[1] Univ Exeter, Ctr Ecol & Conservat, Penryn TR10 9EZ, England
[2] Univ Exeter, Environm & Sustainabil Inst, Penryn TR10 9EZ, England
[3] Univ York, Dept Biol, York YO10 5DD, N Yorkshire, England
[4] Univ Liverpool, Dept Evolut Ecol & Behav, Liverpool L69 7ZB, Merseyside, England
[5] Netherlands Inst Ecol NIOO KNAW, Dept Anim Ecol, NL-6700 AB Wageningen, Netherlands
[6] Sussex Wildlife Trust, Henfield BN5 9SD, W Sussex, England
[7] NERC Ctr Ecol & Hydrol, Wallingford OX10 8BB, Oxon, England
[8] Butterfly Conservat, Wareham BH20 5QP, Dorset, England