1. a Department of Botany , Nelson Mandela Metropolitan University , PO Box 7700, Port Elizabeth , 6031 , South Africa
2. b South African Environmental Observation Network, Elwandle Node , 18 Somerset Street, Grahamstown , 6140 , South Africa
3. c Evolution and Ecology , University of California , 2320 Storer Hall, Davis, One Shields Avenue, Davis , CA , 95616 , USA