Familiar route loyalty implies visual pilotage in the homing pigeon

Wide-ranging animals, such as birds, regularly traverse large areas of the landscape efficiently in the course of their local movement patterns, which raises fundamental questions about the cognitive mechanisms involved. By using precision global-positioning-system loggers, we show that homing pigeons (Columba livia) not only come to rely on highly stereotyped yet surprisingly inefficient routes within the local area but are attracted directly back to their individually preferred routes even when released from novel sites off-route. This precise route loyalty demonstrates a reliance on familiar landmarks throughout the flight, which was unexpected under current models of avian navigation. We discuss how visual landmarks may be encoded as waypoints within familiar route maps.