In Situ Clock Shift Reveals that the Sun Compass Contributes to Orientation in a Pelagic Seabird

Compass orientation is central to the control of animal movement from the scale of local food-caching movements around a familiar area in parids [1] and corvids [2, 3] to the first autumn vector navigation of songbirds embarking on long-distance migration [4-6]. In the study of diurnal birds, where the homing pigeon, Columba livia, has been the main model, a time-compensated sun compass [7] is central to the two-step map-and-compass process of navigation from unfamiliar places, as well as guiding movement via a representation of familiar area landmarks [8-12]. However, its use by an actively navigating wild bird is yet to be shown. By phase shifting an animal's endogenous clock, known as clock-shifting [13-15], sun-compass use can be demonstrated when the animal incorrectly consults the sun's azimuthal position while homing after experimental displacement [15-17]. By applying clock-shift techniques at the nest of a wild bird during natural incubation, we show here that an oceanic navigator-the Manx shearwater, Puffinus puffinus-incorporates information from a time-compensated sun compass during homeward guidance to the breeding colony after displacement. Consistently with homing pigeons navigating within their familiar area [8, 9, 11, 18], we find that the effect of clock shift, while statistically robust, is partial in nature, possibly indicating the incorporation of guidance from landmarks into movement decisions.