[1] Univ Calif Los Angeles, Dept Biomath, Los Angeles, CA 90095 USA
[2] Univ Calif Los Angeles, Dept Biostat, Los Angeles, CA 90095 USA
[3] Univ Calif Los Angeles, Dept Human Genet, Los Angeles, CA 90095 USA
[4] Univ Edinburgh, Inst Evolutionary Biol, Edinburgh EH9 3JT, Midlothian, Scotland