[1] Univ Wageningen & Res Ctr, Math Methods & Models Grp, NL-6703 HA Wageningen, Netherlands
[2] Univ Wageningen & Res Ctr, Eros & Soil & Water Conservat Grp, NL-6709 PA Wageningen, Netherlands
[3] Univ Wageningen & Res Ctr, Trop Nat Conservat & Vertebrate Ecol Grp, NL-6708 PD Wageningen, Netherlands
[4] AFRC, Inst Arable Crops Res, Dept Stat, Rothamsted Expt Stn, Harpenden AL5 2JQ, Herts, England
[5] Univ Wageningen & Res Ctr, Nat Conservat & Plant Ecol Grp, NL-6708 PD Wageningen, Netherlands