1. Laboratory of Evolutionary Ecology, Graduate School of Environmental Science, Okayama University, Tsushima-naka 111, Okayama 700-8530, Japan
2. Department of Biological Sciences, School of Science, Tokyo Metropolitan University, 1-1, Minamiohsawa Hachiohji, Tokyo, 192-0397, Japan
3. Centre for Ecology and Conservation, University of Exeter, Cornwall, Penryn, Cornwall TR10 9EZ, UK