Metabolic rate throughout the annual cycle reveals the demands of an Arctic existence in Great Cormorants

Aquatic endotherms living in polar regions are faced with a multitude of challenges, including low air and water temperatures and low illumination, especially in winter. Like other endotherms from cold environments, Great Cormorants (Phalacrocorax carbo) living in Arctic waters were hypothesized to respond to these challenges through a combination of high daily rate of energy expenditure (DEE) and high food requirements, which are met by a high rate of catch per unit effort (CPUE). CPUE has previously been shown in Great Cormorants to be the highest of any diving bird. In the present study, we tested this hypothesis by making the first measurements of DEE and foraging activity of Arctic-dwelling Great Cormorants throughout the annual cycle. We demonstrate that, in fact, Great Cormorants have surprisingly low rates of DEE. This low DEE is attributed primarily to very low levels of foraging activity, particularly during winter, when the cormorants spent only 2% of their day submerged. Such a low level of foraging activity can only be sustained through consistently high foraging performance. We demonstrate that Great Cormorants have one of the highest recorded CPUEs for a diving predator; 18.6 g per minute submerged (95% prediction interval 13.0-24.2 g/min) during winter. Temporal variation in CPUE was investigated, and highest CPUE was associated with long days and shallow diving depths. The effect of day length is attributed to seasonal variation in prey abundance. Shallow diving leads to high CPUE because less time is spent swimming between the surface and the benthic zone where foraging occurs. Our study demonstrates the importance of obtaining accurate measurements of physiology and behavior from free-living animals when attempting to understand their ecology.