[1] Univ Leeds, Ctr Biodivers & Conservat, Leeds LS2 9JT, W Yorkshire, England
[2] Inst Terr Ecol, Biol Records Ctr, Abbots Ripton, Cambs, England
[3] Ctr Populat Biol, Ascot, Berks, England
[4] Univ Sheffield, Dept Anim & Plant Sci, Biodivers & Macroecol Grp, Sheffield S10 2TN, S Yorkshire, England