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1. Nutrients cause grassland biomass to outpace herbivory

   https://doi.org/10.1038/s41467-020-19870-y  

   Borer, E. T.; Harpole, W. S.; Adler, P. B.; Arnillas, C. A.; Bugalho, M. N.; Cadotte, M. W.; Caldeira, M. C.; Campana, S.; Dickman, C. R.; Dickson, T. L.; et al (December 2020, Nature Communications)

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   Abstract Human activities are transforming grassland biomass via changing climate, elemental nutrients, and herbivory. Theory predicts that food-limited herbivores will consume any additional biomass stimulated by nutrient inputs (‘consumer-controlled’). Alternatively, nutrient supply is predicted to increase biomass where herbivores alter community composition or are limited by factors other than food (‘resource-controlled’). Using an experiment replicated in 58 grasslands spanning six continents, we show that nutrient addition and vertebrate herbivore exclusion each caused sustained increases in aboveground live biomass over a decade, but consumer control was weak. However, at sites with high vertebrate grazing intensity or domestic livestock, herbivores consumed the additional fertilization-induced biomass, supporting the consumer-controlled prediction. Herbivores most effectively reduced the additional live biomass at sites with low precipitation or high ambient soil nitrogen. Overall, these experimental results suggest that grassland biomass will outstrip wild herbivore control as human activities increase elemental nutrient supply, with widespread consequences for grazing and fire risk. 

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2. Global impacts of fertilization and herbivore removal on soil net nitrogen mineralization are modulated by local climate and soil properties

   https://doi.org/10.1111/gcb.15308  

   Risch, Anita C.; Zimmermann, Stefan; Moser, Barbara; Schütz, Martin; Hagedorn, Frank; Firn, Jennifer; Fay, Philip A.; Adler, Peter B.; Biederman, Lori A.; Blair, John M.; et al (July 2020, Global Change Biology)

   Soil nitrogen (N) availability is critical for grassland functioning. However, human activities have increased the supply of biologically-limiting nutrients, and changed the density and identity of mammalian herbivores. These anthropogenic changes may alter net soil N mineralization (soil net Nmin), i.e., the net balance between N mineralization and immobilization, which could severely impact grassland structure and functioning. Yet, to date, little is known about how fertilization and herbivore removal individually, or jointly, affect soil net Nmin across a wide range of grasslands that vary in soil and climatic properties. Here, we collected data from 22 grasslands on five continents, all part of a globally replicated experiment, to assess how fertilization and herbivore removal affected potential (laboratory-based) and realized (field-based) soil net Nmin. Herbivore removal in the absence of fertilization did not alter potential and realized soil net Nmin. However, fertilization alone and in combination with herbivore removal consistently increased potential soil net Nmin. Realized soil net Nmin, in contrast, significantly decreased in fertilized plots where herbivores were removed. Treatment effects on potential and realized soil net Nmin were contingent on site-specific soil and climatic properties. Fertilization effects on potential soil net Nmin were larger at sites with higher mean annual precipitation (MAP) and temperature of the wettest quarter (T.q.wet). Reciprocally, realized soil net Nmin declined most strongly with fertilization and herbivore removal at sites with lower MAP and higher T.q.wet. In summary, our findings show that anthropogenic nutrient enrichment, herbivore exclusion, and alterations in future climatic conditions can negatively impact soil net Nmin across global grasslands under realistic field conditions. This is important context-dependent knowledge for grassland management worldwide. 

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3. Herbivory and nutrients shape grassland soil seed banks

   https://doi.org/10.1038/s41467-023-39677-x  

   Eskelinen, Anu; Jessen, Maria-Theresa; Bahamonde, Hector A.; Bakker, Jonathan D.; Borer, Elizabeth T.; Caldeira, Maria C.; Harpole, W. Stanley; Jia, Meiyu; Lannes, Luciola S.; Nogueira, Carla; et al (December 2023, Nature Communications)

   Abstract Anthropogenic nutrient enrichment and shifts in herbivory can lead to dramatic changes in the composition and diversity of aboveground plant communities. In turn, this can alter seed banks in the soil, which are cryptic reservoirs of plant diversity. Here, we use data from seven Nutrient Network grassland sites on four continents, encompassing a range of climatic and environmental conditions, to test the joint effects of fertilization and aboveground mammalian herbivory on seed banks and on the similarity between aboveground plant communities and seed banks. We find that fertilization decreases plant species richness and diversity in seed banks, and homogenizes composition between aboveground and seed bank communities. Fertilization increases seed bank abundance especially in the presence of herbivores, while this effect is smaller in the absence of herbivores. Our findings highlight that nutrient enrichment can weaken a diversity maintaining mechanism in grasslands, and that herbivory needs to be considered when assessing nutrient enrichment effects on seed bank abundance. 

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4. Top-Down and Bottom-Up Control in the Galápagos Upwelling System

   https://doi.org/10.3389/fmars.2022.845635  

   Brandt, Margarita; Silva-Romero, Isabel; Fernández-Garnica, David; Agudo-Adriani, Esteban; Bove, Colleen B.; Bruno, John F. (May 2022, Frontiers in Marine Science)

   Increased standing macroalgal biomass in upwelling zones is generally assumed to be the result of higher nutrient flux due to upwelled waters. However, other factors can strongly impact macroalgal communities. For example, herbivory and temperature, via their effects on primary producers and the metabolic demands of consumers, can also influence macroalgal biomass and productivity, respectively. We assessed the effects of nutrient availability, temperature, and herbivory on macroalgal biomass on a subtidal nearshore rocky reef in the Galápagos Islands. We manipulated nutrient availability and herbivory in field experiments performed in two seasons: the first during a cool, upwelling season, and the second during a warm, non-upwelling season. Excluding macro-herbivores had a clear effect on standing macroalgal biomass, independent of season or nutrient availability. However, we found different interactive effects of nutrients and macro-herbivores between the two seasons. During the cool season, macroalgal biomass was significantly higher in herbivore exclusions than in open areas under ambient nutrient conditions. However, when nutrients were added, macroalgal biomass was not significantly different across all herbivore treatments, which suggests reduced top-down control of herbivores (hence a greater standing algal biomass) in open areas. In the warm season, macroalgal biomass was significantly higher in herbivore exclusions compared to open treatments, both with and without nutrient addition. Furthermore, biomass reached 11X in herbivore exclusions with nutrient additions, which hints nutrient limitation only during warm, low-upwelling conditions. Overall, our results support the hypothesis that macro-herbivores reduce macroalgal biomass in this system and suggest that nutrient availability, but not temperature, modulate herbivory. 

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5. Soil nutrients cause threefold increase in pathogen and herbivore impacts on grassland plant biomass

   https://doi.org/10.1111/1365-2745.14111  

   Zaret, Max; Kinkel, Linda; Borer, Elizabeth T.; Seabloom, Eric W. (August 2023, Journal of Ecology)

   Abstract A combination of theory and experiments predicts that increasing soil nutrients will modify herbivore and microbial impacts on ecosystem carbon cycling.However, few studies of herbivores and soil nutrients have measured both ecosystem carbon fluxes and carbon pools. Even more rare are studies manipulating microbes and nutrients that look at ecosystem carbon cycling responses.We added nutrients to a long‐term, experiment manipulating foliar fungi, soil fungi, mammalian herbivores and arthropods in a low fertility grassland. We measured gross primary production (GPP), ecosystem respiration (ER), net ecosystem exchange (NEE) and plant biomass throughout the growing season to determine how nutrients modify consumer impacts on ecosystem carbon cycling.Nutrient addition increased above‐ground biomass and GPP, but not ER, resulting in an increase in ecosystem carbon uptake rate. Reducing foliar fungi and arthropods increased plant biomass. Nutrients amplified consumer effects on plant biomass, such that arthropods and foliar fungi had a threefold larger impact on above‐ground biomass in fertilized plots.Synthesis. Our work demonstrates that throughout the growing season soil resources modify carbon uptake rates as well as animal and fungal impacts on plant biomass production. Taken together, ongoing nutrient pollution may increase ecosystem carbon uptake and drive fungi and herbivores to have larger impacts on plant biomass production. 

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