[1] Univ Bourgogne Franche Comte, Biogeosci, UMR 6282, CNRS, 6 Blvd Gabriel, F-21000 Dijon, France
[2] INRAE, UMR, Agroecol 1347, F-21065 Dijon, France
[3] Univ Toulon & Var, AMU, CNRS, IRD,MIO Toulon, CS 60583, F-83041 Toulon 9, France
[4] INRAE, UMR SAS 1069, F-35042 Rennes, France
[5] Museo Nacl Ciencias Nat MNCN CSIC, C Serrano 115-B, Madrid 28006, Spain
[6] Tech Univ Munich, Res Dept Ecol & Ecosyst Management, Chair Soil Sci, 2 Emil Ramann Str, D-85354 Freising Weihenstephan, Germany