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Landscapes are undergoing ecological changes, and how organisms interact with changing habitats has implications for zoogeochemical influences on ecosystem function (processes and properties). This may be especially true for organisms that alter nutrient cycling, such as structure builders, in nutrient‐limited systems such as the Arctic. Our aims were to examine the impact of brown lemming (Lemmus trimucronatus) structures (winter nests, runways, latrines, and burrows) on above‐ and below‐ground processes in habitats that represent contemporary (flat‐centered polygon) and potential future (high‐centered polygon) tundra conditions near Utqiaġvik, Alaska. Above‐ground, structures influenced vegetation community structure at local scales, with winter nests having lower moss and forb cover and burrows, latrines, and runways having lower litter and higher bare ground cover; although effects did not vary by habitat type. Furthermore, structures influenced plant diversity, which may be driven by structure types supporting unique plant species. Below‐ground, burrows had broad effects on soil nutrients, with soils under burrows generally having lower carbon and nitrogen contents and exo‐enzyme activities in both habitat types. Other structures increased nutrient availability, with soils under winter nests having higher ammonium concentrations compared to controls in both habitat types, whereas soils under latrines had higher phosphate and extractable organic carbon than controls, but only in high‐centered polygon tundra. Additionally, soil temperatures under winter nests were lower than at control sites, but only in flat‐centered polygon tundra, and soil pH was higher under winter nests and runways in both habitat types. Effects of structures on soil physical properties probably helped to regulate the effects of structures on soil nutrient availability. Finally, differences in structure effects between habitat types suggest that as high‐centered polygonal tundra becomes more prevalent, this herbivore's influence on ecosystem processes at local scales may feedback to alter the function of future arctic ecosystems. We examined the effects of structure builders on biogeochemical cycling under future arctic tundra conditions. We found that structures influence nutrient availability and vegetation communities differently between contemporary and future conditions. These effects suggest that structure builder effects at local scales will feedback to alter the function of future arctic ecosystems.

Rapid arctic vegetation change as a result of global warming includes an increase in the cover and biomass of deciduous shrubs. Increases in shrub abundance will result in a proportional increase of shrub litter in the litter community, potentially affecting carbon turnover rates in arctic ecosystems. We investigated the effects of leaf and root litter of a deciduous shrub, Betula nana, on decomposition, by examining species-specific decomposition patterns, as well as effects of Betula litter on the decomposition of other species. We conducted a 2-yr decomposition experiment in moist acidic tundra in northern Alaska, where we decomposed three tundra species (Vaccinium vitis-idaea, Rhododendron palustre, and Eriophorum vaginatum) alone and in combination with Betula litter. Decomposition patterns for leaf and root litter were determined using three different measures of decomposition (mass loss, respiration, extracellular enzyme activity). We report faster decomposition of Betula leaf litter compared to other species, with support for species differences coming from all three measures of decomposition. Mixing effects were less consistent among the measures, with negative mixing effects shown only for mass loss. In contrast, there were few species differences or mixing effects for root decomposition. Overall, we attribute longer-term litter mass loss patterns to patterns created by early decomposition processes in the first winter. We note numerous differences for species patterns between leaf and root decomposition, indicating that conclusions from leaf litter experiments should not be extrapolated to below-ground decomposition. The high decomposition rates of Betula leaf litter aboveground, and relatively similar decomposition rates of multiple species below, suggest a potential for increases in turnover in the fast-decomposing carbon pool of leaves and fine roots as the dominance of deciduous shrubs in the Arctic increases, but this outcome may be tempered by negative litter mixing effects during the early stages of encroachment.

1. Ecosystem properties may be determined by the number of different species or groups of species in a community, the identity of those groups, and their relative abundance. The mass ratio theory predicts that the effect of species or groups of species on ecosystem properties will be dependent on their proportional abundance in a community. 2. Single plant functional groups (graminoids, legumes, non-leguminous forbs) were removed from a natural grassland in northern Canada to examine the role of group identity in determining both ecosystem properties and biomass compensation by remaining species. Removals were conducted across two different environmental treatments (fertilization and fungicide) to examine the context dependency of functional group identity effects. 3. The degree of biomass compensation in the first 4 years after removal was influenced by the identity of the functional group removed and also of those remaining. When graminoids were removed, none of the remaining functional groups compensated for the loss of biomass. Graminoids partially compensated for the removal of forbs or legumes, with the degree of compensation depending on environmental treatments. 4. Light interception, soil moisture and soil nutrients were all influenced by functional group identity, with graminoids having a greater impact than expected based on their biomass contribution to the community. Legumes, in contrast, had very little effect on any of the ecosystem properties measured. 5. For most ecosystem properties measured, the role of plant functional groups was not context dependent; functional groups had the same effect on ecosystem properties regardless of fertilization or fungicide treatments. 6. Synthesis. We have shown that the effects of losing a functional group do not solely depend on the group's dominance. In this northern grassland, there are greater effects of losing graminoids than one would predict based on their biomass contributions to the community, and functional group identity plays a critical role in determining the effects of diversity loss.

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 PO4 in 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.

Global change in the Arctic promotes deeper soil thaw and enhanced soil microbial activity, increasing nitrogen (N) and phosphorus (P) availability to plants and microbes in strongly nutrient‐limited ecosystems. This critical, positive climate feedback has been examined through fertilization experiments that describe short‐term (<10 yr) above‐ or belowground responses to combined NP additions, with evidence of enhanced shrub growth, nutrient availability, and soil organic matter decomposition. There has been less opportunity for long‐term comparisons of both above‐ and belowground responses with factorial N and P additions in different systems, despite broad awareness that ecosystem response can shift with time, and the potential for decoupled above‐ vs. belowground or N vs. P responses, currently and with further predicted global change. We examined the response of the plants, soil microbes, and soil nutrients, to factorial N and P additions in the moist acidic tundra (MAT; 26 yr of nutrient additions) and moist non‐acidic tundra (MNT; 16 yr). Aboveground, the MAT plant community continues to change as predicted by earlier studies: Functional groups responded independently to N and P, but NDVI‐biomass, especially of Betula nana, only increased with N addition. Unlike shorter‐term MNT studies, the MNT vegetation, which does not include B. nana, shows few new fertilization responses. Belowground responses were not predicted by aboveground responses in either MAT or MNT. In contrast to the N response aboveground, MAT microbial biomass responded positively and microbial phosphatase activity negatively to P additions, implying possible release from microbial P limitation. Critically, earlier published results of declines in soil total carbon (C) with combined NP addition in the MAT are not present in the long term. We make two conclusions: (1) Arctic ecosystems are not universally N‐limited but also exhibit complex responses to P alone or in combination with N; and (2) the presence or absence of key vegetation species can cascade from aboveground to belowground and restrict the extrapolation of responses of nutrient addition in a single arctic ecosystem to other arctic ecosystems, the short‐term to the long term, or aboveground to belowground.

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.

Herbivores can play an important role in determining arctic ecosystem function with effects determined in part by herbivore identity. We examined the impact of long-term (twenty-two years) small and large mammal herbivore exclusion in two arctic plant communities in northern Alaska: dry heath (DH) and moist acidic tundra (MAT). Our aims were to examine how herbivore exclusion influences (1) plant communities and (2) soil nutrient pools and microbial processes. Though herbivore absence increased moss and decreased evergreen shrub cover in MAT, there were few other significant effects on vegetation in either community. We also observed no influence of exclusion on most soil properties. However, in DH, phosphatase activity was greater in areas where small mammals alone were present, suggesting that they are altering phosphorus (P) availability, perhaps through herbivores' influence on the plant community and subsequently on competition for P with the microbial community. We conclude that herbivore impacts in the Arctic are dependent on both the plant community and herbivore identity (size). We show the importance of understanding the roles of herbivores in the Arctic and contribute to a growing number of herbivore studies in a biome likely to experience future changes in herbivore communities and ecosystem function.

Retrogressive thaw slumps (RTS)—thermal erosion of soil and vegetation after ground ice thaw—are increasing. Recovery of plant biomass after RTS is important for maintaining Arctic carbon (C) stocks and is regulated by nutrient availability for new plant growth. Many RTS are characterized by verdant shrub growth mid-succession, atypical of the surrounding nutrient-limited tundra. Here, we investigated the potential for internal and external sources of nitrogen (N) and phosphorus (P) to support mid-successional shrub growth at three Alaskan RTS chronosequences. We assessed patterns of soil and microbial CNP, soil NP cycling rates and stocks, N inputs via biological N 2 -fixation, and thaw leachate over time after disturbance. We found a clear transfer of P stocks from mineral to organic soils with increasing site age, yet insufficient N from any one source to support observed shrub growth. Instead, multiple mechanisms may have contributed to mid-successional shrub growth, including sustained N-cycling with reduced plant biomass, N leaching from undisturbed tundra, uninvestigated sources of N 2 -fixation, and most promising given the large resource, deep mineral soil N stocks. These potential mechanisms of N supply are critical for the regulation of the Arctic C cycle in response to an increasingly common climate-driven disturbance.

Plant litter decomposition is a critical ecosystem process representing a major pathway for carbon flux, but little is known about how it is affected by changes in plant composition and diversity. Single plant functional groups (graminoids, legumes, non-leguminous forbs) were removed from a grassland in northern Canada to examine the impacts of functional group identity on decomposition. Removals were conducted within two different environmental contexts (fertilization and fungicide application) to examine the context-dependency of these identity effects. We examined two different mechanisms by which the loss of plant functional groups may impact decomposition: effects of the living plant community on the decomposition microenvironment, and changes in the species composition of the decomposing litter, as well as the interaction between these mechanisms. We show that the identity of the plant functional group removed affects decomposition through both mechanisms. Removal of both graminoids and forbs slowed decomposition through changes in the decomposition microenvironment. We found non-additive effects of litter mixing, with both the direction and identity of the functional group responsible depending on year; in 2004 graminoids positively influenced decomposition whereas in 2006 forbs negatively influenced decomposition rate. Although these two mechanisms act independently, their effects may be additive if both mechanisms are considered simultaneously. It is essential to understand the variety of mechanisms through which even a single ecosystem property is affected if we are to predict the future consequences of biodiversity loss.

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.