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We conducted a manipulative experiment to quantify the impact of small mammal herbivores on the belowground biogeochemistry of the tundra at three sites near Toolik Lake, Alaska. At each site we set up grazing fences in July of 2018 to simulate different levels of small mammal herbivore (vole and lemming) activity. Each site had 3 treatment plots and a control plot: 1)Exclosure treatments (EX) were 8 meter (m) x 8m square mesh fences 2) control plots (CT) were 8m x 8m unfenced plots marked with pin flags at corners 3) press treatments (PR) were 20m x 20m square mesh fences stocked with 4 tundra voles (Microtus oeconomus) every summer except for 2024 and 4) pulse treatments (PU) where we stocked the fence with 4 voles in 2018 and then removed and excluded voles from 2019 onward. At each site we collected temperature measurements using iButton data loggers from the soil surface, the soil organic layer, and the soil mineral layer every 4 hours from 2018 - 2024. iButton loggers were removed and replaced after soil thaw every summer. 

We conducted a manipulative experiment to quantify the impact of small mammal herbivores on the belowground biogeochemistry of the tundra at three sites near Toolik Lake, Alaska. At each site we set up grazing fences in July of 2018 to simulate different levels of small mammal herbivore (vole and lemming) activity. Each site had 3 treatment plots and a control plot: 1)Exclosure treatments (EX) were 8 meter (m) x 8m square mesh fences 2) control plots (CT) were 8m x 8m unfenced plots marked with pin flags at corners 3) press treatments (PR) were 20m x 20m square mesh fences stocked with 4 tundra voles (Microtus oeconomus) every summer and 4) pulse treatments (PU) where we stocked the fence with 4 voles in 2018 and then removed and excluded voles from 2019-2022. At each site we collected 10 thaw depth measurements along a transect from each treatment. 

Abstract 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. 

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. 

Abstract Whole‐ecosystem interactions and feedbacks constrain ecosystem responses to environmental change. The effects of these constraints on responses to climate trends and extreme weather events have been well studied. Here we examine how these constraints respond to changes in day‐to‐day weather variability without changing the long‐term mean weather. Although environmental variability is recognized as a critical factor affecting ecological function, the effects of climate change on day‐to‐day weather variability and the resultant impacts on ecosystem function are still poorly understood. Changes in weather variability can alter the mean rates of individual ecological processes because many processes respond non‐linearly to environmental drivers. We assessed how these individual‐process responses to changes in day‐to‐day weather variability interact with one another at an ecosystem level. We examine responses of arctic tundra to changes in weather variability using stochastic simulations of daily temperature, precipitation, and light to drive a biogeochemical model. Changes in weather variability altered ecosystem carbon, nitrogen, and phosphorus stocks and cycling rates in our model. However, responses of some processes (e.g., respiration) were inconsistent with expectations because ecosystem feedbacks can moderate, or even reverse, direct process responses to weather variability. More weather variability led to greater carbon losses from land to atmosphere; less variability led to higher carbon sequestration on land. The magnitude of modeled ecosystem response to weather variability was comparable to that predicted for the effects of climate mean trends by the end of the century. 

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. 

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.