An omnivore vigour hypothesis? Nutrient availability strengthens herbivore suppression by omnivores across 48 field sites.

Nutrients regulate herbivore growth from the 'bottom-up' via improved plant vigour and food quality. Nitrogen also affects 'top-down' control of herbivores by moderating attraction of predators and the rates at which they consume herbivorous prey. Tri-trophic consequences of nitrogen availability are more challenging to predict among omnivorous natural enemies who feed on both plants and herbivores, limiting our ability to predict net outcomes of nutrient availability in food webs. In a two-year field survey of insects on zucchini host plants at 48 sites, I predicted that both herbivores and foliar-feeding omnivores would increase with nutrient availability, while predators would not. My results revealed positive relationships between omnivores and foliar nitrogen concentrations, while predators had neutral responses to foliar N. Surprisingly, herbivores declined with increasing foliar N across the field sites. Greenhouse experiments re-enforced these patterns, as herbivore growth inversely correlated with soil N concentrations in communities that included foliar-feeding omnivores. Conversely, herbivore growth was uncorrelated with soil N on plants with predators, nor on predator-free plants. These results suggest that omnivores mount strong and consistent responses to nitrogen in plant tissues in a variety of ecological contexts. In environments where omnivorous arthropods can thrive, their recruitment to nitrogen-rich plants may increase predation and thereby counterbalance and stabilize 'bottom-up' increases in herbivore performance supported by enhanced foliar nutrition.