[1] Univ Cambridge, Dept Zool, Cambridge CB2 3EJ, England
[2] Univ Stirling, Dept Biol & Mol Sci, Stirling FK9 4LA, Scotland
[3] Zool Soc London, Inst Zool, London NW1 4RY, England
[4] Scottish Nat Heritage, Stilligarry HS8 5RS, Scotland
[5] Inst Terr Ecol, Banchory AB31 4BY, Kincardine, Scotland
[6] Univ Edinburgh, Inst Cell Anim & Populat Biol, Edinburgh EH9 3JT, Midlothian, Scotland
[7] Univ London Imperial Coll Sci Technol & Med, Ascot SL5 7PY, Berks, England