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  1. Abstract

    Climate warming has increased permafrost thaw in arctic tundra and extended the duration of annual thaw (number of thaw days in summer) along soil profiles. Predicting the microbial response to permafrost thaw depends largely on knowing how increased thaw duration affects the composition of the soil microbiome. Here, we determined soil microbiome composition from the annually thawed surface active layer down through permafrost from two tundra types at each of three sites on the North Slope of Alaska, USA. Variations in soil microbial taxa were found between sites up to ~90 km apart, between tundra types, and between soil depths. Microbiome differences at a site were greatest across transitions from thawed to permafrost depths. Results from correlation analysis based on multi‐decadal thaw surveys show that differences in thaw duration by depth were significantly, positively correlated with the abundance of dominant taxa in the active layer and negatively correlated with dominant taxa in the permafrost. Microbiome composition within the transition zone was statistically similar to that in the permafrost, indicating that recent decades of intermittent thaw have not yet induced a shift from permafrost to active‐layer microbes. We suggest that thaw duration rather than thaw frequency has a greater impact on the composition of microbial taxa within arctic soils.

     
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  2. Abstract

    We investigated how lake thermal processes responded to whole lake warming manipulation in an arctic lake through observations and numerical modeling. The warming manipulation was conducted by artificially heating the epilimnion as a proxy for climate warming. We performed numerical modeling with an improved lake scheme based on the Community Land Model (CLM). We simulated a control run (CTL) without warming and a warming manipulation simulation (WARM). Results indicated WARM accurately captured observed temperatures where water stratification was extended in time, and water stability was strengthened. Two additional sensitivity tests with different warming onset dates and of the same warming duration showed that earlier warming onsets are predicted to make the water column more stable and less easily mixed relative to a later onset of warming. The results provide a more complete understanding of lake thermal processes in arctic freshwater lake systems and how they will respond to predicted future warming.

     
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  3. Abstract

    Phosphorus (P) limits or co‐limits plant and microbial life in multiple ecosystems, including the arctic tundra. Although current global carbon (C) models focus on the coupling between soil nitrogen (N) and C, ecosystem P response to climate warming may also influence the global C cycle. Permafrost soils may see enhanced or reduced P availability under climate warming through multiple mechanisms including changing litter inputs through plant community change, changing plant–microbial dynamics, altered rates of mineralization of soil organic P through increased microbial activity, and newly exposed mineral‐bound P via deeper thaw. We investigated the effect of long‐term warming on plant leaf, multiple soil and microbial C, N, and P pools, and microbial extracellular enzyme activities, in Alaskan tundra plots underlain by permafrost. Here, we show that 25 yr of experimental summer warming increases community‐level plant leaf P through changing community composition to favour relatively P‐rich plant species. However, despite associated increases in P‐rich litter inputs, we found only a few responses in the belowground pools of P available for plant and microbial uptake, including a weak positive response for citric acid–extractable PO4in the surface soil, a decrease in microbial biomass P, and no change in soil P (or C or N) stocks. This weak, neutral, or negative belowground P response to warming despite enhanced litter P inputs is consistent with a growing number of studies in the arctic tundra that find no long‐term response of soil C and N stocks to warming.

     
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  4. Abstract

    We use a simple model of coupled carbon and nitrogen cycles in terrestrial ecosystems to examine how “explicitly representing grazers” vs. “having grazer effects implicitly aggregated in with other biogeochemical processes in the model” alters predicted responses to elevated carbon dioxide and warming. The aggregated approach can affect model predictions because grazer‐mediated processes can respond differently to changes in climate compared with the processes with which they are typically aggregated. We use small‐mammal grazers in a tundra as an example and find that the typical three‐to‐four‐year cycling frequency is too fast for the effects of cycle peaks and troughs to be fully manifested in the ecosystem biogeochemistry. We conclude that implicitly aggregating the effects of small‐mammal grazers with other processes results in an underestimation of ecosystem response to climate change, relative to estimations in which the grazer effects are explicitly represented. The magnitude of this underestimation increases with grazer density. We therefore recommend that grazing effects be incorporated explicitly when applying models of ecosystem response to global change.

     
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  5. Abstract

    We use the Multiple Element Limitation (MEL) model to examine responses of 12 ecosystems to elevated carbon dioxide (CO2), warming, and 20% decreases or increases in precipitation. Ecosystems respond synergistically to elevated CO2, warming, and decreased precipitation combined because higher water‐use efficiency with elevated CO2and higher fertility with warming compensate for responses to drought. Response to elevated CO2, warming, and increased precipitation combined is additive. We analyze changes in ecosystem carbon (C) based on four nitrogen (N) and four phosphorus (P) attribution factors: (1) changes in total ecosystem N and P, (2) changes in N and P distribution between vegetation and soil, (3) changes in vegetation C:N and C:P ratios, and (4) changes in soil C:N and C:P ratios. In the combined CO2and climate change simulations, all ecosystems gain C. The contributions of these four attribution factors to changes in ecosystem C storage varies among ecosystems because of differences in the initial distributions of N and P between vegetation and soil and the openness of the ecosystem N and P cycles. The net transfer of N and P from soil to vegetation dominates the C response of forests. For tundra and grasslands, the C gain is also associated with increased soil C:N and C:P. In ecosystems with symbiotic N fixation, C gains resulted from N accumulation. Because of differences in N versus P cycle openness and the distribution of organic matter between vegetation and soil, changes in the N and P attribution factors do not always parallel one another. Differences among ecosystems in C‐nutrient interactions and the amount of woody biomass interact to shape ecosystem C sequestration under simulated global change. We suggest that future studies quantify the openness of the N and P cycles and changes in the distribution of C, N, and P among ecosystem components, which currently limit understanding of nutrient effects on C sequestration and responses to elevated CO2and climate change.

     
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  6. Abstract

    Riveraufeis(ow′ fīse) are widespread features of the arctic cryosphere. They form when river channels become locally restricted by ice, resulting in cycles of water overflow and freezing and the accumulation of ice, with someaufeisattaining areas of ~ 25 + km2and thicknesses of 6+ m. During winter, unfrozen sediments beneath the insulating ice layer provide perennial groundwater‐habitat that is otherwise restricted in regions of continuous permafrost. Our goal was to assess whetheraufeisfacilitate the occurrence of groundwater invertebrate communities in the Arctic. We focused on a singleaufeisecosystem (~ 5 km2by late winter) along the Kuparuk River in arctic Alaska. Subsurface invertebrates were sampled during June and August 2017 from 50 3.5‐cm diameter PVC wells arranged in a 5 × 10 array covering ~ 40 ha. Surface invertebrates were sampled using a quadrat approach. We documented a rich assemblage of groundwater invertebrates (49 [43–54] taxa,[95% confidence limits]) that was distributed below the sediment surface to a mean depth of ~ 69 ± 2 cm (± 1 SE) throughout the entire well array. Although community structure differed significantly between groundwater and surface habitats, the taxa richness from wells and surface sediments (43 [35–48] taxa) did not differ significantly, which was surprising given lower richness in subsurface habitats of large, riverine gravel‐aquifer systems shown elsewhere. This is the first demonstration of a rich and spatially extensive groundwater fauna in a region of continuous permafrost. Given the geographic extent ofaufeisfields, localized groundwater‐dependent ecosystems may be widespread in the Arctic.

     
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  7. Abstract

    Large and small mammalian herbivores are present in most vegetated areas in the Arctic and often have large impacts on plant community composition and ecosystem functioning. The relative importance of different herbivores and especially how their specific impact on the vegetation varies across the Arctic is however poorly understood.

    Here, we investigate how large and small herbivores influence vegetation density and plant community composition in four arctic vegetation types in Scandinavia and Alaska. We used a unique set of exclosures, excluding only large (reindeer and muskoxen) or all mammalian herbivores (also voles and lemmings) for at least 20 years.

    We found that mammalian herbivores in general decreased leaf area index, NDVI, and abundance of vascular plants in all four locations, even though the strength of the effect and which herbivore type caused these effects differed across locations. In three locations, herbivore presence caused contrasting plant communities, but not in the location with lowest productivity. Large herbivores had a negative effect on plant height, whereas small mammalian herbivores increased species diversity by decreasing dominance of the initially dominating plant species. Above‐ or belowground disturbances caused by herbivores were found to play an important role in shaping the vegetation in all locations.

    Synthesis:Based on these results, we conclude that both small and large mammalian herbivores influence vegetation in Scandinavia and Alaska in a similar way, some of which can mitigate effects of climate change. We also see important differences across locations, but these depend rather on local herbivore and plant community composition than large biogeographical differences among continents.

     
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  8. Abstract

    Effects of climate change‐driven disturbance on lake ecosystems can be subtle; indirect effects include increased nutrient loading that could impact ecosystem function. We designed a low‐level fertilization experiment to mimic persistent, climate change‐driven disturbances (deeper thaw, greater weathering, or thermokarst failure) delivering nutrients to arctic lakes. We measured responses of pelagic trophic levels over 12 yr in a fertilized deep lake with fish and a shallow fishless lake, compared to paired reference lakes, and monitored recovery for 6 yr. Relative to prefertilization in the deep lake, we observed a maximum pelagic response in chla(+201%), dissolved oxygen (DO, −43%), and zooplankton biomass (+88%) during the fertilization period (2001–2012). Other responses to fertilization, such as water transparency and fish relative abundance, were delayed, but both ultimately declined. Phyto‐ and zooplankton biomass and community composition shifted with fertilization. The effects of fertilization were less pronounced in the paired shallow lakes, because of a natural thermokarst failure likely impacting the reference lake. In the deep lake there was (a) moderate resistance to change in ecosystem functions at all trophic levels, (b) eventual responses were often nonlinear, and (c) postfertilization recovery (return) times were most rapid at the base of the food web (2–4 yr) while higher trophic levels failed to recover after 6 yr. The timing and magnitude of responses to fertilization in these arctic lakes were similar to responses in other lakes, suggesting indirect effects of climate change that modify nutrient inputs may affect many lakes in the future.

     
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  9. Abstract

    In arctic tundra, large and small mammalian herbivores have substantial impacts on the vegetation community and consequently can affect the magnitude of carbon cycling. However, herbivores are often absent from modern carbon cycle models, partly because relatively few field studies focus on herbivore impacts on carbon cycling. Our objectives were to quantify the impact of 21 years of large herbivore and large and small herbivore exclusion on carbon cycling during peak growing season in a dry heath tundra community. When herbivores were excluded, we observed a significantly greater leaf area index as well as greater vascular plant abundance. While we did not observe significant differences in deciduous dwarf shrub abundance across treatments, evergreen dwarf shrub abundance was greater where large and small herbivores were excluded. Both foliose and fruticose lichen abundance were higher in the large herbivore, but not the small and large herbivore exclosures. Net ecosystem exchange (NEE) likewise indicated the highest carbon uptake in the exclosure treatments and lowest uptake in the control (CT), suggesting that herbivory decreased the capacity of dry heath tundra to take up carbon. Moreover, our calculated NEE for average light and temperature conditions for July 2017, when our measurements were taken, indicated that the tundra was a carbon source in CT, but was a carbon sink in both exclosure treatments, indicating removal of grazing pressure can change the carbon balance of dry heath tundra. Collectively, these findings suggest that herbivore absence can lead to changes in plant community structure of dry heath tundra that in turn can increase its capacity to take up carbon.

     
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  10. Abstract

    The climate of the Arctic region is changing rapidly, with important implications for permafrost, vegetation communities, and transport of solutes by streams and rivers to the Arctic Ocean. While research on Arctic streams and rivers has accelerated in recent years, long‐term records are relatively rare compared to temperate and tropical regions. We began monitoring the upper Kuparuk River in 1983 as part of a long‐term, low‐level, whole‐season phosphorus enrichment of a 4–6 km experimental reach, which was subsequently incorporated into the Arctic Long‐Term Ecological Research (Arctic LTER) programme. The phosphorus enrichment phase of the Upper Kuparuk River Experiment (UKRE) ran continuously for 34 seasons, fundamentally altering the community structure and function of the Fertilized reach. The objectives of this paper are to (a) update observations of the environmental conditions in the Kuparuk River region as revealed by long‐term, catchment‐level monitoring, (b) compare long‐term trends in biogeochemical characteristics of phosphorus‐enriched and reference reaches of the Kuparuk River, and (c) report results from a new ‘ReFertilization’ experiment. During the UKRE, temperature and discharge did not change significantly, though precipitation increased slightly. However, the UKRE revealed unexpected community state changes attributable to phosphorus enrichment (e.g., appearance of colonizing bryophytes) and long‐term legacy effects of these state changes after cessation of the phosphorus enrichment. The UKRE also revealed important biogeochemical trends (e.g., increased nitrate flux and benthic C:N, decreased DOP flux). The decrease in DOP is particularly notable in that this may be a pan‐Arctic trend related to permafrost thaw and exposure to new sources of iron that reduce phosphorus mobility to streams and rivers. The trends revealed by the UKRE would have been difficult or impossible to identify without long‐term, catchment level research and may have important influences on connections between Arctic headwater catchments and downstream receiving waters, including the Arctic Ocean.

     
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