[1] Royal Bot Gardens, Richmond TW9 3DS, Surrey, England
[2] Ctr Evolutionary & Funct Ecol, UMR 5175, F-34293 Montpellier 5, France
[3] Univ Aarhus, Dept Systemat Bot, DK-8000 Aarhus, Denmark
[4] Univ Lausanne, Dept Ecol & Evolut, CH-1015 Lausanne, Switzerland