Extreme intraclutch egg-size dimorphism in Eudyptes penguins, an evolutionary response to clutch-size maladaptation

Eudyptes penguins (six species) are uniquely characterized by a two-egg clutch with extreme intraclutch egg-size dimorphism (ESD): the first-laid A-egg is 17.5%–56.9% smaller than the B-egg. Although A-eggs are viable, they almost never produce fledged chicks (genus average <1%). Using classical life-history theory and phylogenetic comparative methods, we demonstrate a marked slowdown in the life history of Eudyptes: age of first reproduction is 52% later and annual fecundity 48% lower compared with other two-egg clutch penguin species. All six Eudyptes species have retained a two-egg clutch, despite this pronounced life-history slowdown; this suggests evolutionary mismatch between clutch size and chicks fledged per clutch. Consistent with this, we show that Eudyptes fledge 43% fewer chicks per clutch than other two-egg clutch penguin species. Extreme intraclutch ESD in Eudyptes is associated primarily with a uniform (5%) increase in relative B-egg size, and B-egg size has evolved in...