[1] Penn State Univ, Dept Anthropol, University Pk, PA 16802 USA
[2] Penn State Univ, Ctr Conservat & Res, Omahas Henry Doorly Zoo, University Pk, PA 16802 USA
[3] Penn State Univ, Ctr Comparat Genom & Bioinformat, University Pk, PA 16802 USA
[4] Univ Calgary, Dept Anthropol, Calgary, AB T2N 1N4, Canada