[1] Agr Dev & Advisory Serv, Wolverhampton WV9 5AP, W Midlands, England
[2] Univ Southampton, Southampton SO17 1BJ, Hants, England
[3] Rothamsted Res North Wyke, Dept Sustainable Syst & Grassland Sci, Okehampton EX20 2SB, Devon, England
[4] Queen Mary Univ London, Sch Biol & Chem Sci, London E1 4NS, England
[5] CEH Wallingford, Wallingford OX10 8BB, Oxon, England