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Most tundra carbon flux modeling relies on leaf area index (LAI), generally estimated from measurements of canopy greenness using the normalized difference vegetation index (NDVI), to estimate the direction and magnitude of fluxes. However, due to the relative sparseness and low stature of tundra canopies, such models do not explicitly consider the influence of variation in tundra canopy structure on carbon flux estimates. Structure from motion (SFM), a photogrammetric method for deriving three-dimensional (3D) structure from digital imagery, is a non-destructive method for estimating both fine-scale canopy structure and LAI. To understand how variation in 3D canopy structure affects ecosystem carbon fluxes in Arctic tundra, we adapted an existing NDVI-based tundra carbon flux model to include variation in SFM-derived canopy structure and its interaction with incoming sunlight to cast shadows on canopies. Our study system consisted of replicate plots of dry heath tundra that had been subjected to three herbivore exclosure treatments (an exclosure-free control [CT], large mammals exclosure), and a large and small mammal exclosure [ExLS]), providing the range of 3D canopy structures employed in our study. We found that foliage within the more structurally complex surface of CT canopies received significantly less light over the course of the day than canopies within both exclosure treatments. This was especially during morning and evening hours, and was reflected in modeled rates of net ecosystem exchange (NEE) and gross primary productivity (GPP). We found that in the ExLS treatment, SFM-derived estimates of GPP were significantly lower and NEE significantly higher than those based on LAI alone. Our results demonstrate that the structure of even simple tundra vegetation canopies can have significant impacts on tundra carbon fluxes and thus need to be accounted for.

 

Percent cover of tundra vole and brown lemming structures collected from within the Team Vole enclosure/exclosure fences near Nome, Toolik, Utqiagvik, AK 2019. 

We use a simple model of coupled carbon and nitrogen cycles in terrestrial ecosystems to examine how explicitly representing grazers versus 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 from the processes with which they are typically aggregated. We use small-mammal grazers in arctic 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. 

We use a simple model of coupled carbon and nitrogen cycles in terrestrial ecosystems to examine how explicitly representing grazers versus 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 from the processes with which they are typically aggregated. We use small-mammal grazers in arctic 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. 

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.

 

We present a framework for assessing biogeochemical recovery of terrestrial ecosystems from disturbance. We identify three recovery phases. In Phase 1, nitrogen is redistributed from soil organic matter to vegetation, but the ecosystem continues to lose nitrogen because the recovering vegetation cannot take up nitrogen as fast as it is released from soil. In Phase 2, the ecosystem begins re-accumulating nitrogen and converges on a quasi-steady state in which vegetation and soil-microbial processes are in balance. In Phase 3, vegetation and soil-microbial processes remain in balance and the ecosystem slowly re-accumulates the remaining nitrogen. Phase 3 follows a balanced-accumulation trajectory along a continuum of quasi-steady states that approaches the true steady state asymptotically. We examine the effects of three ecosystem properties on recovery: openness of the nitrogen cycle, nitrogen distribution in and turnover between vegetation and soils, and the proportion of nitrogen losses that are in a refractory form. Openness exacerbates Phase 1 nitrogen losses but speeds recovery in Phases 2 and 3. A high fraction of ecosystem nitrogen in vegetation, resulting from nitrogen turnover that is slow in vegetation but fast in soil, exacerbates Phase 1 nitrogen losses but speeds recovery in Phases 2 and 3. A high proportion of nitrogen loss in refractory form mitigates Phase 1 nitrogen losses and speeds recovery in Phases 2 and 3. Application of our conceptual framework requires empirical recognition of the continuum of quasi-steady states constituting the balanced-accumulation trajectory and a distinction between the balanced-accumulation trajectory and the true steady state. 

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.

 

Understanding arctic ecosystem function is key to understanding future global carbon (C) and nutrient cycling processes. However, small mammal herbivores can have effects on ecosystems as structure builders and these effects have been underrepresented in the understanding of arctic systems.

We examined the impact of small mammal structures (hay piles, runways, latrines) on soils and plants in three arctic tundra regions near Utqiaġvik, Toolik Lake, and Nome, Alaska. Our aims were to (1) examine how vole and lemming structures influence plant and soil nutrient pools and microbial processes, (2) determine if structure effects were similar across tundra system types, and (3) understand how changes in the abundance and cover of these structures during different phases of small mammal multi‐annual population cycles might influence biogeochemical cycling.

In general, small mammal structures increased nitrogen (N) availability in soils, although effects varied by study region. Across study regions, hay piles were relatively uncommon (lowest % cover) but increased multiple soil N and P pools, C‐ and N‐acquiring enzyme activities, and leaf phosphorus (P) concentrations, with the specific nutrient variables and size of the effects varying by study region. Latrines had the second highest cover and influenced multiple C, N and P pools, but their effects were mainly observed within a single region. Lastly, runways had the highest % cover of all activity types but increased the fewest number of soil nutrient variables.

We conclude that by influencing soil nutrient availability and biogeochemical cycling, small mammal structures can influence bottom‐up regulation of ecosystem function, particularly during the high phase of the small mammal population cycle. Future changes in these population cycles might alter the role of small mammals in the Arctic and have lasting effects on system processes.

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