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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. 

Understanding 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.

Yukon–Kuskokwim Delta, Alaska.

We 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.

Grazing 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.

We 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.

 

Rapid warming in northern ecosystems over the past four decades has resulted in earlier spring, increased precipitation, and altered timing of plant–animal interactions, such as herbivory. Advanced spring phenology can lead to longer growing seasons and increased carbon (C) uptake. Greater precipitation coincides with greater cloud cover possibly suppressing photosynthesis. Timing of herbivory relative to spring phenology influences plant biomass. None of these changes are mutually exclusive and their interactions could lead to unexpected consequences for Arctic ecosystem function. We examined the influence of advanced spring phenology, cloud cover, and timing of grazing on C exchange in the Yukon–Kuskokwim Delta of western Alaska for three years. We combined advancement of the growing season using passive-warming open-top chambers (OTC) with controlled timing of goose grazing (early, typical, and late season) and removal of grazing. We also monitored natural variation in incident sunlight to examine the C exchange consequences of these interacting forcings. We monitored net ecosystem exchange of C (NEE) hourly using an autochamber system. Data were used to construct daily light curves for each experimental plot and sunlight data coupled with a clear-sky model was used to quantify daily and seasonal NEE over a range of incident sunlight conditions. Cloudy days resulted in the largest suppression of NEE, reducing C uptake by approximately 2 g C m⁻²d⁻¹regardless of the timing of the season or timing of grazing. Delaying grazing enhanced C uptake by approximately 3 g C m⁻²d⁻¹. Advancing spring phenology reduced C uptake by approximately 1.5 g C m⁻²d⁻¹, but only when plots were directly warmed by the OTCs; spring advancement did not have a long-term influence on NEE. Consequently, the two strongest drivers of NEE, cloud cover and grazing, can have opposing effects and thus future growing season NEE will depend on the magnitude of change in timing of grazing and incident sunlight.

 

Climate change is creating phenological mismatches between herbivores and their plant resources throughout the Arctic. While advancing growing seasons and changing arrival times of migratory herbivores can have consequences for herbivores and forage quality, developing mismatches could also influence other traits of plants, such as above‐ and below‐ground biomass and the type of reproduction, that are often not investigated.

In coastal western Alaska, we conducted a 3‐year factorial experiment that simulated scenarios of phenological mismatch by manipulating the start of the growing season (3 weeks early and ambient) and grazing times (3 weeks early, typical, 3 weeks late, or no‐grazing) of Pacific black brant (Branta bernicla nigricans), to examine how the timing of these events influence a primary goose forage species,Carex subspathacea.

After 3 years, an advanced growing season compared to a typical growing season increased stem heights, standing dead biomass, and the number of inflorescences. Early season grazing compared to typical season grazing reduced above‐ and below‐ground biomass, stem height, and the number of tillers; while late season grazing increased the number of inflorescences and standing dead biomass. Therefore, an advanced growing season and late grazing had similar directional effects on most plant traits, but a 3‐week delay in grazing had an impact on traits 3–5 times greater than a similarly timed shift in the advancement of spring. In addition, changes in response to treatments for some variables, such as the number of inflorescences, were not measurable until the second year of the experiment, while other variables, such as root productivity and number of tillers, changed the direction of their responses to treatments over time.

Synthesis. Factors affecting the timing of migration have a larger influence than earlier springs on an important forage species in the breeding and rearing habitats of Pacific black brant. The phenological mismatch prediction for this site of earlier springs and later goose arrival will likely increase above‐ and below‐ground biomass and sexual reproduction of the often‐clonally reproducingC. subspathacea. Finally, the implications of mismatch may be difficult to predict because some variables required successive years of mismatch to respond.

 

The advancement of spring and the differential ability of organisms to respond to changes in plant phenology may lead to “phenological mismatches” as a result of climate change. One potential for considerable mismatch is between migratory birds and food availability in northern breeding ranges, and these mismatches may have consequences for ecosystem function. We conducted a three‐year experiment to examine the consequences for CO₂exchange of advanced spring green‐up and altered timing of grazing by migratory Pacific black brant in a coastal wetland in western Alaska. Experimental treatments represent the variation in green‐up and timing of peak grazing intensity that currently exists in the system. Delayed grazing resulted in greater net ecosystem exchange (NEE) and gross primary productivity (GPP), while early grazing reduced CO₂uptake with the potential of causing net ecosystem carbon (C) loss in late spring and early summer. Conversely, advancing the growing season only influenced ecosystem respiration (ER), resulting in a small increase in ER with no concomitant impact on GPP or NEE. The experimental treatment that represents the most likely future, with green‐up advancing more rapidly than arrival of migratory geese, results in NEE changing by 1.2 µmol m⁻² s⁻¹toward a greater CO₂sink in spring and summer. Increased sink strength, however, may be mitigated by early arrival of migratory geese, which would reduce CO₂uptake. Importantly, while the direct effect of climate warming on phenology of green‐up has a minimal influence on NEE, the indirect effect of climate warming manifest through changes in the timing of peak grazing can have a significant impact on C balance in northern coastal wetlands. Furthermore, processes influencing the timing of goose migration in the winter range can significantly influence ecosystem function in summer habitats.