[1] Royal Bot Gardens, Richmond TW9 3AB, Surrey, England
[2] Museum Natl Hist Nat, UPMC, EPHE, Inst Systemat Evolut Biodivers,CNRS,UMR 7205, F-75231 Paris, France
[3] Parc Bot & Zool Tsimbazaza, Dept Flore, Herbier, Antananarivo 101, Madagascar
[4] Univ Sheffield, Dept Anim & Plant Sci, Sheffield S10 2TN, S Yorkshire, England
[5] Univ Toulouse, ENFA, EDB Lab Evolut & Divers Biol, CNRS,UMR5174, F-31062 Toulouse, France