Gulls of a feather do not sleep whenever—circadian rhythm of activity of black-headed gulls Chroicocephalus ridibundus during the incubation period

Many vertebrates exhibit a diel activity, steered by light–dark cycle. However, some colonial waterbirds, in that several species of gulls, are active not only in day hours but also at night. In this study, we aimed to investigate 24 h cycle of black-headed gulls (BHG) Chroicocephalus ridibundus activity with focus on sleep behaviour. We expected that 24 h patterns of activity differ between colonies located in various habitats, and within a colony between nests located in the centre vs at the edge. We studied behaviour based on 9600 of 30 s videos from camera-traps taken in six colonies and data from 10 GPS-tracked individuals from one colony recorded during incubation. BHGs stayed active on average during 48.1% of a night, mainly spent on passive and active nest defence, and on nest maintenance. BHGs spent similar time on these activities in day hours. Individuals breeding in the colony centre slept at night longer than those at its edge. BHGs stayed active during on average 76.5% of daytime. In two urban colonies with the highest nest densities and highest level of light pollution birds slept less during the day than in other studied colonies (three rural and one urban) characterized by lower densities and light intensity after sunset near the colony. Knowledge of nocturnal behaviour is crucial to comprehend 24 h activity patterns of an organism, especially to understand flexibility of behaviour crucial for restoration, like sleep.