[1] Univ London Royal Holloway & Bedford New Coll, Sch Biol Sci, Egham TW20 0EX, Surrey, England
[2] Max Planck Inst Verhaltensphysiol, D-82319 Seewiesen, Germany
[3] Max Planck Res Ctr Ornithol, D-78215 Radolfzell am Bodensee, Germany
[4] Cornell Univ, Dept Psychol, Ithaca, NY 14853 USA