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Aix Marseille Univ, CNRS, Equipe Evolut Genome & Environm, Ctr St Charles,IRD,IMEP,UMR 6116, F-13331 Marseille 3, France Aix Marseille Univ, CNRS, Equipe Evolut Genome & Environm, Ctr St Charles,IRD,IMEP,UMR 6116, F-13331 Marseille 3, France