Abstract
1. For seabirds that forage at sea but breed while fasting on land, successful reproduction depends on the effective management of energy stores. Additionally, breeding often means aggregating in dense colonies where social stress may affect energy budgets. 2. Male king penguins (Aptenodytes patagonicus) fast for remarkably long periods (up to 1 5 months) while courting and incubating ashore. Although their fasting capacities have been well investigated in captivity, we still know very little about the energetics of freely breeding birds. 3. We monitored heart rate (HR, a proxy to energy expenditure), body temperature and physical activity of male king penguins during their courtship and first incubation shift in a colony of some 24 000 freely breeding pairs. Males were breeding either under low but increasing colony density (early breeders) or at high and stable density (late breeders). 4. In early breeders, daily mean and resting HR decreased during courtship but increased again 3 days before egg laying and during incubation. In late breeders, HR remained stable throughout this same breeding period. Interestingly, the daily increase in resting HR we observed in early breeders was strongly associated with a marked increase in colony density over time. This finding remained significant even after controlling for climate effects. 5. In both early and late breeders, courtship and incubation were associated with a progressive decrease in physical activity, whereas core body temperature remained unchanged. 6. We discuss the roles of decreased physical activity and thermoregulatory strategies in sustaining the long courtship-incubation fast of male king penguins. We also draw attention to a potential role of conspecific density in affecting the energetics of breeding-fasting seabirds, that is, a potential energy cost to coloniality.