[1] Max Planck Inst Evolutionary Biol, Dept Theoret Biol, D-24306 Plon, Germany
[2] Princeton Univ, Dept Ecol & Evolutionary Biol, Princeton, NJ 08544 USA
[3] Princeton Univ, Lewis Sigler Inst Integrat Genom, Princeton, NJ 08544 USA
[4] Univ Calif Berkeley, Miller Inst Basic Res Sci, Berkeley, CA 94720 USA
[5] Univ Calif Berkeley, Dept Integrat Biol, Berkeley, CA 94720 USA