[1] Univ London Imperial Coll Sci Technol & Med, Dept Biol Sci, London SW7 2AZ, England
[2] Syngenta, Jealotts Hill Res Ctr, Bracknell RG42 6EY, Berks, England
[3] NERC, Ctr Ecol & Hydrol, Oxford OX1 3SR, England
[4] Wageningen Univ, Virol Lab, NL-6709 PD Wageningen, Netherlands