[1] Univ Wyoming, Sch Comp, Laramie, WY 82071 USA
[2] Univ Wyoming, Haub Sch Environm & Nat Resources, Laramie, WY 82072 USA
[3] Max Planck Inst Anim Behav, Collect Migrat Grp, D-78315 Radolfzell am Bodensee, Germany
[4] Univ Konstanz, Ctr Adv Study Collect Behav, D-78468 Constance, Germany
[5] Max Planck Inst Anim Behav, Dept Migrat, D-78315 Radolfzell am Bodensee, Germany
[6] Univ Konstanz, Dept Biol, D-78457 Constance, Germany