[1] Univ York, Dept Biol, Stockholm Environm Inst, SEI York Ctr, York YO10 5DD, N Yorkshire, England
[2] Univ York, Dept Biol, York YO10 5YW, N Yorkshire, England
[3] Alice Holt Lodge, Forest Res, Biometr Survey & Stat Div, Farnham GU10 4LH, Surrey, England
[4] Univ Jaen, Dept Biol Anim Biol Vegetal & Ecol, Jaen 23071, Spain