[1] Univ London Imperial Coll Sci Technol & Med, Ascot SL5 7PY, Berks, England
[2] Royal Bot Gardens, Jodrell Lab, Richmond TW9 3DS, Surrey, England
[3] Univ Reading, Sch Plant Sci, Ctr Plant Divers & Systemat, Reading RG6 6AS, Berks, England
[4] Univ Costa Rica, Lankester Bot Garden, Cartago, Costa Rica
[5] Univ Lausanne, Dept Ecol & Evolut, CH-1015 Lausanne, Switzerland
[6] Queen Mary Univ London, Sch Biol & Chem Sci, London E1 4NS, England
[7] Univ Florida, Florida Museum Nat Hist, Gainesville, FL 32611 USA