Resting costs too: the relative importance of active and resting energy expenditure in a sub-arctic seabird.

Breeding is costly for many animals, including birds that must deliver food to a central place (i.e. nest). Measuring energy expenditure throughout the breeding season can provide valuable insight into physiological limitations by highlighting periods of high demand, and ultimately allows improvement of conservation strategies. However, quantifying energy expenditure in wildlife can be challenging, as existing methods do not measure both active (e.g. foraging) and resting energy costs across short and long time scales. Here, we developed a novel method for comparing active and resting costs in 66 pre-breeding and breeding seabirds (black-legged kittiwakes, Rissa tridactyla) by combining accelerometry and triiodothyronine (T3) as proxies for active and resting costs, respectively. Active energy costs were higher during incubation (P=0.0004) and chick rearing (P<0.0001) than during pre-laying, because of an increase in the time spent in flight of 11% (P=0.0005) and 15% (P<0.0001), respectively. Levels of T3, reflecting resting costs, peaked marginally during incubation with a mean (±s.d.) concentration of 4.71±1.97 pg ml-1 in comparison to 2.66±1.30 pg ml-1 during pre-laying (P=0.05) and 3.16±2.85 pg ml-1 during chick rearing (P=0.11). Thus, although chick rearing is often assumed to be the costliest breeding stage by multiple studies, our results suggest that incubation could be more costly as a result of high resting costs. We highlight the importance of accounting for both active and resting costs when assessing energy expenditure.