Assessing the Consequences of Brood Parasitism and Nest Predation on Seasonal Fecundity in Passerine Birds

-Brood parasites and nest predators reduce the seasonal fecundity and, hence, the population growth rates of their victims. However, most field studies do not measure directly how parasites and predators decrease seasonal fecundity, but instead measure the impact of these organisms on individual nesting attempts. Because a female may renest after losing a nest to predation, abandoning a parasitized nest, or successfully fledging a brood, knowing how brood parasites and nest predators reduce the number of offspring fledged from individual nesting attempts is not equivalent to knowing their impact on seasonal fecundity. We address this problem by developing a mathematical model that: estimates several parameters describing the natural history of this system, including the brood-parasitism rate, nest-predation rate, and probability of nest abandonment in response to a parasitism event; and extrapolates to seasonal fecundity from these parameters and others describing the length of the breeding season, the timing of events in the nesting cycle, and the productivity of parasitized and unparasitized nests. We also show how different researchers using different observational methodologies to study exactly the same population likely would arrive at noticeably different conclusions regarding the intensity of brood parasitism, and we provide mathematical formulas for comparing among several of these measures of parasitism. Our procedures extend Mayfield's method for calculating nest-success rates from nest-history data in that we simultaneously estimate parameters describing nest predation and brood parasitism, predict seasonal fecundity from these parameters, and provide confidence intervals on all parameter estimates. The model should make the design and interpretation of logistically difficult empirical studies more efficient. It also can be specialized to species affected by nest predators but not brood parasites. We use the model to analyze Prairie Warbler (Dendroica discolor) and Black-capped Vireo (Vireo atricapillus) nesting data. We estimate the model's parameters for these species and use the resulting estimates to predict seasonal fecundity. For both species, the predicted seasonal fecundity closely matches the value measured directly. Received 30 November 1993, accepted 15 February 1994. BROOD PARASITES AND NEST PREDATORS cause many passerine nests to fail before any young are fledged (Ricklefs 1969). In addition, when a parasitized nest is not immediately abandoned, the number of host young that can be fledged is often greatly reduced. As Rothstein (1990) summarized, about 80 bird species are interspecific brood parasites, including cowbirds (Emberizidae, Icterinae), cuckoos (Cuculidae, Cuculinae and Neomorphinae), the Cuckoo-finch (Anomalospiza imberbis), whydahs (Ploceidae, Viduinae), honeyguides (Indicatoridae), and the Black-headed Duck (Heteronetta atricapillus; Friedmann 1929, 1955, Weller 1959, Payne 1977, Wyllie 1981). Nest predation is known from all passerines studied in any detail, and includes predation by small mammals, birds,