1. 2Department of Botany, University of Toronto, 25 Willcocks Street, Toronto, Ontario M5S 3B2, Canada;
2. 4Current address: Department of Ecology and Evolutionary Biology, Cornell University, Corson Hall, Ithaca, New York 14853 USA
3. 3Section of Evolution and Ecology and Center for Population Biology, One Shields Avenue, University of California, Davis, California 95616 USA;