[1] Microsoft Res Ltd, Computat Ecol & Environm Sci, Cambridge CB3 0FB, England
[2] Ctr Ecol & Hydrol, Banchory AB31 4BW, Kincardine, Scotland
[3] Univ Leeds, Inst Integrat & Comparat Biol, Fac Biol Sci, Leeds LS2 9JT, W Yorkshire, England
[4] Univ Aberdeen, Inst Biol Sci, Aberdeen AB24 2TZ, Scotland