively buffered from contemporary climate change”) (Morelli et al. 2016). These “slow lanes” for biodiversity will be especially important for conservation and management of boreal species and ecosystems (Morelli et al. 2020). Practically speaking, the refugia concept can translate into specific sites or regions that are expected to be more resistant to the influence of climate change than other areas (“in situ refugia”; Ashcroft 2010). Refugia may also encompass sites or regions to which species may more readily retreat as climate conditions change (“ex situ refugia”; Ashcroft 2010; Keppel et al. 2012), as well as temporary “stepping stones” (Hannah et al. 2014) linking current and future habitats. In addition to areas that are climatically buffered, fire refugia – “places that are disturbed less frequently or less severely by wildfire” (Krawchuk et al. 2016) – may also play key roles in promoting ecosystem persistence under changing conditions (Meddens et al. 2018). Previous examinations of climatechange refugia have primarily emphasized external, terrainmediated mechanisms. Factors such as topographic shading and temperature inverClimatechange refugia in boreal North America: what, where, and for how long?

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Fuel availability not fire weather controls boreal wildfire severity and carbon emissions
Xanthe J. Walker, Brendan M. Rogers, Sander Veraverbeke, Jill F. Johnstone, Jennifer L. Baltzer, Kirsten Barrett, Laura Bourgeau‐Chavez, Nicola J. Day, William J. de Groot, Catherine M. Dieleman, S. J. Goetz, Elizabeth Hoy, Liza K. Jenkins, Evan S. Kane, Marc‐André Parisien, Stefano Potter, Edward A. G. Schuur, M. R. Turetsky, Ellen Whitman, Michelle C. Mack
Nature Climate Change, Volume 10, Issue 12

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.

2019

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Wildfire severity reduces richness and alters composition of soil fungal communities in boreal forests of western Canada
Nicola J. Day, Kari E. Dunfield, Jill F. Johnstone, Michelle C. Mack, M. R. Turetsky, Xanthe J. Walker, Alison L. White, Jennifer L. Baltzer
Global Change Biology, Volume 25, Issue 7

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.

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Increasing wildfires threaten historic carbon sink of boreal forest soils
Xanthe J. Walker, Jennifer L. Baltzer, Steven G. Cumming, Nicola J. Day, Christopher Ebert, S. J. Goetz, Jill F. Johnstone, Stefano Potter, Brendan M. Rogers, Edward A. G. Schuur, M. R. Turetsky, Michelle C. Mack
Nature, Volume 572, Issue 7770

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.

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Tall Shrubs Mediate Abiotic Conditions and Plant Communities at the Taiga–Tundra Ecotone
Cory A. Wallace, Jennifer L. Baltzer
Ecosystems, Volume 23, Issue 4

Shrub expansion has occurred across much of the arctic tundra over the past century. Increasing dominance of woody vegetation is expected to have global influences on climate patterns and lead to local changes in hydrological function and nutrient cycling. Changing abiotic conditions associated with shrubs will likely alter the relative fitness of neighbouring plants resulting in distinct community composition. Here, we use an extensive set of paired abiotic and biotic data to investigate the capacity for Alnus alnobetula (green alder) patches to modify the habitat of the local plant community at the taiga–tundra ecotone of the Northwest Territories, Canada. Plots were established across topographic positions in ten alder patches and adjacent, alder-free tundra. Habitat corresponded to the strongest gradient of among-site variation in abiotic measures and plant community composition, indicating that alder patch growing conditions were distinct from those of alder-free tundra. Slope position was generally unimportant in determining environmental conditions. Alder patches changed the vertical structure of the understory by increasing the maximum height of birch. Tall shrubs also decreased the richness of tundra specialists, suggesting that these species face competitive pressures from shrub expansion at the southern edge of their ranges. Our findings demonstrate that tall shrub patches can substantially modify their local environment in taiga–tundra ecotone systems, altering available habitat and acting as niche constructors for the local plant community. These habitats will therefore be important to consider in regional predictions of hydrology, nutrient cycling, and biodiversity as shrubs continue to expand across the arctic.

2018

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Environmental and taxonomic controls of carbon and oxygen stable isotope composition in Sphagnum across broad climatic and geographic ranges
Gustaf Granath, Håkan Rydin, Jennifer L. Baltzer, Fia Bengtsson, Nicholas Boncek, Luca Bragazza, Zhao‐Jun Bu, Simon J. M. Caporn, Ellen Dorrepaal, О. В. Галанина, Mariusz Gałka, Anna Ganeva, David P. Gillikin, Irina Goia, Nadezhda Goncharova, Michal Hájek, Akira Haraguchi, Lorna I. Harris, Elyn Humphreys, Martin Jiroušek, Katarzyna Kajukało, Edgar Karofeld, Natalia G. Koronatova, Natalia P. Kosykh, Mariusz Lamentowicz, Е. Д. Лапшина, Juul Limpens, Maiju Linkosalmi, Jinze Ma, Marguerite Mauritz, Tariq M. Munir, Susan M. Natali, Rayna Natcheva, Maria Noskova, Richard J. Payne, Kyle Pilkington, Sean Robinson, Bjorn J. M. Robroek, Line Rochefort, David Singer, Hans K. Stenøien, Eeva‐Stiina Tuittila, Kai Vellak, Anouk Verheyden, J. M. Waddington, Steven K. Rice

Abstract. Rain-fed peatlands are dominated by peat mosses (Sphagnum sp.), which for their growth depend on elements from the atmosphere. As the isotopic composition of carbon (12,13C) and oxygen (16,18O) of these Sphagnum mosses are affected by environmental conditions, the dead Sphagnum tissue accumulated in peat constitutes a potential long-term archive that can be used for climate reconstruction. However, there is a lack of adequate understanding of how isotope values are influenced by environmental conditions, which restricts their current use as environmental and palaeoenvironmental indicators. Here we tested (i) to what extent C and O isotopic variation in living tissue of Sphagnum is species-specific and associated with local hydrological gradients, climatic gradients (evapotranspiration, temperature, precipitation), and elevation; (ii) if the C isotopic signature can be a proxy for net primary productivity (NPP) of Sphagnum; and (iii) to what extent Sphagnum tissue δ18O tracks the δ18O isotope signature of precipitation. In total, we analysed 337 samples from 93 sites across North America and Eurasia using two important peat-forming Sphagnum species (S. magellanicum, S. fuscum) common to the Holartic realm. There were differences in δ13C values between species. For S. magellanicum δ13C decreased with increasing height above the water table (HWT, R2 = 17 %) and was positively correlated to productivity (R2 = 7 %). Together these two variables explained 46 % of the between-site variation in δ13C values. For S. fuscum, productivity was the only significant predictor of δ13C (total R2 = 6 %). For δ18O values, ca. 90 % of the variation was found between sites. Globally-modelled annual δ18O values in precipitation explained 69% of the between-site variation in tissue δ18O. S. magellanicum showed lower δ18O enrichment than S. fuscum (−0.83 ‰ lower) . Elevation and climatic variables were weak predictors of tissue δ18O values after controlling for δ18O values of the precipitation. To summarise, our study provides evidence for (a) good predictability of tissue δ18O values from modelled annual δ18O values in precipitation, and (b) the possibility to relate tissue δ13C values to HWT and NPP, but this appears to be species-dependent. These results suggest that isotope composition can be used at a large scale for climatic reconstructions but that such models should be species-specific.

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Minor contribution of overstorey transpiration to landscape evapotranspiration in boreal permafrost peatlands
Rebecca K. Warren, Christoforos Pappas, Manuel Helbig, L. Chasmer, Aaron Berg, Jennifer L. Baltzer, W. L. Quinton, Oliver Sonnentag
Ecohydrology, Volume 11, Issue 5

Evapotranspiration (ET) is a key component of the water cycle, whereby accurate partitioning of ET into evaporation and transpiration provides important information about the intrinsically coupled carbon, water, and energy fluxes. Currently, global estimates of partitioned evaporative and transpiration fluxes remain highly uncertain, especially for high‐latitude ecosystems where measurements are scarce. Forested peat plateaus underlain by permafrost and surrounded by permafrost‐free wetlands characterize approximately 60% (7.0 × 107 km2) of Canadian peatlands. In this study, 22 Picea mariana (black spruce) individuals, the most common tree species of the North American boreal forest, were instrumented with sap flow sensors within the footprint of an eddy covariance tower measuring ET from a forest–wetland mosaic landscape. Sap flux density (JS), together with remote sensing data and in situ measurements of canopy structure, was used to upscale tree‐level JS to overstorey transpiration (TBS). Black spruce trees growing in nutrient‐poor permafrost peat soils were found to have lower mean JS than those growing in mineral soils. Overall, TBS contributed less than 1% to landscape ET. Climate‐change‐induced forest loss and the expansion of wetlands may further minimize the contributions of TBS to ET and increase the contribution of standing water.

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Cross‐scale controls on carbon emissions from boreal forest megafires
Xanthe J. Walker, Brendan M. Rogers, Jennifer L. Baltzer, Steven G. Cumming, Nicola J. Day, S. J. Goetz, Jill F. Johnstone, Edward A. G. Schuur, M. R. Turetsky, Michelle C. Mack
Global Change Biology, Volume 24, Issue 9

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.

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Boreal tree hydrodynamics: asynchronous, diverging, yet complementary
Christoforos Pappas, Ashley M. Matheny, Jennifer L. Baltzer, Alan Barr, T. Andrew Black, Gil Bohrer, Matteo Detto, Jason Maillet, Alexandre Roy, Oliver Sonnentag, Jilmarie Stephens
Tree Physiology, Volume 38, Issue 7

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.

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Quantification of uncertainties in conifer sap flow measured with the thermal dissipation method
Richard L. Peters, Patrick Fonti, David Frank, Rafael Poyatos, Christoforos Pappas, Ansgar Kahmen, Vinicio Carraro, Angela Luisa Prendin, Loïc Schneider, Jennifer L. Baltzer, Greg A. Baron‐Gafford, Lars Dietrich, Ingo Heinrich, R. L. Minor, Oliver Sonnentag, Ashley M. Matheny, Maxwell G. Wightman, Kathy Steppe
New Phytologist, Volume 219, Issue 4

Trees play a key role in the global hydrological cycle and measurements performed with the thermal dissipation method (TDM) have been crucial in providing whole-tree water-use estimates. Yet, different data processing to calculate whole-tree water use encapsulates uncertainties that have not been systematically assessed. We quantified uncertainties in conifer sap flux density (Fd ) and stand water use caused by commonly applied methods for deriving zero-flow conditions, dampening and sensor calibration. Their contribution has been assessed using a stem segment calibration experiment and 4 yr of TDM measurements in Picea abies and Larix decidua growing in contrasting environments. Uncertainties were then projected on TDM data from different conifers across the northern hemisphere. Commonly applied methods mostly underestimated absolute Fd . Lacking a site- and species-specific calibrations reduced our stand water-use measurements by 37% and induced uncertainty in northern hemisphere Fd . Additionally, although the interdaily variability was maintained, disregarding dampening and/or applying zero-flow conditions that ignored night-time water use reduced the correlation between environment and Fd . The presented ensemble of calibration curves and proposed dampening correction, together with the systematic quantification of data-processing uncertainties, provide crucial steps in improving whole-tree water-use estimates across spatial and temporal scales.

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Assessing local adaptation vs. plasticity under different resource conditions in seedlings of a dominant boreal tree species
Anastasia E. Sniderhan, Gordon G. McNickle, Jennifer L. Baltzer
AoB PLANTS, Volume 10, Issue 1

Under changing climate conditions, understanding local adaptation of plants is crucial to predicting the resilience of ecosystems. We selected black spruce (Picea mariana), the most dominant tree species in the North American boreal forest, in order to evaluate local adaptation vs. plasticity across regions experiencing some of the most extreme climate warming globally. Seeds from three provenances across the latitudinal extent of this species in northwestern Canada were planted in a common garden study in growth chambers. Two levels of two resource conditions were applied (low/high nutrient and ambient/elevated CO2) in a fully factorial design and we measured physiological traits, allocational traits, growth and survival. We found significant differences in height, root length and biomass among populations, with southern populations producing the largest seedlings. However, we did not detect meaningful significant differences among nutrient or CO2 treatments in any traits measured, and there were no consistent population-level differences in physiological traits or allocation patterns. We found that there was greater mortality after simulated winter in the high nutrient treatment, which may reflect an important shift in seedling growth strategies under increased resource availability. Our study provides important insight into how this dominant boreal tree species might respond to the changing climate conditions predicted in this region.

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Soil organic layer combustion in boreal black spruce and jack pine stands of the Northwest Territories, Canada
Xanthe J. Walker, Jennifer L. Baltzer, Steven G. Cumming, Nicola J. Day, Jill F. Johnstone, Brendan M. Rogers, Kylen Solvik, M. R. Turetsky, Michelle C. Mack
International Journal of Wildland Fire, Volume 27, Issue 2

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.

2016

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Losing Legacies, Ecological Release, and Transient Responses: Key Challenges for the Future of Northern Ecosystem Science
M. R. Turetsky, Jennifer L. Baltzer, Jill F. Johnstone, Michelle C. Mack, Kevin S. McCann, Edward A. G. Schuur
Ecosystems, Volume 20, Issue 1

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.