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Herbivorous insects can influence grassland ecosystem functions in several ways, notably by altering primary production and nutrient turnover. Interactions between above- and belowground herbivory could affect these functions; an effect that might be modified by nitrogen (N) addition, an important global change driver. To explore this, we added above- (grasshoppers) and belowground (wireworms) insect herbivores and N into enclosed, equally composed, grassland plant communities in a fully factorial field experiment. N addition substantially altered the impact of above- and belowground herbivory on ecosystem functioning. Herbivory and N interacted such that biomass was reduced under above ground herbivory and high N input, while plant biomass remained stable under simultaneous above- and belowground herbivory. Aboveground herbivory lowered nutrient turnover rate in the soil, while belowground herbivory mitigated the effect of aboveground herbivory. Soil decomposition potential and N mineralization rate were faster under belowground herbivory at ambient N, but at elevated N this effect was only observed when aboveground herbivores were also present. We found that N addition does not only influence productivity directly (repeatedly shown by others), but also appears to influence productivity by herbivory mediated effects on nutrient dynamics, which highlights the importance of a better understanding of complex biotic interactions.

Understanding the role that species interactions play in determining the rate and direction of ecosystem change due to nitrogen (N) eutrophication is important for predicting the consequences of global change. Insects might play a major role in this context. They consume substantial amounts of plant biomass and can alter competitive interactions among plants, indirectly shaping plant community composition. Nitrogen eutrophication affects plant communities globally, but there is limited experimental evidence of how insect herbivory modifies plant community response to raised N levels. Even less is known about the roles of above- and belowground herbivory in shaping plant communities, and how the interaction between the two might modify a plant community's response to N eutrophication. We conducted a 3-yr field experiment where grassland plant communities were subjected to above- and belowground insect herbivory with and without N addition, in a full-factorial design. We found that herbivory modified plant community responses to N addition. Aboveground herbivory decreased aboveground plant community biomass by 21%, but only at elevated N. When combined, above- and belowground herbivory had a stronger negative effect on plant community biomass at ambient N (11% decrease) than at elevated N (4% decrease). In addition, herbivory shifted the functional composition of the plant community, and the magnitude of the shifts depended on the N level. The N and herbivory treatments synergistically conferred a competitive advantage to forbs, which benefited when both herbivory types were present at elevated N. Evenness among the plant species groups increased when aboveground herbivory was present, but N addition attenuated this increase. Our results demonstrate that a deeper understanding of how plant-herbivore interactions above and below ground shape the composition of a plant community is crucial for making reliable predictions about the ecological consequences of global change., (© 2016 by the Ecological Society of America.)