Nest predation by cowbirds and its consequences for passerine demography.

Brood parasitism by brown-headed cowbirds (Molothrus ater) reduces reproductive success in many passerines that nest in fragmented habitats and ecological edges, where nest predation is also common. We tested the hypothesis that parasitism and predation are often linked because cowbirds depredate nests discovered late in the host's nesting cycle to enhance future opportunities for parasitism. Over a 20-year study period, brood parasitism by cowbirds was a prerequisite to observing marked inter- and intraannual variation in the rate of nest failure in an insular song sparrow (Melospiza melodia) population. Nest failure increased with the arrival and laying rate of cowbirds and declined when cowbirds ceased laying. The absence or removal of cowbirds yielded the lowest nest failure rates recorded in the study. The absence of cowbirds also coincided with the absence of an otherwise strong positive correlation between host numbers and the annual rate of nest failure. Host numbers, cowbird parasitism, and nest failure may be correlated because cowbirds facilitate nest failure rather than cause it directly. However, an experiment mimicking egg ejection by cowbirds did not affect nest failure, and, contrary to the main prediction of the predation facilitation hypothesis, naturally parasitized nests failed less often than unparasitized nests. Higher survival of parasitized nests is expected under the cowbird predation hypothesis when female cowbirds defend access to hosts because cowbirds should often depredate unparasitized nests but should not depredate nests they have laid in. Where female cowbirds have overlapping laying areas, we expect parasitized nests to fail more often than others if different cowbirds often discover the same nests. We suggest that nest predation by cowbirds represents an adaptation for successful parasitism and that cowbirds influence host demography via nest predation.