[1] Univ Edinburgh, Sch Geosci, Edinburgh EH9 3JN, Midlothian, Scotland
[2] Catholic Univ Louvain, Dept Geog, B-1348 Louvain, Belgium
[3] Univ Aberdeen, Sch Biol Sci, Aberdeen AB24 3UU, Scotland
[4] Katholieke Univ Leuven, Dept Land Management & Econ, B-3001 Louvain, Belgium
[5] Observ Midi Pyrenees, CNRS, IRD, LMTG, F-31400 Toulouse, France