There is abundant evidence in many taxa for positive directional selection on body size, and yet little evidence for microevolutionary change. In many species, variation in body size is partly determined by the actions of parents, so a proposed explanation for stasis is the presence of a negative genetic correlation between direct and parental effects. Consequently, selecting genes for increased body size would result in a correlated decline in parental effects, reducing body size in the following generation. We show that these arguments implicitly assume that parental care is cost free, and that including a cost alters the predicted genetic architectures needed to explain stasis. Using a large cross-fostered population of blue tits, we estimate direct selection on parental effects for body mass, and show it is negative. Negative selection is consistent with a cost to parental care, mainly acting through a reduction in current fecundity rather than survival. Under these conditions, evolutionary stasis is possible for moderately negative genetic correlations between direct and parental effects. This is in contrast to the implausibly extreme correlations needed when care is assumed to be cost free. Thus, we highlight the importance of accounting correctly for complete selection acting on traits across generations.

Main DataframeLongform data frame used to analyse selection on offspring mass and parental performance. Dataframe is ordered as chick masses, chick survival, adult survival, adult fecundity.chicks.RDataSelection gradient estimates through female parental performanceThe output from estimation of selection gradients for each iteration of the model, through female parental performance traits.betasF.emn.RDataSelection gradient estimates through male parental performanceThe output from estimation of selection gradients for each iteration of the model, through female parental performance traits.betasM.emn.RDatatBIRDSContains fixed information for each bird, such as sex, parentage, hatch date, death datetMORPHMorphological measurements, including weight.tEGGSInformation about each egg, such as weight, laying date, hatching date.Main Analysis R ScriptR Script for carrying out the main analysesmain_analyses.RSelection Gradients R ScriptR Script for estimating selection gradients from the output of the modelsselection_gradients.R