[1] Royal Bot Gardens, Richmond TW9 3AE, Surrey, England
[2] Queen Mary Univ London, Sch Biol & Chem Sci, Mile End Rd, London E1 4NS, England
[3] Kew Madagascar Conservat Ctr, Lot 2 J 131 B Ambodivoanjo, Antananarivo 101, Madagascar
[4] Univ Antananarivo, BP 906, Antananarivo, Madagascar
[5] Parc Bot & Zool Tsimbazaza,BP 4096,Rue Kasanga, Antananarivo, Madagascar
[6] Univ Ghent, Dept Biol, Systemat & Evolutionary Bot Lab, KL Ledeganckstr 35, B-9000 Ghent, Belgium