He received a B.S. degree in Biology from the Massachusetts Institute of Technology in 1989 and a Ph.D. in Genetics from the University of California, Berkeley in 1995. He did postdoctoral research in the Department of Genetics at the University of Cambridge with support from a NSF / Sloan Foundation Fellowship in Molecular Evolution, and in the Section of Evolution and Ecology at the University of California, Davis with support from the NSF. He joined the Cornell faculty in 2004.