Burrowing seabird effects on invertebrate communities in soil and litter are dominated by ecosystem engineering rather than nutrient addition

Vertebrate consumers can be important drivers of the structure and functioning of ecosystems, including the soil and litter invertebrate communities that drive many ecosystem processes. Burrowing seabirds, as prevalent vertebrate consumers, have the potential to impact consumptive effects via adding marine nutrients to soil (i.e. resource subsidies) and non-consumptive effects via soil disturbance associated with excavating burrows (i.e. ecosystem engineering). However, the exact mechanisms by which they influence invertebrates are poorly understood. We examined how soil chemistry and plant and invertebrate communities changed across a gradient of seabird burrow density on two islands in northern New Zealand. Increasing seabird burrow density was associated with increased soil nutrient availability and changes in plant community structure and the abundance of nearly all the measured invertebrate groups. Increasing seabird densities had a negative effect on invertebrates that were strongly influenced by soil-surface litter, a positive effect on fungal-feeding invertebrates, and variable effects on invertebrate groups with diverse feeding strategies. Gastropoda and Araneae species richness and composition were also influenced by seabird activity. Generalized multilevel path analysis revealed that invertebrate responses were strongly driven by seabird engineering effects, via increased soil disturbance, reduced soil-surface litter, and changes in trophic interactions. Almost no significant effects of resource subsidies were detected. Our results show that seabirds, and in particular their non-consumptive effects, were significant drivers of invertebrate food web structure. Reductions in seabird populations, due to predation and human activity, may therefore have far-reaching consequences for the functioning of these ecosystems.