1. Department of Polar Science, The Graduate University for Advanced Studies, Sokendai, 10-3 Midori-cho, Tachikawa, Tokyo 190-8518, Japan
2. National Institute of Polar Research, 10-3 Midori-cho, Tachikawa, Tokyo 190-8518, Japan
3. School of Biology, University of St Andrews, Scottish Oceans Institute, East Sands, St Andrews, Fife KY16 9TS UK
4. Department of Ecology and Evolutionary Biology, Institute of Marine Science, University of California Santa Cruz, Santa Cruz CA, 95060, USA