[1] Michigan State Univ, Dept Integrat Biol, E Lansing, MI 48824 USA
[2] Michigan State Univ, Program Ecol Evolut & Behav, E Lansing, MI 48824 USA
[3] Univ Colorado, Dept Ecol & Evolutionary Biol, Boulder, CO 80309 USA
[4] Mara Hyena Project, Maasai Mara, Narok County, Kenya
[5] Nova Southeastern Univ, Halmos Coll Nat Sci & Oceanog, Dept Biol Sci, Ft Lauderdale, FL 33314 USA
[6] Max Planck Inst Anim Behav, Dept Ecol Anim Soc, Constance, Germany