Differences in egg size, shell thickness, pore density, pore diameter and water vapour conductance between first and second eggs of Snares Penguins Eudyptes robustus and their influence on hatching asynchrony

Brood reduction in birds is frequently induced by hatching asynchrony. Crested penguins (genus Eudyptes) are obligate brood reducers, but in contrast to most other birds, first-laid eggs are considerably smaller in size than second-laid eggs; furthermore, first-laid eggs hatch after their siblings. The mechanisms underlying this reversal in size and hatching order remain unclear. In this study, we tested whether the second-laid eggs of Snares Penguins Eudyptes robustus have a higher eggshell porosity allowing them to maintain a higher metabolic rate throughout incubation and to hatch before their first-laid siblings. We investigated differences in egg size, shell thickness, pore density, pore diameter and water vapour conductance between first and second eggs within clutches and examined the influence of these shell characteristics on hatching asynchrony. First-laid eggs of Snares Penguins were approximately 78% of the size of the larger second eggs. Second-laid eggs had considerably thicker shells and more pores per cm² than first eggs, whereas pore diameter did not differ between eggs. Water vapour conductance was greater in second- (16.8 mg/day/torr) than in first-laid eggs (14.9 mg/day/torr). The difference in water vapour conductance between first- and second-laid eggs within clutches was related to hatching patterns. In nests where second eggs hatched before first-laid eggs, second eggs had a considerably greater water conductance than their sibling, whereas in nests where both eggs hatched on the same day, the difference in water conductance between eggs was very small, and in a few nests where small first eggs hatched before their larger sibling, they had a greater water conductance than their larger second-laid nestmate. Surprisingly few studies have investigated differences in shell characteristics between eggs within clutches and associated effects on hatching asynchrony. This study has demonstrated that such differences exist between eggs within clutches and that they can influence hatching patterns.