[1] UCL, Dept Biol, London WC1E 6BT, England
[2] Univ London Royal Holloway & Bedford New Coll, Sch Biol Sci, Egham TW20 0EX, Surrey, England
[3] Univ Oxford, Dept Zool, Oxford OX1 3PS, England
[4] Nat Hist Museum, Dept Palaeontol, London SW7 5BD, England
[5] Moscow MV Lomonosov State Univ, Dept Paleontol, Moscow 119992, Russia
[6] Russian Acad Sci, Severtsov Inst Ecol & Evolut, Moscow 119071, Russia
[7] Univ Alaska, Inst Arctic Biol, Fairbanks, AK 99709 USA