[1] Swedish Univ Agr Sci, Dept Soil & Environm, POB 7014, S-75007 Uppsala, Sweden
[2] Lund Univ, Ctr Environm & Climate Res, S-22362 Lund, Sweden
[3] Swedish Univ Agr Sci, Dept Ecol, POB 7044, S-75007 Uppsala, Sweden
[4] James Hutton Inst, Aberdeen AB15 8QH, Scotland