[1] Penn State Univ, Dept Anthropol, University Pk, PA 16802 USA
[2] Penn State Univ, Dept Biol, University Pk, PA 16802 USA
[3] Univ Tubingen, DFG Ctr Adv Studies Words Bones Genes Tools, Tubingen, Germany
[4] Liverpool John Moores Univ, Res Ctr Evolutionary Anthropol & Palaeoecol, Liverpool, Merseyside, England