title: Intraclutch differences in egg characteristics mitigate the consequences of age-related hierarchies in a wild passerine.
abstract: 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.
authors.affiliation: Department of Zoology, Edward Grey Institute, University of Oxford, Oxford, OX1 3PS, United Kingdom; Institute of Evolutionary Biology, School of Biological Sciences, University of Edinburgh, Edinburgh, EH9 3JT, United Kingdom. jarrod.hadfield@zoo.ox.ac.uk.
authors.forename: Jarrod D
authors.initials: JD
authors.lastname: Hadfield
authors.forename: Elizabeth A
authors.initials: EA
authors.lastname: Heap
authors.forename: Florian
authors.initials: F
authors.lastname: Bayer
authors.forename: Elizabeth A
authors.initials: EA
authors.lastname: Mittell
authors.forename: Nicholas M A
authors.initials: NM
authors.lastname: Crouch
mesh_terms: D000367:Age Factors; D000818:Animals; D004625:Embryo, Nonmammalian / Q000033:anatomy & histology*; D014644:Genetic Variation; D046109:Passeriformes / Q000033:anatomy & histology / Q000235:genetics* / Q000502:physiology; D011153:Population / Q000235:genetics; D012098:Reproduction / Q000235:genetics; D012641:Selection, Genetic
keywords: Cyanistes caeruleus; development; eggs; maternal effects; quantitative genetics; selection