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Hokkaido Univ, Grad Sch Vet Med, Dept Environm Vet Sci, Lab Wildlife Biol & Med, Sapporo, Hokkaido 0600818, Japan Hokkaido Univ, Grad Sch Vet Med, Dept Environm Vet Sci, Lab Wildlife Biol & Med, Sapporo, Hokkaido 0600818, Japan