[1] UCL, Res Dept Genet Evolut & Environm, London WC1E 6BT, England
[2] Univ Oxford, Dept Zool, Oxford OX1 3PS, England
[3] Australian Natl Univ, Res Sch Biol, Ctr Macroevolut & Macroecol, Canberra, ACT 0200, Australia
[4] Royal Holloway Univ London, Sch Biol Sci, Egham TW20 0EX, Surrey, England
[5] Australian Natl Univ, Sch Archaeol & Anthropol, Canberra, ACT 0200, Australia