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We used a supplemental-feeding experiment, the doubly labeled water technique, and a model-selection approach based upon the Akaike Information Criterion to examine effects of food availability on energy expenditure rates of Black-legged Kittiwakes, Rissa tridactyla, raising young in Middleton Island, Alaska. Energy expenditure rates of females were more responsive to fluctuations in food availability than those of males. Our objectives were to determine if the energy expenditure rates of parent kittiwakes provided with supplemental food, differed from those of control parents, and to determine the strength of the relationship between parental energy expenditure rates and a suite of extrinsic and intrinsic covariates and factors. Fed males likely expended more energy while off the nest than fed females, possibly because of nest defense. Energy expenditure rates of fed kittiwakes were similar to values reported for kittiwakes that were either not raising young or not foraging. Fed kittiwakes increased their reproductive output while simultaneously decreasing parental effort and increasing body condition. Parent kittiwakes, therefore, adjusted parental effort in response to variation in breeding conditions due to changes in food availability. Adjustments in reproductive effort in response to variable foraging conditions may have significant effects on the survival and productivity of individuals, and thus provide substantial fitness benefits for long-lived seabirds such as Black-legged Kittiwakes. Differences in survival rates between the sexes may occur if each sex experiences a different survival curve with respect to field metabolic rate or parental effort.

Abstract: We investigated the relative roles of bottom-up and top-down factors in limiting productivity of an upper trophic level marine predator. Our primary working hypothesis was that the reproductive success of black-legged kittiwakes (Rissa tridactyla) a piscivorous, colonial-nesting seabird, was most limited by the abundance, distribution, and species composition of surface-schooling forage fishes. A secondary working hypothesis was that reproductive loss to kittiwake nest predators was greatest during years of reduced prey availability. We report on a broad-scale, integrated study of kittiwakes and their prey in Prince William Sound, Alaska. Our study spanned five breeding seasons (1995–1999) and focused on three colonies that differed in size (ranging from ca. 220 to ca. 7000 breeding pairs) and proximity to each other (50–135km apart). Kittiwakes in PWS encountered a variety of aquatic habitats, creating a complex foraging environment for breeding birds. We measured kittiwake reproductive success and foraging activities, while simultaneously measuring the abundance of surface schooling forage fishes throughout the foraging range of breeding kittiwakes. The abundance of primary prey species for kittiwakes (Pacific herring Clupea pallasi, Pacific sand lance Ammodytes hexapterus, and capelin Mallotus villosus) varied both annually and regionally, with no one region consistently having the greatest abundance of prey. Likewise, kittiwake reproductive success varied considerably among colonies and years. We found that bottom-up, top-down, timing mismatch, and colony-specific effects were all important to kittiwake productivity. Although bottom-up effects appeared to be strongest, they were not evident in some cases until other effects, such as geographic location (proximity of colony to prey concentrations) and top-down predation, were considered. Important bottom-up effects on kittiwake reproductive success were not only total prey abundance and distribution, but also species, age composition, and chronology of prey occurrence (match/mismatch of timing with critical brood-rearing periods); these effects varied by colony. Top-down effects of predation on kittiwake nest contents (independent of prey abundance) confounded seabird-forage fish relationships. Ultimately, when confounding factors were minimized, non-linear asymptotic relationships were identified between kittiwakes and their prey, with an asymptotic threshold of fish school surface area density of ca. 5m2/km2, beyond which top-down, physiological, or phylogenetic constraints likely restrict further reproductive output. The integrated approach of our investigations provided a more thorough understanding of the mechanisms underlying predator–prey relationships in the complex marine environment. However, such mechanistic theories can only be tested and refined through long-term research and monitoring of much greater duration than the 5-year study reported herein. [Copyright &y& Elsevier]