Potential mode of clutch size determination and follicle development in Eudyptes penguins

It has long been held that Eudyptes penguins will only ever develop a maximum of two mature yolky follicles to match their invariant two-egg clutch, an idea inferred largely from egg removal studies. Combining our own data with those from a previous but rarely cited study and by applying these to a simple developmental model, we show that macaroni penguins (Eudyptes chrysolophus) develop up to four large, yolky vitellogenic follicles, and they do so despite the fact that they will never lay more than two eggs or rear more than one chick, a tactic that seems maladaptive given their realized reproductive success. We discuss these results within the context of the usual pattern of reproductive investment in Eudyptes penguins and suggest a broader significance to modes of clutch size determination among all penguins (order Sphenisciformes).