1. Department of Ecology and Evolutionary Biology, University of Toronto , 25 Willcocks Street, Toronto, ON, M5S 3B2 , Canada
2. Department of Ecology and Evolutionary Biology, Yale University , New Haven, CT 06511 , USA
3. Department of Biology, University of Oxford , South Parks Road, Oxford, OX1 3RB , UK
4. Department of Botany and Plant Sciences, University of California, Riverside , CA 92521 , USA