[1] Fisheries Res Agcy, Natl Res Inst Fisheries Sci, Freshwater Fisheries Res Div, Ueda, Nagano 3860031, Japan
[2] Osaka City Univ, Lab Anim Sociol, Dept Biol & Geosci, Grad Sch Sci, Osaka 558, Japan
[3] Ehime Univ, Ctr Marine Environm Studies, Matsuyama, Ehime, Japan
[4] Kyoto Univ, Anim Ecol Lab, Grad Sch Sci, Kyoto, Japan
[5] Univ Bern, Dept Behav Ecol, Inst Ecol & Evolut, Hinterkappelen, Switzerland