[1] Spanish Natl Res Council, Computat Biol Lab, Estn Biol Donana, Seville, Spain
[2] Univ Lyon, INRIA Beagle Team, INSA Lyon, LIRIS UMR CNRS 5205, Lyon, France
[3] Michigan State Univ, BEACON Ctr Study Evolut Act, E Lansing, MI USA
[4] Michigan State Univ, Ecol Evolut Behav Program, E Lansing, MI USA
[5] Michigan State Univ, Dept Comp Sci & Engn, E Lansing, MI USA