1. Department of Biology, Faculty of Science, Hokkaido University, Kita-ku, Sapporo, Hokkaido, Japan
2. Department of Evolutionary Ecology, Estación Biológica de Doñana-CSIC, c/Americo Vespucio, Seville, Spain
3. MTA-ELTE, Theoretical Biology and Evolutionary Ecology Research Group, Department of Plant Systematics, Ecology and Theoretical Biology, Eötvös Loránd University, Pázmány Péter sétány, Budapest, Hungary