1. Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, UK
2. Institut de Recherche pour le Développement (IRD), UMR-DIADE, 911 Avenue Agropolis, BP 64501, Montpellier 34394, France
3. Kew Madagascar Conservation Centre, Lot II J 131 B Ambodivoanjo, Ivandry, Antananarivo 101, Madagascar
4. Mention Biologie et Ecologie Végétales, Faculté des Sciences, Université d'Antananarivo, Antananarivo BP 906101, Madagascar
5. Department of Biology, Aarhus University, Aarhus, Denmark
6. Molecular Ecology and Evolution Bangor, Environment Centre Wales, School of Natural Sciences, Bangor University, Bangor LL57 2UW, UK