[1] Univ Otago, Fac Dent, Sir John Walsh Res Inst, Dunedin, New Zealand
[2] Liverpool John Moores Univ, Sch Biol & Environm Sci, Res Ctr Evolutionary Anthropol & Palaeoecol, Liverpool, Merseyside, England
[3] Univ Witwatersrand, Ctr Explorat Deep Human Journey, Johannesburg, South Africa
[4] Tufts Univ, Dept Anthropol, Medford, MA 02155 USA
[5] Tufts Univ, Dept Biol, Medford, MA USA