[1] Georgia Inst Technol, Sch Biol, Atlanta, GA 30332 USA
[2] Univ Minnesota, St Paul, MN 55108 USA
[3] Max Planck Inst Evolut Biol, D-24306 Plon, Germany
[4] UCL, Dept Genet Evolut & Environm, London WC1N 6BT, England
[5] Univ Minnesota, Dept Ecol Evolut & Behav, St Paul, MN 55108 USA
[6] Univ Minnesota, BioTechnol Inst, St Paul, MN 55108 USA