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  1. 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 Δ14C (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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  2. Abstract

    In post‐fire Siberian larch forests, where tree density can vary within a burn perimeter, shrubs constitute a substantial portion of the vegetation canopy. Leaf area index (LAI), defined as the one‐sided total green leaf area per unit ground surface area, is useful for characterizing variation in plant canopies. We estimated LAI with allometry for trees and tall shrubs (>0.5 and <1.5 m) across 26 sites with varying tree stem density (0.05–3.3 stems/m2) and canopy cover (4.6%–76.9%) in a uniformly‐aged mature Siberian larch forest that regenerated following a fire ∼75 years ago. We investigated relationships between tree density, tree LAI, and tall shrub LAI, and between LAI and satellite observations of Normalized Difference and Enhanced Vegetation Indices (NDVI and EVI). Across the density gradient, tree LAI increases with increasing tree density, while tall shrub LAI decreases, exhibiting no patterns in combined tree‐shrub LAI. We also found significant positive relationships between tall shrub LAI and NDVI/EVI from PlanetScope and Landsat imagery. These findings suggest that tall shrubs compensate for lower tree LAI in tree canopy gaps, forming a canopy with contiguous combined tree‐shrub LAI across the density gradient. Our findings suggest that NDVI and EVI are more sensitive to variation in tall shrub canopies than variation in tree canopies or combined tree‐shrub canopies in these ecosystems. The results improve our understanding of the relationships between forest density and tree and shrub leaf area and have implications for interpreting spatial variability in LAI, NDVI, and EVI in Siberian boreal forests.

     
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  3. Abstract

    Foundation species have disproportionately large impacts on ecosystem structure and function. As a result, future changes to their distribution may be important determinants of ecosystem carbon (C) cycling in a warmer world. We assessed the role of a foundation tussock sedge (Eriophorum vaginatum) as a climatically vulnerable C stock using field data, a machine learning ecological niche model, and an ensemble of terrestrial biosphere models (TBMs). Field data indicated that tussock density has decreased by ∼0.97 tussocks per m2over the past ∼38 years on Alaska’s North Slope from ∼1981 to 2019. This declining trend is concerning because tussocks are a large Arctic C stock, which enhances soil organic layer C stocks by 6.9% on average and represents 745 Tg C across our study area. By 2100, we project that changes in tussock density may decrease the tussock C stock by 41% in regions where tussocks are currently abundant (e.g. −0.8 tussocks per m2and −85 Tg C on the North Slope) and may increase the tussock C stock by 46% in regions where tussocks are currently scarce (e.g. +0.9 tussocks per m2and +81 Tg C on Victoria Island). These climate-induced changes to the tussock C stock were comparable to, but sometimes opposite in sign, to vegetation C stock changes predicted by an ensemble of TBMs. Our results illustrate the important role of tussocks as a foundation species in determining future Arctic C stocks and highlight the need for better representation of this species in TBMs.

     
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  4. Abstract

    As climate warms, tree density at the taiga–tundra ecotone (TTE) is expected to increase, which may intensify competition for belowground resources in this nitrogen (N)‐limited environment. To determine the impacts of increased tree density on N cycling and productivity, we examined edaphic properties indicative of soil N availability along with aboveground and belowground tree‐level traits and stand characteristics related to carbon (C) and N cycling across a tree density gradient of monodominant larch (Larix cajanderi) at the TTE in far northeastern Siberia. We found no consistent evidence from soil, tree, or stand‐level N cycling characteristics of lower N availability or greater intraspecific competition for N with increased density. Active layer thickness declined, but resin‐sorbed N and soil organic layer thickness did not covary with increased tree density. There was, however, greater allocation belowground to stand‐level coarse and fine roots with increased tree density, an allocation pattern suggestive of limited soil resources. Foliar traits related to C (%C, δ13C, and resorption) were responsive to density indicating the importance of non‐nutrient resources, like light, to foliar stoichiometry. As tree density increased and individual trees had lower productivity, tree‐level N and biomass pools aboveground and belowground declined tracking decreases in N uptake, N resorption, N use efficiency, and allocation to slow cycling tissues like wood. At the stand level, our findings show high N turnover with increased N acquisition, allocation to short‐lived tissues with relatively high N content and reduced N residence time, and greater stand productivity as tree density increased. Yet, these positive relationships were curtailed at the highest tree densities. Our observations of shifts in biomass, C and N allocation, and loss aboveground, along with greater root density with increased tree density, could have strong impacts on C and N cycling and should be represented in models of TTE dynamics and feedbacks to climate.

     
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  5. Abstract

    Greater tree density and forest productivity at the tundra–taiga ecotone (TTE) are expected with climate warming, with potential feedbacks to the climate system. Yet, competition for nitrogen (N) may impact TTE dynamics. Greater tree density will likely increase N demand, while reducing N supply through soil shading and slower decomposition rates. We explored whether characteristics of roots and root‐associated fungi important to N acquisition responded to changes in density at the TTE and were related to above‐ground stand productivity and N cycling.

    We measured rooting depth, uptake of N forms among soil layers and ectomycorrhizal (EcM) colonization and composition along a natural tree density gradient of monodominant larchLarix cajanderiin northeastern Siberia. We tested relationships between larch root and fungal characteristics, above‐ground productivity and stand‐level N cycling parameters.

    Overall, there was preferential uptake of ammonium compared to glycine or nitrate. Nitrogen uptake was greatest in shallow soils of the organic horizon and related to root chemistry, root‐associated fungi and above‐ground N cycling parameters, but the direction of these relationships depended on N form. Uptake of different N forms, rooting depth and EcM colonization and composition were not related to tree density, but fungal composition was correlated with root N chemistry and above‐ground N cycling parameters. In addition to EcM, the abundance of dark septate endophytes and other ascomycetous taxa was positively related to N uptake and above‐ground N cycling parameters.

    Synthesis. There was little impact of tree density on root and fungal parameters related to N acquisition suggesting intraspecific larch competition for N was not amplified with increased density. There was, however, a strong impact of root‐associated fungi on N uptake and stand N dynamics regardless of tree density. Together, this suggests an important role of root‐associated fungi on broadscale patterns of N cycling in TTE larch forests independent of changes in tree density expected with climate warming.

     
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  6. Vast amounts of organic carbon are stored in Arctic soils. Much of this is in the form of peat, a layer of decomposing plant matter. Arctic wildfires release this carbon to the atmosphere as carbon dioxide (CO 2 ) ( 1 ) and contribute to global warming. This creates a feedback loop in which accelerated Arctic warming ( 2 ) dries peatland soils, which increases the likelihood of bigger, more frequent wildfires in the Arctic and releases more CO 2 , which further contributes to warming. Although this feedback mechanism is qualitatively understood, there remain uncertainties about its details. On page 532 of this issue, Descals et al. ( 3 ) analyze data from the 2019 and 2020 wildfire seasons in the Siberian Arctic and predict the extent of carbon-rich soils likely to burn in the area with future warming. Critically, they suggest that even minor increases in temperature above certain thresholds may promote increasingly larger wildfires. 
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  7. 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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  8. null (Ed.)
    Cajander larch (Larix cajanderi Mayr.) forests of the Siberian Arctic are experiencing increased wildfire activity in conjunction with climate warming. These shifts could affect postfire variation in the density and arrangement of trees and understory plant communities. To better understand how understory plant composition, abundance, and diversity vary with tree density, we surveyed understory plant communities and stand characteristics (e.g., canopy cover, active layer depth, and soil organic layer depth) within 25 stands representing a density gradient of similarly-aged larch trees that established following a 1940 fire near Cherskiy, Russia. Understory plant diversity and mean total plant abundance decreased with increased canopy cover. Canopy cover was also the most important variable affecting individual species’ abundances. In general, tall shrubs (e.g., Betula nana subsp. exilis) were more abundant in low-density stands with high light availability, and mosses (e.g., Sanionia spp.) were more abundant in high-density stands with low light availability. These results provide evidence that postfire variation in tree recruitment affects understory plant community composition and diversity as stands mature. Therefore, projected increases in wildfire activity in the Siberian Arctic could have cascading impacts on forest structure and composition in both overstory and understory plant communities. 
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  9. null (Ed.)
    The transition zone between the northern boreal forest and the arctic tundra, known as the tundra-taiga ecotone (TTE) has undergone rapid warming in recent decades. In response to this warming, tree density, growth, and stand productivity are expected to increase. Increases in tree density have the potential to negate the positive impacts of warming on tree growth through a reduction in the active layer and an increase in competitive interactions. We assessed the effects of tree density on tree growth and climate-growth responses of Cajander larch (Larix cajanderi) and on trends in the normalized difference vegetation index (NDVI) in the TTE of Northeast Siberia. We examined 19 mature forest stands that all established after a fire in 1940 and ranged in tree density from 300 to 37,000 stems ha-1. High density stands with shallow active layers had lower tree growth, higher stand productivity, and more negative growth responses to growing season temperatures compared to low density stands with deep active layers. Variation in stand productivity across the density gradient was not captured by Landsat derived NDVI, but NDVI did capture annual variations in stand productivity. Our results suggest that the expected increases in tree density following fires at the TTE may effectively limit tree growth and that NDVI is unlikely to capture increasing productivity associated with changes in tree density. 
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