[1] Univ Paris 06, Sorbonne Univ, LOCEAN Lab, CNRS,IRD,MNHN, F-75005 Paris, France
[2] Univ Toulon & Var, Aix Marseille Univ, MIO UM110, CNRS,IRD, F-13288 Marseille, France
[3] Vrije Univ Brussel, B-1050 Brussels, Belgium
[4] Royal Museum Cent Africa, Dept Earth Sci, B-3080 Tervuren, Belgium