Environmental and not maternal effects determine variation in offspring phenotypes in a passerine bird

Maternal and environmental effects can profoundly influence offspring phenotypes, independent of genetic effects. Within avian broods, both the asymmetric post-hatching environment created by hatching asynchrony and the differential maternal investment through the laying sequence have important consequences for individual nestlings in terms of the allocation of resources to body structures with different contributions to fitness. The purpose of this study was to evaluate the relative importance of post-hatching environmental and maternal effects in generating variation in offspring phenotypes. First, an observational study showed that within blue tit, Cyanistes caeruleus, broods, late-hatched nestlings allocated resources to tarsus development, maintained mass gain and head-bill growth and directed resources away from the development of fourth primary feathers. Second, a hatching order manipulation experiment resulted in nestlings from first-laid eggs hatching last, thereby allowing comparison with both late and early-hatched nestlings. Experimental nestlings had growth patterns which were closer to late-hatched nestlings, suggesting that within-brood growth patterns are determined by post-hatching environmental effects. Therefore, we conclude that post-hatching environmental effects play an important role in generating variation in offspring phenotypes.