Survival costs of chick rearing in black-legged kittiwakes

1. We tested for costs of chick rearing in the black-legged kittiwake Rissa tridactyla (Linnaeus) by removing entire clutches from 149 of 405 randomly selected nests, in which one or both mates was colour-banded. After the manipulation, we monitored adult nest attendance and body condition at unmanipulated and manipulated nests, and measured the survival and fecundity of these adults the following year. 2. Late in the chick-rearing period, adults from unmanipulated nests (i.e. with chicks) went on significantly longer foraging trips, and were significantly lighter for their size, than adults from manipulated nests (i.e. without chicks). 3. Adults from unmanipulated nests also survived to the following nesting season at a significantly lower rate than those from the manipulated nests (0.898 vs. 0.953), suggesting that attempting to raise chicks can reduce life expectancy by 55%. 4. There was a tendency for adults from nests that were unmanipulated in year one to have lower reproductive success in year two, primarily because of reduced hedging success, and a higher incidence of non-breeding. 5. These findings suggest that mass loss in kittiwakes during chick rearing may not be adaptive. Raising chicks can lead to reproductive costs, and the causal mechanism appears to be a reduction in body condition. 6. We compare our results with previous brood (or clutch) size manipulation experiments that have measured adult body condition, survival and/or future fecundity. Although the empirical evidence suggests that long-lived species are more likely to experience survival costs than short-lived species, we believe the opposite may be true. We suggest that shifting the experimental protocol of cost of reproduction studies from brood enlargements (an approach taken in most prior studies) to brood reductions will provide more accurate quantifications of naturally occurring costs. 7. The cost of reproduction is one mechanism proposed to explain the reduced survival rates reported for kittiwake populations in the North Atlantic relative to those in the North Pacific ocean. Oceanographic data, however, suggest that lower food availability may limit survival of kittiwakes in the North Atlantic where a deeper mixed layer and reduced primary production combine to make conditions less favourable for this seabird during the winter months.