To better understand how fluctuations in prey abundance may impact seabird reproductive success, we studied short-term changes in prey populations and their effect on prey selection and brood-rearing in the black-legged kittiwake Rissa tridactyla, a predator of near-surface-schooling forage fishes. Our fine-scale approach involved a weekly assessment of forage fish abundance and brood-rearing conditions during 4 consecutive years (1996 to 1999) at the Shoup Bay kittiwake colony in Prince William Sound, Alaska. We conducted forage fish surveys from a fixed-wing aircraft to determine weekly prey abundance throughout the known foraging range of breeding kittiwakes. Our results provide clear evidence that short-term fluctuations in prey availability are responsible for dramatic, within-season changes in the breeding conditions of black-legged kittiwakes. Adult kittiwakes often showed immediate response to changes in the prey base by altering prey selection; however, there were instances when kittiwakes selected prey species disproportionate to their availability (typically selecting for Pacific herring Clupea pallasi and against Pacific sand lance Ammodytes hexapterus). Changes in prey selection often resulted in striking differences in the amount of time required to obtain a load of food. The cascading effects of longer foraging trips was translated into reduced nestling growth and survival. Of the 3 components of energy provisioning to nestlings (meal delivery rate, meal size, and energy density), meal delivery rate had the strongest and most consistent positive effect on nestling growth and survival. Overall, these results demonstrate that complex foraging conditions limit the reproductive success of a central place-foraging species relying on an ephemeral food source. Moreover, we demonstrated that feeding conditions during the first 2 wk of brood-rearing were most critical for survival of the brood. Given the potential for such marked within-season variation in breeding conditions, it is critical that investigators adequately sample throughout the brood-rearing period, or, alternatively, select that portion that is germane to their study.
We sought to determine the effect of variation in time-activity budgets (TABs) and foraging behavior on energy expenditure rates of parent black-legged kittiwakes (Rissa tridactyla). We quantified TABs using direct observations of radio-tagged adults and simultaneously measured field metabolic rates (FMR) of these same individuals (n=20) using the doubly labeled water technique. Estimated metabolic rates of kittiwakes attending their brood at the nest or loafing near the colony were similar (ca. 1.3 x basal metabolic rate [BMR]), although loafing during foraging trips was more costly (2.9 x BMR). Metabolic rates during commuting flight (7.3 x BMR) and prey-searching flight (6.2 x BMR) were similar, while metabolic rates during plunge diving were much higher (ca. 47 x BMR). The proportion of the measurement interval spent foraging had a positive effect on FMR (R2=0.68), while the combined proportion of time engaged in nest attendance and loafing near the colony had a negative effect on FMR (R2=0.72). Thus, more than two-thirds of the variation in kittiwake FMR could be explained by the allocation of time among various activities. The high energetic cost of plunge diving relative to straight flight and searching flight suggests that kittiwakes can optimize their foraging strategy under conditions of low food availability by commuting long distances to feed in areas where gross foraging efficiency is high.
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.
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