hiː wʌz ɡɹˈæntᵻd hɪz pˌiːˈeɪtʃ.dˈiː. ɪn pˌɑːpjʊlˈeɪʃən baɪˈɑːlədʒi fɹʌm stˈænfɚd jˌuːnɪvˈɜːsᵻɾi ɪn nˈaɪntiːnhˈʌndɹɪd sˈɛvənti tˈuː, hɪz θˈiːsɪs bˌiːɪŋ ˌɔn dʒɪnˈɛɾɪk vˌɛɹɪˈeɪʃən ɪn ˈælpaɪn bˈʌɾɚflˌaɪz.
He was granted his Ph.D. in population biology from Stanford University in 1972, his thesis being on genetic variation in alpine butterflies.