A central issue in the study of evolution in natural populations is how morphology evolves in response to environmental complexity, especially when environmental variation results in contrasting selective pressures (Levins 1968; Boyce 1979; Gomulkiewicz and Kirkpatrick 1992). In birds, contrasting selective pressures associated with seasonal fluctuations are common (e.g., Fretwell 1972; Newton 1972; Baker and Baker 1973; Price and Grant 1984; Schluter and Smith 1986). Here, we explore the consequences of seasonal fluctuations on the evolution of morphological traits related to foraging in the red crossbill (Loxia curvirostra). Crossbills are highly specialized for foraging on seeds in conifer cones (Benkman 1988, 1993; Benkman and Lindholm 1991) and rely on conifer seeds year-round (Newton 1972; Benkman 1987a, 1990, 1992). In seed-eating birds, bill size is the likely target of selection and so provides a logical focal trait for a study of the evolution of morphology related to foraging (e.g., Grant 1986). Here, we consider the evolution of bill size of one sibling species of red crossbill (L. curvirostra complex) that Groth (1993) designates as type 5. (Call types are recognized by their vocalizations, but also differ in bill and body size; Groth 1993.) Morphological traits associated with avian foraging ecology are, with few exceptions (e.g., Smith 1993), polygenically inherited (Boag and van Noordwijk 1987; Grant and Grant 1989). To make predictions concerning the evolution of quantitative traits, estimates of both the amount of genetic variation for the trait and the pattern of selection acting on the trait are needed. In this study, we assume that evolution of bill size is not genetically constrained and is determined mostly by selection. Based on the results of other studies of seed-eating birds (Boag and van Noordwijk 1987; Grant and Grant 1989) this simplifying assumption is unlikely to interfere with our predictions. The form of selection depends on the relationship between the trait and fitness. Field estimates of fitness are difficult to obtain for nomadic species such as red crossbills. Therefore, we make use of the partitioning suggested by Arnold (1983) where first the relationship between the trait and some measure of performance is estimated and then performance is related to fitness. We use the time to extract seeds from cones as a measure of performance, which can be measured in aviaries (Benkman 1993). Furthermore, we adjust foraging ef-