Abstract
Density‐dependent competition for food resources influences both foraging ecology and reproduction in a variety of animals. The relationship between colony size, local prey depletion, and reproductive output in colonial central‐place foragers has been extensively studied in seabirds; however, most studies have focused on effects of intraspecific competition during the breeding season, while little is known about whether density‐dependent resource depletion influences individual migratory behavior outside the breeding season. Using breeding colony size as a surrogate for intraspecific resource competition, we tested for effects of colony size on breeding home range, nestling health, and migratory patterns of a nearshore colonial seabird, the brown pelican (Pelecanus occidentalis), originating from seven breeding colonies of varying sizes in the subtropical northern Gulf of Mexico. We found evidence for density‐dependent effects on foraging behavior during the breeding season, as individual foraging areas increased linearly with the number of breeding pairs per colony. Contrary to our predictions, however, nestlings from more numerous colonies with larger foraging ranges did not experience either decreased condition or increased stress. During nonbreeding, individuals from larger colonies were more likely to migrate, and traveled longer distances, than individuals from smaller colonies, indicating that the influence of density‐dependent effects on distribution persists into the nonbreeding period. We also found significant effects of individual physical condition, particularly body size, on migratory behavior, which in combination with colony size suggesting that dominant individuals remain closer to breeding sites during winter. We conclude that density‐dependent competition may be an important driver of both the extent of foraging ranges and the degree of migration exhibited by brown pelicans. However, the effects of density‐dependent competition on breeding success and population regulation remain uncertain in this system.
Highlights
Density dependence, or the feedback between population size and population growth rate, acts as a stabilizing mechanism in ecological communities by altering individual behavior and fitness (Fowler, 1987; Mylis & Diekmann, 1995)
One of the principal mechanisms underlying the shift from positive to negative density-dependent effects is the increase in intraspecific competition coincident with increasing population size, which can result in lower resource availability and reduced individual fitness (Fowler, 1987)
Because the foraging ranges of central-place foragers are limited at some level by energetic constraints (Orians & Pearson, 1979), individuals in more numerous colonies experience intensified localized competition for food resources, which results in reduced foraging success or increased foraging costs due to direct resource depletion, conspecific interference, and altered prey behavior (Kuhn, Baker, Towell, & Ream, 2014; Lewis, Sherratt, Hamer, & Wanless, 2001)
Summary
The feedback between population size and population growth rate, acts as a stabilizing mechanism in ecological communities by altering individual behavior and fitness (Fowler, 1987; Mylis & Diekmann, 1995). Individual foraging ranges during the breeding period are often constrained by the need to return to the colony site at regular intervals to feed offspring, a process known as central-place foraging (Orians & Pearson, 1979) Both foraging effort and offspring condition are frequently used to test for the presence and direction of density- dependent effects in colonial breeders. Diamond (1978) tested colony size relationships across several tropical species and found that species that bred in larger colonies were more likely to migrate than species with smaller average colony sizes This remains the only example of density-dependent constraints on migratory patterns in seabirds, and it focused on species-wide patterns rather than individual strategies. Laughing gulls, which are kleptoparasitic feeders and may target brown pelicans, were not accurately censused in the study region; our observations suggest they were present at similar densities across all study colonies
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