[1] Hokkaido Univ, Grad Sch Sci, Dept Nat Hist Sci, Sapporo, Hokkaido, Japan
[2] Hokkaido Univ, Grad Sch Agr, Frontiers Environm Sci, Sapporo, Hokkaido, Japan
[3] Russian Acad Sci, Fed Sci Ctr East Asia Terr Biodivers, Lab Evolutionary Zool & Genet, Far Eastern Branch, Vladivostok, Russia
[4] Hokkaido Univ, Fac Sci, Dept Nat Hist Sci, Sapporo, Hokkaido, Japan