Aggregative responses of marine predators to a pulsed resource.

Pulsed resources resulting from animal migrations represent important, transient influxes of high resource availability into recipient communities. The ability of predators to respond and exploit these large increases in background resource availability, however, may be constrained when the timing and magnitude of the resource pulse vary across years. In coastal Newfoundland, Canada, we studied aggregative responses of multiple seabird predators to the annual inshore pulse of a key forage fish species, capelin (Mallotus villosus). Seabird aggregative responses to fish biomass were quantified from weekly hydroacoustic and seabird surveys during July-August within an annually persistent foraging area (10 km²) associated with a cluster of capelin spawning sites across 10 years (2009-2010, 2012, 2014-2020). Seabird predators included breeding members of the families Alcidae (Common Murres Uria aalge, Razorbills Alca torda, Atlantic Puffins Fratercula arctica) and Laridae (Great Black-backed Gulls Larus marinus, American Herring Gulls L. argentatus smithsonianus) and Northern Gannets Morus bassanus, along with non-breeding, moulting members of the Family Procellariidae (Sooty Shearwaters Ardenna griseus, Great Shearwaters A. gravis). The inshore migration of spawning capelin resulted in 5-619 times (mean ± SE, 146 ± 59 times) increase in coastal fish biomass along with a shift towards more, larger and denser fish shoals. Within years, seabird abundance did not increase with inshore fish biomass but rather peaked near the first day of spawning, suggesting that seabirds primarily respond to the seasonal resource influx rather than short-term variation in fish biomass. Across years, the magnitude of the seabird aggregative response was lower during low-magnitude resource pulse years, suggesting that predators are unable to perceive low-magnitude pulses, avoid foraging under high competitor densities, and/or shift dietary reliance away from capelin under these conditions. The seabird response magnitude, however, was higher when the resource pulse was delayed relative to the long-term average, suggesting that predators increase exploitation during years of minimal overlap between the resource pulse and energetically demanding periods (e.g. breeding, moulting). This long-term study quantifying responses of multiple predators to a pulsed resource illustrates the ability of natural systems to tolerate natural and human-induced disturbance events.