Foraging Strategies of Glaucous-Winged Gulls in a Rocky Intertidal Community

Abstract

Foraging strategies of Glaucous-winged Gulls (Larus glaucescens) were studied in rocky intertidal habitats of the western Aleutian Islands, Alaska. Daily foraging activity was most intense at maximum low tide, and was concentrated in the lowest intertidal zones available to the birds. Barnacles (Balanus glandula) and mussels (Mytilus edulis) comprised most of the gull's diet during neap low tides, but these species were almost entirely abandoned during spring low tides in favor of sea urchins (Strongylocentrotus polyacanthus), chitons (Katharina tunicata), and limpets (Collisella pelta and Notoacmaea scutum). Sea urchins, chitons, and limpets, which had positive prey selection indices, were most abundant in the lower intertidal zones; barnacles and mussels, which had negative prey selection indices, were most abundant in the upper zones. Gulls also generally selected the larger individuals from each prey species, although sea urchins larger than the commissural bill width were avoided and limpets were selected in proportion to availability. Variation in prey availability also occurred among study areas with varying densities of sea otters (Enhydra lutris). With increasing depression of invertebrate prey by sea otter predation, gulls fed on a more diverse prey resource, and they switched to neritic fishes under intense sea otter predation. Preference experiments were conducted in the field, in which the common species and sizes of prey were made equally available to foraging gulls, thus eliminating search and capture times. In comparison with natural food choice, where sea urchins were most preferred, chitons became most preferred. We suggest that chitons are infrequent in natural diets because they adhere more strongly to the substratum than do sea urchins. Benefits of selective foraging were determined by comparing the net rate of energy gain of simulated random foragers with energy gained by selection of intertidal zones, prey species, and prey sizes. Observed selection patterns provided increased energy as gulls became more selective, and averaged 155% more than that obtained by the simulated random foragers. Results of the study support the two main predictions of optimal foraging theory in that (1) foraging patches (intertidal zones) and diets were selected such that net rates of energy gain were maximized, and (2) gulls became more selective foragers when energetically more profitable prey were more available.