[1] Univ Aberdeen, Inst Biol & Environm Sci, 23 St Machar Dr, Aberdeen AB24 3UU, Scotland
[2] INRA, 147 Rue Univ, F-75338 Paris 07, France
[3] Lawrence Berkeley Natl Lab, Earth & Environm Sci Div, Berkeley, CA 94720 USA
[4] Univ Zurich, Dept Geog, CH-8057 Zurich, Switzerland
[5] Univ Zurich, Res Prior Program Global Change & Biodivers, URPP, GCB, CH-8057 Zurich, Switzerland