1. Institute of Genetics, School of Biology, University of Nottingham, Nottingham, NG7 2UH, U.K.
2. Institute of Evolutionary Biology, School of Biological Sciences, University of Edinburgh, Edinburgh, EH9 3JT, U.K.
3. Graduate School of Life Sciences, Tohoku University, Aoba‐ku, Sendai 980 8578, Japan
4. Department of Zoology, The Natural History Museum, London, SW7 5BD, U.K.