[1] Univ Antananarivo, Fac Sci, Ment Biol & Ecol Vegetales, BP 906, Antananarivo 101, Madagascar
[2] Madagascar Res & Conservat Program, Missouri Bot Garden, BP 3391, Antananarivo 101, Madagascar
[3] Missouri Bot Garden, 4344 Shaw Blvd, St Louis, MO 63110 USA
[4] Univ Antilles, Sorbonne Univ, Ecole Prat Hautes Etud, Museum Natl Hist Nat,Ctr Natl Rech Sci,Inst Syste, CP 39,57 Rue Cuvier, F-75005 Paris, France