Habitat use across the tidal cycle by black-headed gulls breeding in the Wadden Sea

Movements of birds foraging in intertidal areas are often strongly linked to the tidal cycle, as water levels determine where and when birds can forage. The strength of this link likely depends on the ability to forage in habitats other than intertidal areas and on constraints imposed by breeding duties. Few studies have focused on the use of intertidal areas by generalists, such as the black-headed gull Chroicocephalus ridibundus, that occupy a wide variety of habitats within and beside intertidal mudflats. We investigated to what degree black-headed gulls (1) use intertidal mudflats versus terrestrial habitats during different phases of the tidal cycle, (2) follow the tidal wave to exploit recently exposed mudflats and (3) whether these behaviours are influenced by central place foraging. For this, 11 black-headed gulls breeding on the Wadden Sea island Griend were tracked during two years using GPS loggers. When commuting to and from Griend, up to 75