[1] Univ London Imperial Coll Sci Technol & Med, Dept Biol, Ascot SL5 7PY, Berks, England
[2] Univ London Imperial Coll Sci Technol & Med, NERC, Ctr Populat Biol, Ascot SL5 7PY, Berks, England
[3] Nat Hist Museum, Dept Entomol, London SW7 5BD, England
[4] Univ Bristol, Sch Biol Sci, Bristol BS8 1UG, Avon, England