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Abstract AimsHerbivores create large differences in litter decomposition rates, but identifying how they do this can be difficult because they simultaneously influence both biotic and abiotic factors. In the Yukon-Kuskokwim (Y-K) River Delta in western Alaska, geese are dominant herbivores in wet-sedge meadows, where they create ‘grazing lawns’ that have nutrient-rich litter and an open habitat structure. To understand how geese affect decomposition, we tested the effects of litter quality and habitat type on litter decomposition over one year. MethodsWe performed a litter bag study in which we collected two litter types representing grazed and ungrazed vegetation conditions (high quality litter similar to grazed litter, and lower quality senesced, ungrazed litter), then incubated them in ‘grazing lawn’ and ungrazed meadows. Litter mass loss, carbon, nitrogen, cellulose and lignin content were measured after 3, 6, 9, and 52 weeks. We also monitored abiotic conditions (i.e., soil temperature, UV radiation, throughfall, and soil moisture content) in each habitat type. ResultsHigh-quality litter (lower lignin:N ratios) lost more mass than low-quality ungrazed litter over the whole study. However, at different times during the decomposition process, lower quality litter decomposed faster in grazed habitat, whereas higher quality litter decomposed faster in ungrazed habitat. This occurred despite abiotic conditions in grazed habitat that generally promote faster decomposition. ConclusionResults suggest that herbivore-induced increases in litter quality increase decomposition rates, and that the accumulation of the low-quality litter in ungrazed habitats is partly due to slow decomposition rates. While herbivores influence habitat conditions, the effects of habitat on decomposition differed across litter qualities, which suggests that other variables, such as differing microbial communities, play a role in decomposition processes. Graphical abstract 

Abstract High latitude wetlands are ecologically important ecosystems due to their large carbon (C) storage capacity and because they serve as breeding and nesting habitat for large populations of migratory birds. Goose herbivory in wetland meadows affects leaf chemical and morphological traits and also influences soil properties by increasing soil temperature and depositing faeces. Grazing‐induced changes to above‐ground traits and soil properties impact C cycling, but the influence of grazing on root‐mediated C and nitrogen (N) cycling has not been explored.We investigated how goose herbivory in a low‐Arctic coastal wetland in western Alaska affected root morphological, physiological and chemical traits of a dominant graminoid by assessing plant traits in ungrazed versus heavily grazed sedge meadows. We also performed a 11‐week lab‐based root incubation experiment to determine how grazing affects CO₂‐C efflux, the size and decay rate of the fast‐cycling C pool (i.e. C with a mean residence time of days to weeks, determined via CO₂‐C efflux), and patterns of N mineralization during root decomposition.Goose grazing altered root chemical traits by increasing root N by 7%, cellulose by 12%, and ash content by 17%, indicating that grazing shifted root chemical traits towards a resource‐acquisition strategy. Grazing did not alter root biomass, morphology or bulk C exudation. In our root incubation, soils that included the roots of grazed plants tended to exhibit greater CO₂‐C efflux than those containing ungrazed plant roots due to a larger fast‐cycling C pool. Additionally, grazing‐induced increases in soil temperature led to greater CO₂‐C efflux due to a faster decay rate of the fast‐cycling C pool. Finally, compared with ungrazed roots, we found that the decomposition of grazed roots resulted in more N being transferred to root necromass from the surrounding soil, suggesting that microbial communities decomposing grazed roots immobilized N.Synthesis. Overall, our results indicate that goose grazing increased C‐cycling rates by influencing soil environmental conditions and by altering the ecological strategy of grazed plants. In contrast, grazing decreased net N mineralization by promoting N immobilization. These results suggest that changing patterns and abundances of herbivores can have substantial effects on elemental cycles. 

Abstract QuestionUnderstanding the sensitivity and magnitude of plant community responses in tundra wetlands to herbivory and warming is pressing as these ecosystems are increasingly threatened by changes in grazing pressure and higher temperatures. Here, we ask to what extent different low‐Arctic coastal wetland plant communities are affected by short‐term goose grazing and warming, and whether these communities differ in their responses. LocationYukon–Kuskokwim Delta, Alaska. MethodsWe conducted an experiment where we simulated goose grazing by clipping the vegetation and summer warming by using open‐top chambers in three plant communities along a 6‐km coastal–inland gradient. We assessed plant community compositional changes following two years of treatments. ResultsGrazing had stronger effects than warming on both plant functional group and species composition. Overall, grazing decreased the abundance of grasses and sedges and increased the abundance of forbs, whereas warming only caused a decrease in forb abundance. However, plant communities and functional groups, both within and across communities, varied widely in their responses to treatments. Grazing decreased grass abundance (−25%) and increased forb abundance (+44%) in the two more coastal communities, and reduced sedge abundance (−22%) only in the most inland community. Warming only decreased forb abundance (−18%) in the most coastal community, which overall was the most responsive to treatments. ConclusionsWe show that short‐term goose grazing predominates over short‐term summer warming in eliciting compositional changes in three different low‐Arctic coastal wetland plant communities. Yet, responses varied among communities and the same functional groups could respond differently across them, highlighting the importance of investigating the effects of biotic and abiotic drivers in different contexts. By showing that tundra wetland plant communities can differ in their immediate sensitivity to goose grazing and, though to a lesser extent, warming, our findings have implications for the functioning of these rapidly changing high‐latitude ecosystems. 

Abstract Global change drivers that modify the quality and quantity of litter inputs to soil affect greenhouse gas fluxes, and thereby constitute a feedback to climate change. Carbon cycling in the Yukon–Kuskokwim (Y–K) River Delta, a subarctic wetland system, is influenced by landscape variations in litter quality and quantity generated by herbivores (migratory birds) that create ‘grazing lawns’ of short stature, nitrogen-rich vegetation. To identify the mechanisms by which these changes in litter inputs affect soil carbon balance, we independently manipulated qualities and quantities of litter representative of levels found in the Y–K Delta in a fully factorial microcosm experiment. We measured CO₂fluxes from these microcosms weekly. To help us identify how litter inputs influenced greenhouse gas fluxes, we sequenced soil fungal and bacterial communities, and measured soil microbial biomass carbon, dissolved carbon, inorganic nitrogen, and enzyme activity. We found that positive correlations between litter input quantity and CO₂flux were dependent upon litter type, due to differences in litter stoichiometry and changes to the structure of decomposer communities, especially the soil fungi. These community shifts were particularly pronounced when litter was added in the form of herbivore feces, and in litter input treatments that induced nitrogen limitation (i.e., senesced litter). The sensitivity of carbon cycling to litter quality and quantity in this system demonstrates that herbivores can strongly impact greenhouse gas fluxes through their influence on plant growth and tissue chemistry. Graphical abstract 

Abstract Understanding environmental drivers of species diversity has become increasingly important under climate change. Different trophic groups (predators, omnivores and herbivores) interact with their environments in fundamentally different ways and may therefore be influenced by different environmental drivers. Using random forest models, we identified drivers of terrestrial mammals' total and proportional species richness within trophic groups at a global scale. Precipitation seasonality was the most important predictor of richness for all trophic groups. Richness peaked at intermediate precipitation seasonality, indicating that moderate levels of environmental heterogeneity promote mammal richness. Gross primary production (GPP) was the most important correlate of the relative contribution of each trophic group to total species richness. The strong relationship with GPP demonstrates that basal‐level resource availability influences how diversity is structured among trophic groups. Our findings suggest that environmental characteristics that influence resource temporal variability and abundance are important predictors of terrestrial mammal richness at a global scale. 

In the Yukon-Kuskokwim (YK) Delta, geese create grazing lawns in Carex subspathacea meadows. Geese annually maintain the grazing lawns, resulting in different aboveground morphological expressions for grazed Carex subspathacea compared to ungrazed Carex subspathacea. Grazed C. subspathacea tends to grow to an average of 1.5 centimeters (cm) in height and has a floret growth form, while ungrazed C. subspathacea reaches an average height of nearly 15.5 cm. Additionally, grazed C. subspathacea has lower Carbon : Nitrogen (C:N) content than ungrazed C. subspathacea. Furthermore, both the physical alterations to Carex subspathacea and the changes to the soil physiochemical environment caused by grazing suggest that aboveground herbivory may affect root trait expression of C. subspathacea, which in turn may influence biogeochemical processes such as soil respiration and decomposition rates. This data set contains information on Carex subspathacea root traits, including root morphology (total length, surface area, and volume), root exudates (dissolved organic carbon concentration), and root chemistry (carbon, nitrogen, phosphorus, lignin, cellulose, and acid fiber detergent) collected in Western Alaska's Yukon-Kuskokwim's delta. The samples were collected from two Carex subspathacea habitat types (grazed or ungrazed). 

Global change drivers that modify the quality and quantity of litter inputs to soil affect greenhouse gas fluxes, and thereby constitute a feedback to climate change. Carbon cycling in the Yukon-Kuskokwim (Y-K) River Delta, a subarctic wetland system, is influenced by landscape variations in litter quality and quantity generated by herbivores (migratory birds) that create ‘grazing lawns’ of short stature, nitrogen-rich vegetation. To identify the mechanisms by which these changes in litter inputs affect soil carbon balance, we independently manipulated qualities and quantities of litter representative of levels found in the Y-K Delta in a fully factorial microcosm experiment. We measured carbon dioxide (CO2) fluxes from these microcosms weekly. To help us identify how litter inputs influenced greenhouse gas fluxes, we sequenced soil fungal and bacterial communities, and measured soil microbial biomass carbon, dissolved carbon, inorganic nitrogen, and enzyme activity. We found that positive correlations between litter input quantity and CO2 flux were dependent upon litter type, due to differences in litter stoichiometry and changes to the structure of decomposer communities, especially the soil fungi. These community shifts were particularly pronounced when litter was added in the form of herbivore feces, and in litter input treatments that induced nitrogen limitation (i.e., senesced litter). The sensitivity of carbon cycling to litter quality and quantity in this system demonstrates that herbivores can strongly impact greenhouse gas fluxes through their influence on plant growth and tissue chemistry. 

Vertebrate herbivore excrement is thought to influence nutrient cycling, plant nutrition, and growth; however, its importance is rarely isolated from other aspects of herbivory, such as trampling and leaf removal, leaving questions about the extent to which herbivore effects are due to feces. We hypothesized that as a source of additional nutrients, feces would directly increase soil N concentrations and N2O emission, alleviate plant, and microbial nutrient limitations, resulting in increased plant growth and foliar quality, and increase CH4 emissions. We tested these hypotheses using a field experiment in coastal western Alaska,USA, where we manipulated goose feces such that naturally grazed areas received three treatments:feces removal, ambient amounts of feces, or double ambient amounts of feces. Doubling feces marginally increased NH4 +-N in soil water, whereas both doubled feces and feces removal significantly increased NO3--N; N2O flux was also higher in removal plots. Feces removal marginally reduced root biomass and significantly reduced productivity (that is, GPP) in the second year, measured as greater CO2 emissions. Doubling feces marginally increased foliar chemical quality by increasing %N and decreasing C:N. Treatments did not influence CH4 flux. In short, feces removal created sites poorer in nutrients, with reduced root growth, graminoid nutrient uptake, and productivity. While goose feces alone did not create dramatic changes in nutrient cycling in western Alaska, they do appear to be an important source of nutrients for grazed areas and to contribute to greenhouse gas exchange as their removal increased emissions of CO2 and N2O to the atmosphere. 

Herbivory can have strong impacts on greenhouse gas fluxes in high-latitude ecosystems. For example, in the Yukon-Kuskokwim (Y-K) Delta in western Alaska, migratory goose grazing affects the magnitude of soil carbon dioxide (CO2) and methane (CH4) fluxes. However, the underlying drivers of this relationship are unclear, as few studies systematically tease apart the processes by which herbivores influences soil biogeochemistry. To examine these mechanisms in detail, we conducted a laboratory incubation experiment to quantify changes in greenhouse gas fluxes in response to three parameters altered by herbivores in situ: temperature, soil moisture content, and nutrient inputs. These treatments were applied to soils collected in grazing lawns and nearby ungrazed habitat, allowing us to assess how variation in microbial community structure influenced observed responses. We found pronounced differences in both fungal and prokaryotic community composition between grazed and ungrazed areas. In the laboratory incubation experiment, CO2 and CH4 fluxes increased with temperature, soil moisture, and goose fecal addition, suggesting that grazing-related changes in the soil abiotic environment may enhance soil C losses. Yet, these abiotic drivers were insufficient to explain variation in fluxes between soils with and without prior grazing. Differences in trace gas fluxes between grazed and ungrazed areas may result both from herbivore-induced shifts in abiotic parameters and grazing-related alterations in microbial community structure. Our findings suggest that relationships among herbivores and soil microbial communities could mediate carbon-climate feedbacks in rapidly changing high-latitude ecosystems.