Fundação Calouste Gulbenkian - Instituto Gulbenkian de Ciência, University of Mahajanga, affiliation not provided to SSRN, affiliation not provided to SSRN, University of Mahajanga, University of Mahajanga, affiliation not provided to SSRN, University of Toulouse 3, Paul Sabatier University - Laboratoire Évolution & Diversité Biologique - UMR 5174 (EDB), Université de Toulouse, University of Antsiranana, University of Mahajanga, University of Mahajanga and affiliation not provided to SSRN