1School of Biology, University of St Andrews, Harold Mitchel Building, St Andrews, Fife, KY16 9TH, Scotland
2Japan Society for the Promotion of Science, Kojimachi, Chiyoda-ku, Tokyo, 102-0083, Japan
3Department of Evolutionary Studies of Biosystems, School of Advanced Sciences, SOKENDAI (The Graduate University for Advanced Studies), Shonan Village, Hayama, Kanagawa 240-0193, Japan