[1] Univ Cambridge, Dept Zool, Conservat Biol Grp, Cambridge CB2 3EJ, England
[2] Univ Copenhagen, Dept Comp Sci, DK-2100 Copenhagen, Denmark
[3] Univ Copenhagen, Museum Zool, DK-2100 Copenhagen, Denmark
[4] Nat Hist Museum, Biogeog & Conservat Lab, London SW7 5BD, England