1. Department of Ecology and Evolutionary Biology, University of California, Los Angeles, California 90095, USA;
2. Department of Molecular and Cell Biology, University of California, Merced, California 95343, USA
3. Current affiliation: Department of Ecology and Evolutionary Biology, Brown University, Providence, Rhode Island 02912, USA
4. Interdepartmental Program in Bioinformatics and Department of Human Genetics, David Geffen School of Medicine, University of California, Los Angeles, California 90095, USA