Active territory defence at a low energy cost in a colonial seabird

Aggressive behaviour associated with the defence of a territory is thought to impose substantial energy costs and thus to represent a trade-off with other energy-demanding activities. The energy costs of aggressive behaviours, however, have rarely been estimated in the wild, and the overall contribution of territorial defence to daily energy expenditure has never been determined. We studied the activity budget of breeding king penguins, Aptenodytes patagonicus, equipped with heart rate data loggers to estimate the energy costs associated with territory defence in this colonial bird exhibiting very high rates of agonistic interactions. We also assessed whether threat displays imposed lower energy costs than attacks with body contact. During territorial defence (i.e. threats and physical attacks combined), energy expenditure averaged 1.27 times resting metabolic rate. Defence accounted for 13% of the daily time budget and contributed to 2.7% of the total daily energy expenditure. Interactions with body contact cost three times more than threat displays, but accounted for only 16% of the aggressive behaviours recorded. Neither did body mass, body size, penguin sex or breeding stage affect the cost of aggressiveness. Our results are consistent with previous research reporting that fighting imposes significant metabolic costs. However, we found that aggressive behaviour in king penguins was not an expensive activity compared to the total energy budget. Because king penguins go without food and are sleep deprived while breeding, they may have developed behavioural strategies (e.g. lower rates of attacks with body contact) allowing them to defend their territory efficiently at a low energy cost.