Parental effort measures the total amount of time and energy allocated toward care of offspring, and it is often an obvious and variable component of an animal's life history (Winkler and Wilkinson 1988, Clutton-Brock 1991). Among birds, intraspecific variation in parental effort is more quantitative than qualitative. Individual adults differ in feeding rate (Yasukawa et al. 1990, Wright and Cuthill 1992), intensity of offspring defense (Winkler 1992), and tendency to engage in prolonged associations with young (Weatherhead and McRae 1990). Like all quantitative traits, intraspecific variation in parental effort can be attributed to genetic and environmental causes. However, recent work has focused mostly on environmental sources of variation (Ketterson and Nolan 1994, Gowaty 1996a), and individual birds often are viewed as decision-making machines that respond adaptively to changes in their physical or social environments (Winkler and Wilkinson 1988). Indeed, birds modify parental effort in response to numerous proximate factors, including genetic parentage (Lifjeld et al. 1998), brood size (Ruusila and P6ysa 1998), and offspring age and condition (Whittingham and Robertson 1993). In no case have environmental factors accounted completely for differences among individuals in the intensity of parental care, however, and some role for genetic or cultural determination seems likely. For quantitative traits, genetic and environmental effects can be distinguished by examining the phenotypic resemblance of relatives. In cases of high narrow-sense heritability (h2), additive genetic effects contribute substantially to overall phenotypic variation, and relatives strongly covary (Falconer and Mackay 1996). Among vertebrates, estimates of heritability are lacking for measures of parental effort and for most quantitative behaviors in the field (Hailman 1986, Boag and van Noordwijk 1987, Mousseau and Roff 1987; but see Waser and Jones 1989). In general, heritabilities for behavioral traits are low (Lemon 1993, Berthold and Pulido 1994), suggesting that quantitative differences among individuals are largely driven by the environment. Here, we examine the degree to which male and female feeding rates are heritable in an insular population of Savannah Sparrows (Passerculus sandwichensis). We show that quantitative differences in feeding rates among males can be attributed to the be-
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