Yolk androgen deposition in rockhopper penguins, a species with reversed hatching asynchrony

To maximize fitness, females should invest optimally in the siblings within a litter or brood and adapt this investment to environmental conditions. Chick mass and yolk androgens have been shown to influence the outcome of sibling competition. In birds, asynchronous hatching plays a major role in this process and often leads to brood reduction. We studied maternal deposition of yolk androgens in eggs of southern rockhopper penguins (Eudyptes chrysocome chrysocome). Contrary to other avian models, laying and hatching sequences do not coincide in this species, which exhibits reversed hatching asynchrony. This provides a unique model to test whether the first egg to hatch (B-egg), which is the most likely to survive, differs in composition from the second egg to hatch (A-egg). We found that B-eggs had higher egg masses, yolk masses, yolk androgen concentrations and total yolk androgen amounts than A-eggs. This was observed consistently for the three androgens analyzed (testosterone, androstenedione and 5α-dihydrotestosterone). Laying date affected androgen deposition into A- and B-eggs differently. Interestingly, late clutches had proportionally higher androgen levels in the B-egg compared to the A-egg than early clutches. We discuss these results in relation to the chronology of egg formation and the potential effect of the observed differences on embryo development and brood reduction.