[1] Univ Cambridge, Dept Zool, Large Anim Res Grp, Cambridge CB2 3EJ, England
[2] Univ London Imperial Coll Sci Technol & Med, Royal Sch Mines, TH Huxley Sch Environm Earth Sci & Engn, London SW7 2BP, England
[3] Scottish Nat Heritage, Edinburgh EH6 5NP, Midlothian, Scotland