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1. Herbivory changes soil microbial communities and greenhouse gas fluxes in a high-latitude wetland

   https://doi.org/10.1007/s00248-021-01733-8  

   Foley, Karen M. ; Beard, Karen H. ; Atwood, Trisha B. ; Waring, Bonnie G. ( March 2021 , Microbial Ecology)

   null (Ed.)

   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. 

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2. Nutrient availability controls the impact of mammalian herbivores on soil carbon and nitrogen pools in grasslands

   https://doi.org/10.1111/gcb.15023  

   Sitters, Judith ; Wubs, E. R. Jasper ; Bakker, Elisabeth S. ; Crowther, Thomas W. ; Adler, Peter B. ; Bagchi, Sumanta ; Bakker, Jonathan D. ; Biederman, Lori ; Borer, Elizabeth T. ; Cleland, Elsa E. ; et al ( February 2020 , Global Change Biology)

   Grasslands are subject to considerable alteration due to human activities globally, including widespread changes in populations and composition of large mammalian herbivores and elevated supply of nutrients. Grassland soils remain important reservoirs of carbon (C) and nitrogen (N). Herbivores may affect both C and N pools and these changes likely interact with increases in soil nutrient availability. Given the scale of grassland soil fluxes, such changes can have striking consequences for atmospheric C concentrations and the climate. Here, we use the Nutrient Network experiment to examine the responses of soil C and N pools to mammalian herbivore exclusion across 22 grasslands, under ambient and elevated nutrient availabilities (fertilized with NPK + micronutrients). We show that the impact of herbivore exclusion on soil C and N pools depends on fertilization. Under ambient nutrient conditions, we observed no effect of herbivore exclusion, but under elevated nutrient supply, pools are smaller upon herbivore exclusion. The highest mean soil C and N pools were found in grazed and fertilized plots. The decrease in soil C and N upon herbivore exclusion in combination with fertilization correlated with a decrease in aboveground plant biomass and microbial activity, indicating a reduced storage of organic matter and microbial residues as soil C and N. The response of soil C and N pools to herbivore exclusion was contingent on temperature – herbivores likely cause losses of C and N in colder sites and increases in warmer sites. Additionally, grasslands that contain mammalian herbivores have the potential to sequester more N under increased temperature variability and nutrient enrichment than ungrazed grasslands. Our study highlights the importance of conserving mammalian herbivore populations in grasslands worldwide. We need to incorporate local‐scale herbivory, and its interaction with nutrient enrichment and climate, within global‐scale models to better predict land–atmosphere interactions under future climate change.

    

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3. Long‐term herbivore removal experiments reveal how geese and reindeer shape vegetation and ecosystem CO2 ‐fluxes in high‐ Arctic tundra

   https://doi.org/10.1111/1365-2745.14200  

   Petit Bon, Matteo ; Hansen, Brage B. ; Loonen, Maarten J. J. E. ; Petraglia, Alessandro ; Bråthen, Kari Anne ; Böhner, Hanna ; Layton‐Matthews, Kate ; Beard, Karen H. ; Le Moullec, Mathilde ; Jónsdóttir, Ingibjörg S. ; et al ( September 2023 , Journal of Ecology)

   Given the current rates of climate change, with associated shifts in herbivore population densities, understanding the role of different herbivores in ecosystem functioning is critical for predicting ecosystem responses. Here, we examined how migratory geese and resident, non‐migratory reindeer—two dominating yet functionally contrasting herbivores—control vegetation and ecosystem processes in rapidly warming Arctic tundra.

   We collected vegetation and ecosystem carbon (C) flux data at peak plant growing season in the two longest running, fully replicated herbivore removal experiments found in high‐Arctic Svalbard. Experiments had been set up independently in wet habitat utilised by barnacle geeseBranta leucopsisin summer and in moist‐to‐dry habitat utilised by wild reindeerRangifer tarandus platyrhynchusyear‐round.

   Excluding geese induced vegetation state transitions from heavily grazed, moss‐dominated (only 4 g m⁻²of live above‐ground vascular plant biomass) to ungrazed, graminoid‐dominated (60 g m⁻²after 4‐year exclusion) and horsetail‐dominated (150 g m⁻²after 15‐year exclusion) tundra. This caused large increases in vegetation C and nitrogen (N) pools, dead biomass and moss‐layer depth. Alterations in plant N concentration and CN ratio suggest overall slower plant community nutrient dynamics in the short‐term (4‐year) absence of geese. Long‐term (15‐year) goose removal quadrupled net ecosystem C sequestration (NEE) by increasing ecosystem photosynthesis more than ecosystem respiration (ER).

   Excluding reindeer for 21 years also produced detectable increases in live above‐ground vascular plant biomass (from 50 to 80 g m⁻²; without promoting vegetation state shifts), as well as in vegetation C and N pools, dead biomass, moss‐layer depth and ER. Yet, reindeer removal did not alter the chemistry of plants and soil or NEE.

   Synthesis. Although both herbivores were key drivers of ecosystem structure and function, the control exerted by geese in their main habitat (wet tundra) was much more pronounced than that exerted by reindeer in their main habitat (moist‐to‐dry tundra). Importantly, these herbivore effects are scale dependent, because geese are more spatially concentrated and thereby affect a smaller portion of the tundra landscape compared to reindeer. Our results highlight the substantial heterogeneity in how herbivores shape tundra vegetation and ecosystem processes, with implications for ongoing environmental change.

    

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4. Hubbard Brook Experimental Forest: Soil Freezing Study (SFS) In Situ Measurements of Snow and Soil Frost Depth

   https://doi.org/10.6073/pasta/207b884c7510aca0aa9bf2c5ed93e432  

   Templer, Pamela H ; Socci, Anne ; Campbell, John L ( January 2021 , Environmental Data Initiative)

   Climate models for the northeastern United States (U.S.) over the next century predict an increase in air temperature between 2.8 and 4.3 °C and a decrease in the average number of days per year when a snowpack will cover the forest floor (Hayhoe et al. 2007, 2008; Campbell et al. 2010). Studies of forest dynamics in seasonally snow-covered ecosystems have been primarily conducted during the growing season, when most biological activity occurs. However, in recent years considerable progress has been made in our understanding of how winter climate change influences dynamics in these forests. The snowpack insulates soil from below-freezing air temperatures, which facilitates a significant amount of microbial activity. However, a smaller snowpack and increased depth and duration of soil frost amplify losses of dissolved organic C and NO3- in leachate, as well as N2O released into the atmosphere. The increase in nutrient loss following increased soil frost cannot be explained by changes in microbial activity alone. More likely, it is caused by a decrease in plant nutrient uptake following increases in soil frost. We conducted a snow-removal experiment at Hubbard Brook Experimental Forest to determine the effects of a smaller winter snowpack and greater depth and duration of soil frost on trees, soil microbes, and arthropods. A number of publications have been based on these data: Comerford et al. 2013, Reinmann et al. 2019, Templer 2012, and Templer et al. 2012. These data were gathered as part of the Hubbard Brook Ecosystem Study (HBES). The HBES is a collaborative effort at the Hubbard Brook Experimental Forest, which is operated and maintained by the USDA Forest Service, Northern Research Station. Campbell JL, Ollinger SV, Flerchinger GN, Wicklein H, Hayhoe K, Bailey AS. Past and projected future changes in snowpack and soil frost at the Hubbard Brook Experimental Forest, New Hampshire, USA. Hydrological Processes. 2010; 24:2465–2480. Comerford DP, PG Schaberg, PH Templer, AM Socci, JL Campbell, and KF Wallin. 2013. Influence of experimental snow removal on root and canopy physiology of sugar maple trees in a northern hardwood forest. Oecologia 171:261-269. Hayhoe K, Wake CP, Huntington TG, Luo LF, Schwartz MD, Sheffield J, et al. Past and future changes in climate and hydrological indicators in the US Northeast. Climate Dynamics. 2007; 28:381–407. Hayhoe, K., Wake, C., Anderson, B. et al. Regional climate change projections for the Northeast USA. Mitig Adapt Strateg Glob Change 13, 425–436 (2008). https://doi.org/10.1007/s11027-007-9133-2. Reinmann AB, J Susser, EMC Demaria, PH Templer. 2019. Declines in northern forest tree growth following snowpack decline and soil freezing.  Global Change Biology 25:420-430. Templer PH. 2012. Changes in winter climate: soil frost, root injury, and fungal communities (Invited). Plant and Soil 35: 15-17 Templer PH , AF Schiller, NW Fuller, AM Socci, JL Campbell, JE Drake, and TH Kunz. 2012. Impact of a reduced winter snowpack on litter arthropod abundance and diversity in a northern hardwood forest ecosystem. Biology and Fertility of Soils 48:413-424. 

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5. Evidence, causes, and consequences of declining nitrogen availability in terrestrial ecosystems

   https://doi.org/10.1126/science.abh3767  

   Mason, Rachel E. ; Craine, Joseph M. ; Lany, Nina K. ; Jonard, Mathieu ; Ollinger, Scott V. ; Groffman, Peter M. ; Fulweiler, Robinson W. ; Angerer, Jay ; Read, Quentin D. ; Reich, Peter B. ; et al ( April 2022 , Science)

   BACKGROUND The availability of nitrogen (N) to plants and microbes has a major influence on the structure and function of ecosystems. Because N is an essential component of plant proteins, low N availability constrains the growth of plants and herbivores. To increase N availability, humans apply large amounts of fertilizer to agricultural systems. Losses from these systems, combined with atmospheric deposition of fossil fuel combustion products, introduce copious quantities of reactive N into ecosystems. The negative consequences of these anthropogenic N inputs—such as ecosystem eutrophication and reductions in terrestrial and aquatic biodiversity—are well documented. Yet although N availability is increasing in many locations, reactive N inputs are not evenly distributed globally. Furthermore, experiments and theory also suggest that global change factors such as elevated atmospheric CO 2 , rising temperatures, and altered precipitation and disturbance regimes can reduce the availability of N to plants and microbes in many terrestrial ecosystems. This can occur through increases in biotic demand for N or reductions in its supply to organisms. Reductions in N availability can be observed via several metrics, including lowered nitrogen concentrations ([N]) and isotope ratios (δ 15 N) in plant tissue, reduced rates of N mineralization, and reduced terrestrial N export to aquatic systems. However, a comprehensive synthesis of N availability metrics, outside of experimental settings and capable of revealing large-scale trends, has not yet been carried out. ADVANCES A growing body of observations confirms that N availability is declining in many nonagricultural ecosystems worldwide. Studies have demonstrated declining wood δ 15 N in forests across the continental US, declining foliar [N] in European forests, declining foliar [N] and δ 15 N in North American grasslands, and declining [N] in pollen from the US and southern Canada. This evidence is consistent with observed global-scale declines in foliar δ 15 N and [N] since 1980. Long-term monitoring of soil-based N availability indicators in unmanipulated systems is rare. However, forest studies in the northeast US have demonstrated decades-long decreases in soil N cycling and N exports to air and water, even in the face of elevated atmospheric N deposition. Collectively, these studies suggest a sustained decline in N availability across a range of terrestrial ecosystems, dating at least as far back as the early 20th century. Elevated atmospheric CO 2 levels are likely a main driver of declines in N availability. Terrestrial plants are now uniformly exposed to ~50% more of this essential resource than they were just 150 years ago, and experimentally exposing plants to elevated CO 2 often reduces foliar [N] as well as plant-available soil N. In addition, globally-rising temperatures may raise soil N supply in some systems but may also increase N losses and lead to lower foliar [N]. Changes in other ecosystem drivers—such as local climate patterns, N deposition rates, and disturbance regimes—individually affect smaller areas but may have important cumulative effects on global N availability. OUTLOOK Given the importance of N to ecosystem functioning, a decline in available N is likely to have far-reaching consequences. Reduced N availability likely constrains the response of plants to elevated CO 2 and the ability of ecosystems to sequester carbon. Because herbivore growth and reproduction scale with protein intake, declining foliar [N] may be contributing to widely reported declines in insect populations and may be negatively affecting the growth of grazing livestock and herbivorous wild mammals. Spatial and temporal patterns in N availability are not yet fully understood, particularly outside of Europe and North America. Developments in remote sensing, accompanied by additional historical reconstructions of N availability from tree rings, herbarium specimens, and sediments, will show how N availability trajectories vary among ecosystems. Such assessment and monitoring efforts need to be complemented by further experimental and theoretical investigations into the causes of declining N availability, its implications for global carbon sequestration, and how its effects propagate through food webs. Responses will need to involve reducing N demand via lowering atmospheric CO 2 concentrations, and/or increasing N supply. Successfully mitigating and adapting to declining N availability will require a broader understanding that this phenomenon is occurring alongside the more widely recognized issue of anthropogenic eutrophication. Intercalibration of isotopic records from leaves, tree rings, and lake sediments suggests that N availability in many terrestrial ecosystems has steadily declined since the beginning of the industrial era. Reductions in N availability may affect many aspects of ecosystem functioning, including carbon sequestration and herbivore nutrition. Shaded areas indicate 80% prediction intervals; marker size is proportional to the number of measurements in each annual mean. Isotope data: (tree ring) K. K. McLauchlan et al. , Sci. Rep. 7 , 7856 (2017); (lake sediment) G. W. Holtgrieve et al. , Science 334 , 1545–1548 (2011); (foliar) J. M. Craine et al. , Nat. Ecol. Evol. 2 , 1735–1744 (2018) 

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