Intraclutch differences in egg characteristics mitigate the consequences of age-related hierarchies in a wild passerine.

The relative age of an individual's siblings is a major cause of fitness variation in many species. In Blue tits (Cyanistes caeruleus), we show that age hierarchies are predominantly caused by incubation preclutch completion, such that last laid eggs hatch later than early laid eggs. However, after statistically controlling for incubation behavior late laid eggs are shown to hatch more quickly than early laid eggs reducing the amount of asynchrony. By experimentally switching early and late laid eggs between nests on the day they were laid, we controlled for the effect of differential incubation and found that the faster hatching times of late laid eggs remains. Chicks that hatched earlier were heavier and had higher probability of fledgling, and chicks that hatched from experimental eggs had patterns of growth and survival consistent with this. Egg mass explained a small part of this variation, but the remainder must be due to egg composition. These results are consistent with the idea that intrinsic differences between eggs across the laying sequence serve to mitigate the effects of age-related hierarchies. We also show that between-clutch variation in prenatal developmental rate exists and that it is mainly environmental in origin rather than genetic.

Disentangling genetic and prenatal sources of familial resemblance across ontogeny in a wild passerine.

Cross-fostering experiments are widely used by quantitative geneticists to study genetics and by behavioral ecologists to study the effects of prenatal investment. Generally, the effects of genes and prenatal investment are confounded and the interpretation given to such experiments is largely dependent on the interests of the researcher. Using a large-scale well-controlled experiment on a wild population of blue tits (Cyanistes caeruleus), we are able to partition variation in body mass across ontogeny into the effects of genes and the effects of between-clutch variation in egg characteristics. We show that although egg effects are important early in ontogeny they quickly dissipate, suggesting that the genetic interpretation of cross-fostering experiments may be preferable for many types of trait. However, the heritability of body mass is smaller than has previously been reported. Our results suggest that this is due to a combination of controlling postnatal environmental effects more carefully and accounting for viability selection operating early in ontogeny.