[1] Royal Holloway Univ London, Sch Biol Sci, Egham TW20 0EX, Surrey, England
[2] Royal Holloway Univ London, Dept Geog, Egham TW20 0EX, Surrey, England
[3] Swedish Museum Nat Hist, Dept Mol Systemat, S-10405 Stockholm, Sweden
[4] Stockholm Univ, Dept Zool, S-10691 Stockholm, Sweden
[5] Nat Hist Museum, Dept Earth Sci, London SW7 5BD, England
[6] Univ Liege, Serv Prehist, B-4000 Liege, Belgium
[7] Royal Belgian Inst Nat Sci, Dept Palaeontol, B-1000 Brussels, Belgium
[8] Bournemouth Univ, Sch Appl Sci, Poole BH12 5BB, Dorset, England