༤ኈㄽᩥ ᕞᮾ㒊࠾ࡅࡿ࣐ࢦࡢಖ㛵ࡍࡿ◊✲ ᖹᡂ 27 ᖺ 9 ᭶ ᗈᓥᏛᏛ㝔⏕≀ᅪ⛉Ꮫ◊✲⛉ ⏕≀㈨※⛉Ꮫᑓᨷ ᮌᮏᆂ㍜ ┠ ḟ ➨ 1 ❶ ⥴ゝ 㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃 1 ➨ 2 ❶ ὶ⛬ἢࡗࡓ㨶㢮⩌㞟ࡢኚ 㺃㺃㺃㺃㺃㺃㺃 8 ➨ 3 ❶ ࣐ࢦᾋୖ⛶㨶ࡢὶ⛬ศᕸ 㺃㺃㺃㺃㺃㺃㺃㺃㺃 28 ➨ 4 ❶ ࣐ࢦᾋୖ⛶㨶ࡢ⏕ᜥሙᡤ⏝ 㺃㺃㺃㺃㺃 50 ➨ 5 ❶ ࣐ࢦಶయ⩌ࡢ㑇ఏⓗᵓ㐀 㺃㺃㺃㺃㺃㺃㺃㺃㺃 74 ➨ 6 ❶ ⥲ྜ⪃ᐹ 㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃 91 せ⣙ 㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃 102 ㅰ㎡ 㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃 106 ᘬ⏝ᩥ⊩ 㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃㺃 107 ➨ 1 ❶ ⥴ゝ ࣐ࢦ Oncorhynchus masou ishikawae㸦Figs. 1-1, 1-2㸧ࡣࢧࢣ⛉ᒓࡍࡿ᪥ᮏ ᅛ᭷ࡢள✀࡛࠶ࡾ㸪⚄ዉᕝ┴௨すࡢᮏᕞኴᖹὒᓊ㸪ᅄᅜᇦ㸪ᕞࡢ℩ᡞෆᾏ ὶධἙᕝ⮬↛ศᕸࡍࡿ㸦ᓥ㸪1957㸧㸬ᮏள✀ࡣ㐳Ἑᅇ㐟㨶࡛࠶ࡾ㸪ᮏᕞࡣ 㝆ᾏᆺಶయ⩌࡛࠶ࡿࢧࢶ࣐࢟ࢫࡀࡳࡽࢀࡿࡀ㸦୰㔝㸪1993㸧 㸪ศᕸ༡㝈ᇦ࡛ ࠶ࡿᕞࡣἙᕝෆ࡛⏕άྐࢆ⤖ࡍࡿἙᕝᆺಶయ⩌ࡔࡅࡀ⏕ᜥࡍࡿ㸦Kimura, 1989㸹๓ᕝ࣭୰㔝㸪1994㸧 㸬୍᪉㸪࣐ࢦࡀ⏕ᜥࡋ࡞࠸ᾏ㐨ࡽᕞ⮳ࡿᅜ ෆࡢᗈ࠸⠊ᅖࡣ㸪ᇶள✀࡛࠶ࡿࢧࢡ࣐ࣛࢫ㸦Ἑᕝᆺࡣ࣐࣓ࣖ㸧Oncorhynchus masou masou ࡀ⏕ᜥࡍࡿ㸦ᓥ㸪1957㸧㸬㏆ᖺ㸪࣐ࢦࡣࡢἙᕝᛶࢧࢣ⛉㨶㢮 ྠᵝ㸪◁㜵ࢲ࣒ࡢタ⨨㸪Ἑ␁ᯘࡢఆ᥇㸪Ỉ㉁ởᰁ➼ࡼࡿ⏕ᜥ⎔ቃࡢᝏ 㸦๓ᕝ㸪2000㸹㧗ᶫ㸪2000㸹Yoshimura et al., 2005㸧 㸪⁺ᴗ࣭㐟⁺ࡼࡿ⋓➼ ࡼࡾῶᑡࡋ࡚࠸ࡿ㸦୰㔝㸪1993㸹୰ᮧ㸪2009㸧 㸬 ࣐ࢦࡣ㣗ᮦࡋ࡚ࡢ౯್ࡸ⨾ࡋ࠸ጼែࡢࡓࡵ⁺ᴗ࣭㐟⁺࠾ࡅࡿ⏝౯ ್ࡀ㧗ࡃ㸦Kimura, 1989㸧 㸪ከࡃࡢ⏕ᜥἙᕝ࠾࠸࡚ෆỈ㠃⁺ᴗ༠ྠ⤌ྜࡢ⁺ᴗ ᶒᑐ㇟㨶✀࡞ࡗ࡚࠸ࡿ㸦㣤⏣࣭୰ᮧ㸪2009㸧 㸬᪉㸪ෆỈ㠃ࡢỈ⏘ື≀ࡢ㈨※ 㔞ࡣᾏ㠃ẚ࡚ᑡ࡞ࡃ㸪⋓ࡼࡗ࡚ᐜ᫆ᯤῬࡍࡿࡓࡵ㸪⁺ᴗἲ࡛ࡣ⁺ᴗ ᶒࡢචチࢆཷࡅࡓෆỈ㠃⁺ᴗ༠ྠ⤌ྜᑐࡋ࡚㸪⁺ᴗᶒ㨶✀ࢆ✚ᴟⓗቑṪࡍ ࡿ⩏ົࢆㄢࡋ࡚࠸ࡿ㸦㣤⏣࣭୰ᮧ㸪2009㸧㸬࣐ࢦࢆྵࡴᅾ᮶ࢧࢣ⛉㨶㢮࡛ࡣ㸪 ✀ⱑ⏕⏘ᢏ⾡ࡀ☜❧ࡉࢀࡓ 1970 ᖺ௦௨㝆㸪 ࠾ࡶ㣴Ṫ㨶ࡢᨺὶࡼࡗ࡚ቑṪ⩏ ົࡀᒚ⾜ࡉࢀ㸪㈨※ࡢ⥔ᣢ࣭ቑṪຠᯝࢆୖࡆ࡚ࡁࡓ㸦❧ᕝ࣭ᮏⲮ㸪1976㸧㸬 ୍᪉㸪ࢧࢣ⛉㨶㢮࡛ࡣᵝࠎ࡞ᙧ㉁㛵ࡋ࡚ᒁᡤ㐺ᛂࡢᏑᅾࡀሗ࿌ࡉࢀ࡚࠾ࡾ 㸦Taylor, 1991㸹Garcia de Leaniz et al., 2007㸹Kikko et al., 2008㸹Fraser et al., 2011㸹 Primmer, 2011㸹Drinan et al., 2012㸧㸪ᨺὶࡉࢀࡓ㣴Ṫ㨶ࡣ㸪㔝⏕ಶయ⩌ࡢ୰㐺 ᛂⓗ࡛࡞࠸㑇ఏᏊࢆὶධࡉࡏ㸪␗⣔㓄ᙅໃࡸᒁᡤ㐺ᛂࡢᔂቯࢆࡶࡓࡽࡍྍ⬟ ᛶࡀᣦࡉࢀ࡚࠸ࡿ㸦Miller et al., 2004㸹Garcia de Leaniz et al., 2007㸹Tymchuk et al., 2007㸧㸬㣴Ṫ㨶ࡢᨺὶࡀ㛗ᮇࢃࡓࡾᅜⓗつᶍ࡛ᐇࡉࢀࡓ⤖ᯝ㸪⌧ᅾ࡛ ࡣ㸪࣐ࢦࢆྵࡴᅜෆࡢἙᕝᛶࢧࢣ⛉㨶㢮ࡢᅾ᮶ಶయ⩌㸦ᨺὶࡉࢀࡓ㣴Ṫ㨶 1 㓄ࡋ࡚࠸࡞࠸ᆅࡁࡢಶయ⩌㸧ࡣ⤯⁛ࡢ༴ᶵ࠶ࡾ㸪⮬↛ࡸࢲ࣒࡛㝸㞳ࡉ ࢀࡓἙᕝ※ὶ㒊ࡔࡅṧᏑࡍࡿ㸦Kawamura et al., 2007㸹Kubota et al., 2007㸹Sato et al., 2010㸧 㸬ࡲࡓ㸪እ㒊ᙧែࡀ㢮ఝࡍࡿ࣐ࢦ࣐࣓࡛ࣖࡣ㸪⮬↛ศᕸᇦࡀ༑ ศ⪃៖ࡉࢀࡎᨺὶࡀ⾜ࢃࢀࡓࡓࡵ㸪⌧ᅾ࡛ࡣ୧ள✀ࡢศᕸࡀⴭࡋࡃࢀ࡚ ࠸ࡿ㸦Kimura, 1989㸹୰㔝㸪1993㸧 㸬ࡉࡽ㸪ᨺὶࡉࢀࡓ㣴Ṫ㨶ࡣ㸪➇தࢆ ㏻ࡌ࡚㔝⏕ࡢࢧࢣ⛉㨶㢮ࡢᡂ㛗࣭⏕ṧࢆపୗࡉࡏࡿ㸦Nakamura and Doi, 2014㸧㸬 ࡇࡢࡼ࠺㣴Ṫ㨶ࡢᨺὶࡣ㸪࣐ࢦࢆྵࡴ㔝⏕ࡢἙᕝᛶࢧࢣ⛉㨶㢮ࡢᏑ⥆ᝏ ᙳ㡪ࢆ࠼ࡿྍ⬟ᛶࡀ᫂ࡽ࡞ࡗ࡚ࡁࡓࡇࡽ㸪㛗ᮇⓗ࡞ಶయ⩌ࡢ⥔ᣢࡢ ࡓࡵࡣᐇࡍࡁ࡛࡞࠸⪃࠼ࡽࢀࡿ㸬ࡋࡋ㸪ෆỈ㠃㈨※ࡢ⬤ᙅᛶࢆ⪃៖ ࡍࡿ㸪⁺ᴗ࣭㐟⁺ࡼࡿ▷ᮇⓗ࡞⏝ࡢࡓࡵࡣᨺὶࡀᚲせ࡛࠶ࡿ㸬 ㏆ᖺ㸪Ἑᕝᛶࢧࢣ⛉㨶㢮ࡢಖ⏝ࡢ୧❧ࢆᅗࡿࡓࡵࡢ᪉ἲࡋ࡚㸪ࢰ࣮ ࢽࣥࢢࡼࡿ⁺ሙ⟶⌮ࡀὀ┠ࡉࢀ࡚࠸ࡿ㸦ຍ⸨㸪1990㸹ᒣᮏ㸪1991㸹ᮌᮧ㸪2004㸹 ୰ᮧ㸪2009㸧㸬ࢰ࣮ࢽࣥࢢࡣ㸪ᅾ᮶ಶయ⩌ࡢ⏕ᜥỈᇦ⮬↛⏕⏘ࡢᐇែᇶ ࡙࠸࡚Ἑᕝࢆ༊ศࡋ㸪ࡑࢀࡒࢀࡢỈᇦࡢ⮬↛᮲௳♫᮲௳ྜࢃࡏ࡚ᨺὶ ⁺ᴗ࣭㐟⁺⟶⌮ᡭἲࢆ࠸ศࡅࡿ࠸࠺⪃࠼᪉࡛࠶ࡿ㸦୰ᮧ㸪2009㸧 㸬ᮏ◊✲ࡣ㸪 ࢰ࣮ࢽࣥࢢ⟶⌮ࢆ⌧ሙᑟධࡍࡿ♫ᐇ㦂ࡋ࡚Ỉ⏘ᗇ᱂ᇵ㣴Ṫㄢࡀ 2003– 2007 ᖺᗘᐇࡋࡓࠕ࡞ෆỈ㠃⏕ែ⣔ඖ᥎㐍ጤクᴗࠖࡢ࠺ࡕࠕὶᇦ ⟶⌮యไᵓ⠏ᴗࠖࢆศ┴ࡀཷクࡋ㸪ᙜࡢศ┴ᾏὒỈ⏘◊✲ࢭࣥࢱ࣮ෆ Ỉ㠃◊✲ᡤ㸦2003–2004 ᖺᗘ㸧ศ┴㎰ᯘỈ⏘◊✲ࢭࣥࢱ࣮Ỉ⏘ヨ㦂ሙෆỈ 㠃◊✲ᡤ㸦2005–2007 ᖺᗘ㸧ᡤᒓࡋ࡚࠸ࡓ➹⪅ࡽࡀᐇࡋࡓࡶࡢ࡛࠶ࡿ㸬 ᮏ◊✲ඛ❧ࡕ㸪 ศ┴ෆࡢෆỈ㠃⁺ᴗ༠ྠ⤌ྜࢆᑐ㇟ࣥࢣ࣮ࢺࢆ⾜࠸㸪 ᅾ᮶࣐ࢦಶయ⩌ࡢᏑྰࢆᑜࡡࡓࡇࢁ㸪၏୍㸪㔝ᕝỈ⣔ࡢ୍ᨭὶ࡛࠶ࡿ⚄ ཎᕝࡢ※ὶ㒊⏕ᜥࡍࡿྍ⬟ᛶࡀ♧ࡉࢀࡓ㸦ᚨගಇ㹙ศ┴ᾏὒỈ⏘◊✲ࢭ ࣥࢱ࣮ෆỈ㠃◊✲ᡤ㹛㸪ᮍⓎ⾲㸪1999 ᖺ㸧 㸬㔝ᕝỈ⣔ࡣᕞ࠾ࡅࡿ࣐ࢦࡢ ⮬↛ศᕸᇦࡢ༡㝈࡛࠶ࡾ㸦ᓥ㸪1957㸧 㸪ࡑࡢ※ࢆ㜿⸽እ㍯ᒣ࠾ࡼࡧஂఫᒣⓎ ࡋ㸪ศ┴ࡢ୰ኸ㒊ࢆὶࡋ࡚℩ᡞෆᾏ㸦ูᗓ‴㸧ὀࡄ㸪ὶ㊰ᘏ㛗 128 km㸪 ὶᇦ㠃✚ 1,455 km2㸪ᖹᆒὶ㔞 300 t/᪥ࡢ┴ෆ᭱ࡢἙᕝ࡛࠶ࡿ㸦ᚿ㈡࣭ᕝ㔝㸪 1977㸧㸬ᮏỈ⣔ᒓࡍࡿ⚄ཎᕝࡣ㸪ศ┴㸪ᐑᓮ┴㸪⇃ᮏ┴ࡢ┴ቃ⨨ࡍࡿ♽ 2 ẕᒣ㸦ᶆ㧗 1,756 m㸧ࡢᩳ㠃Ⓨࡋ㸪⣙ 5 km ୗὶ࡛♽ẕᒣࡢᮾᩳ㠃Ỉ※ ࢆࡶࡘἼᮌྜᕝྜὶࡋࡓࡢࡕ㸪 ⣙ 5 km ὶࡋ࡚⥴᪉ᕝྜὶࡍࡿ㸦Fig. 1-3㸧㸬 ⥴᪉ᕝࡣ➉⏣ᕷࢆᮾ᪉ྥὶࢀ㸪 ⣙ 7 km ୗὶ࡛ᨭὶ༑ゅᕝྜὶࡋࡓࡢࡕ㸪 ᮾὶࡋ࡚Ἑཱྀࡽ 52 km ᆅⅬࡢ㔝ᕝྜὶࡍࡿ㸬⚄ཎᕝ⥴᪉ᕝ࡛ࡣ㸪ᅾ᮶ ࢧࢣ⛉㨶㢮ࡢ✀ⱑ⏕⏘ᢏ⾡ࡢ☜❧௨๓ࡽ࣐ࢦࡢ⮬↛ศᕸࡀ☜ㄆࡉࢀ࡚࠸ࡿ 㸦ᮌᮧ㸪1959㸧 㸬㞄᥋ᨭὶࡢἼᮌྜᕝ※ὶ㒊㸦࣓ࣥࣀࢶࣛ㇂㸧ࡣ㸪࣐ࢦࡢ↓ ᩬᆺ࡛࠶ࡿ࣓࣡㸦Fig. 1-1㸧ࡀ⏕ᜥࡋ࡚࠸ࡿ㸦Kimura and Nakamura, 1961㸧 㸬 ᮏ◊✲࡛ࡣ㸪⚄ཎᕝ⥴᪉ᕝࡢ࣐ࢦ⏕ᜥᇦయࢆ⥙⨶ࡍࡿࡓࡵ㸪Ἑཱྀࡽ 70 km ᆅⅬ࡞ࡿ༑ゅᕝࡢྜὶⅬࡽ⚄ཎᕝࡢྜὶⅬࡲ࡛ࡢ⥴᪉ᕝ㸪⥴᪉ ᕝࡢྜὶⅬࡽ㨶㢮⏕ᜥୖ㝈㸦ᮌᮧ㸪1959㸹ᮌᮏ㸪ᮍⓎ⾲㸧࡛࠶ࡿྜ ┠㸦ⴠᕪ 11 m㸧ࡲ࡛ࡢ⚄ཎᕝ㸪࠾ࡼࡧᨭὶἼᮌྜᕝࡢ୍㒊༊㛫ࢆㄪᰝᑐ㇟Ỉ ᇦࡋࡓ㸦Fig. 1-3㸧㸬ᮏㄪᰝỈᇦࡣἙᕝ※ὶ㒊ࡽᡪ≧ᆅཬࡪࡓࡵ㸦ὶ⛬㛗 ⣙ 16.4 km㸧㸪ࡑࡢᬒほࡣὶ⛬ἢࡗ࡚ࡁ࡞ኚࢆ♧ࡍ㸦㧗ᶫ࣭ኴ⏣㸪2000㸹 ⴛሙ㸪2013㸧 㸬ୗὶ➃ࡢᶆ㧗ࡣ⣙ 230 m ࡛㸪࿘㎶ࡢἙᗋ໙㓄ࡣ 1%๓ᚋ㸪Ἑᕝᙧ ែᆺ㸦ྍඣ㸪1978㸧ࡣ Bb ᆺࢆ♧ࡋ㸪୧ᓊࡣ㛤ࡅ࡚Ỉ⏣⏝ࡉࢀ࡚࠸ࡿ㸬⥴ ᪉ᕝࡢἢᓊࡣỈ⩌ࡀⅬᅾࡋ㸪ㄪᰝỈᇦୗὶഃࡢỈ ᙳ㡪ࢆ࠼࡚࠸ࡿ㸬 ⥴᪉ᕝࡢྜὶⅬࡼࡾୖὶࡢ⚄ཎᕝὶᇦࡣࠕ⚄ཎ㇂ࠖࡤࢀ㸪Ἑᕝᙧែᆺ ࡣ Aa-Bb ⛣⾜ᆺࢆ♧ࡋ㸪୧ᓊࡀ㏕ࡗ࡚㇂ࡢᵝ┦ࢆ࿊ࡍࡿ㸬⚄ཎᕝἼᮌྜᕝ ࡢྜὶⅬࡢᶆ㧗ࡣ⣙ 400 m ࡛㸪ࡇࢀࡼࡾୖὶ࡛ࡣἙᗋ໙㓄ࡀࡁࡃ㸪Ἑᕝᙧែ ᆺࡀ Aa ᆺࢆ♧ࡍᆺⓗ࡞ᒣᆅὶ࡞ࡿ㸬ࡇࡢỈᇦࡣ⮬↛ࡀぢࡽࢀࡿ 㸪ሙᡤࡼࡗ࡚ࡣከᩘࡢ◁㜵ࢲ࣒➼ࡀタ⨨ࡉࢀ࡚࠸ࡿ㸬ୖὶ➃ࡢᶆ㧗ࡣ⣙ 820 m㸪㏆ࡢἙᗋ໙㓄ࡣ⣙ 20%࡛࠶ࡿ㸬➹⪅ࡽࡣ㸪ᮏㄪᰝỈᇦࢰ࣮ࢽࣥࢢ⟶⌮ ࢆᑟධࡍࡿࡓࡵᅾ᮶࣐ࢦಶయ⩌ࡢ⏕ᜥỈᇦ⮬↛⏕⏘ࡢᐇែ➼ࢆㄪࡓ㸬 ㄪᰝ⤖ᯝᇶ࡙࠸࡚సᡂࡉࢀࡓࢰ࣮ࢽࣥࢢࡣ㸪ᆅඖఫẸ㸪㔝ᕝ⁺ᴗ༠ྠ⤌ ྜ㸪㐟⁺⪅㸪ᆅඖ⾜ᨻ㸦➉⏣ᕷ㸧➼ࢆᵓᡂဨࡍࡿጤဨ࡛༠㆟ࡉࢀ㸪᭱⤊ⓗ ᅾ᮶ಶయ⩌ಖࡢࡓࡵࡢ⚗⁺༊ࡀタ⨨ࡉࢀࡓ㸦ᮌᮏ㸪2008㸧 㸬ᮏ༤ኈㄽᩥࡣ㸪 ࢰ࣮ࢽࣥࢢࢆసᡂࡍࡿࡓࡵᐇࡋࡓ」ᩘࡢ◊✲ᡂᯝࢆྲྀࡾ⧳ࡵࡓࡶࡢ࡛࠶ ࡾ㸪ᮏ❶㸦➨ 1 ❶㸪ᗎㄽ㸧ࢆྵࡴ 6 ❶ࡽᵓᡂࡉࢀࡿ㸬 3 ᮏㄪᰝỈᇦࡢࡼ࠺㸪※ὶ㒊ࡽᡪ≧ᆅࡅ࡚ࡁ࡞⎔ቃࡢኚࡀぢࡽࢀ ࡿሙᡤ࡛ࡣ㸪ୖὶഃࡢࢧࢣ⛉㨶㢮ࢆ୰ᚰࡍࡿ㨶㢮⩌㞟ࡽୗὶഃࡢࢥ⛉㨶 㢮ࢆ୰ᚰࡍࡿ⩌㞟ࡢ⛣⾜ࡀ⏕ࡌࡿࡇࡀ▱ࡽࢀ࡚࠸ࡿ㸦Rahel and Hubert, 1991㸹Torgersen et al., 2006㸧 㸬ࡲࡓ㸪⩌㞟ࡢ⛣⾜ࡀ⏕ࡌࡿỈᇦ㸦⛣⾜ᖏ㸧࡛ࡣ㸪 ᵓᡂ✀ࡀྛࠎࡢศᕸࡢ࿘⦕㒊⏕ᜥࡋ࡚࠸ࡿࡓࡵ㸪⎔ቃᨵኚᑐࡍࡿឤཷᛶࡀ 㧗࠸ᣦࡉࢀ࡚࠸ࡿ㸦Naiman et al., 1988㸹Aarts and Nienhuis, 2003㸹Lasne et al., 2007㸧㸬ࢰ࣮ࢽࣥࢢ⟶⌮ࡣ㸪ᑐ㇟㨶✀ࡢ⮬↛⏕⏘ࡢᐇែᇶ࡙࠸࡚⾜ࢃࢀࡿࡀ 㸦୰ᮧ㸪2009㸧 㸪⮬↛⏕⏘ࢆྵࡴࢧࢣ⛉㨶㢮ࡢ⏕άྐ⯡ࡣ㸪ࡇ࠺ࡋࡓࡁ࡞ ✵㛫ࢫࢣ࣮ࣝࡢ⎔ቃ໙㓄ࡁࡃつᐃࡉࢀࡿ⪃࠼ࡽࢀࡿ㸬➨ 2 ❶㸦ὶ⛬ἢ ࡗࡓ㨶㢮⩌㞟ࡢኚ㸧࡛ࡣ㸪⏕ᜥᇦయࢃࡓࡿ࣐ࢦࡢὶ⛬ศᕸࡢ≉ᚩࢆᢕ ᥱࡍࡿࡓࡵ㸪ὶ⛬ἢࡗࡓ㨶㢮⩌㞟ࡢኚ⎔ቃኚ㔞ࡢ㛵ಀࢆㄪࡓ㸬 ➨ 3 ❶㸦࣐ࢦᾋୖ⛶㨶ࡢὶ⛬ศᕸ㸧࡛ࡣ㸪ᮏㄪᰝỈᇦ࠾ࡅࡿ࣐ࢦࡢ⮬ ↛⏕⏘ࡢᐇែࢆᢕᥱࡍࡿࡓࡵ㸪ὶ⛬ࡢ₯Ỉ┠どࡼࡾᾋୖ⛶㨶㸦Fig. 1-2㸧 ࡢὶ⛬ศᕸࢆㄪࡓ㸬ࡲࡓ㸪࣐ࢦᾋୖ⛶㨶ᗂ㨶ᮇ௨㝆ࡢಶయࡢὶ⛬ศᕸࢆ ẚ㍑ࡋ㸪୧⪅ࡢ㛵ಀᇶ࡙࠸࡚ᾋୖ⛶㨶ࡢ⛣ືᵝᘧࡘ࠸࡚ࡶ⪃ᐹࡋࡓ㸬 ➨ 4 ❶㸦࣐ࢦᾋୖ⛶㨶ࡢ⏕ᜥሙᡤ⏝㸧࡛ࡣ㸪➨ 3 ❶࡛⮬↛⏕⏘ࡀపㄪ ᥎ᐃࡉࢀࡓୗὶഃỈᇦ࡛㸪⮬↛⏕⏘ࡸቑṪࢆಁ㐍ࡍࡿࡓࡵࡢᇶ♏ⓗ▱ぢࢆ ᚓࡿࡇࢆ┠ⓗ㸪ὶ㊰༢ࢫࢣ࣮ࣝ㸦Frissell et al., 1986㸧࠾ࡅࡿᾋୖ⛶㨶 ࡢ⏕ᜥሙᡤ⏝ࢆㄪࡓ㸬ᮏ❶ࡣࢰ࣮ࢽࣥࢢᑟධᚋࡢᑐ⟇ࡋ࡚ᐇࡋࡓ㸬 ➨ 5 ❶㸦࣐ࢦಶయ⩌ࡢ㑇ఏⓗᵓ㐀㸧࡛ࡣ㸪ᅾ᮶࣐ࢦಶయ⩌ࡢ⏕ᜥỈᇦࢆ ≉ᐃࡍࡿࡓࡵ㸪⁺ᴗ༠ྠ⤌ྜࡢࣥࢣ࣮ࢺㄪᰝ࡛ᣦࡉࢀࡓ⚄ཎᕝ※ὶ㒊ࡢ ࣐ࢦಶయ⩌ຍ࠼㸪㞄᥋ᨭὶἼᮌྜᕝࡢ࣓ࣥࣀࢶࣛ㇂ࡢ࣓࣡ಶయ⩌࠾ࡼࡧ Ἴᮌྜᕝࡲࢇࡾࡻ࠺㇂ࡢ࣐ࢦಶయ⩌ࡢ㑇ఏⓗᵓ㐀ࢆ mtDNA ࡼࡾゎᯒࡋࡓ㸬 ➨ 6 ❶ࡣ⥲ྜ⪃ᐹࡋ࡚㸪ྛ❶ࡢෆᐜࢆᕞᮾ㒊࠾ࡅࡿ࣐ࢦࡢಖ ࠸࠺┠ⓗἢࡗ࡚ᨵࡵ࡚ホ౯㸪ゎ㔘ࡍࡿࡶ㸪ྛ❶࡛ಶูᚓࡽࢀࡓ⤖ᯝ ࡢ㛵㐃ᛶࢆ᳨ウࡋ㸪ᮏㄪᰝỈᇦ࠾ࡅࡿ࣐ࢦಖ⟇ࡢయീࢆᑟฟࡋࡓ㸬 ࡞࠾㸪ᮏㄽᩥࡢෆᐜࡢ୍㒊ࡣ㸪ᮌᮏ㸦2013㸪2015㸪༳ๅ୰㸧࠾ࡼࡧ Kimoto et al.㸦༳ๅ୰㸧ࡼࡗ࡚Ꮫ⾡㞧ㄅࡍ࡛බหࡉࢀ࡚࠸ࡿ㸬 4 Fig. 1-1. Two fluvial forms of the red-spotted masu salmon (Oncorhynchus masou ishikawae), amago (top; 142 mm in fork length) and iwame (bottom; 179 mm in fork length), captured by bait fishing from the uppermost location of the Kohbaru Stream in September 2007 and the Mennotsura-dani of the Hakiai Stream in February 2008, respectively. 5 Fig. 1-2. Newly emerged fry of amago captured by dip nets from the Kohbaru Stream in March 2004 (top; 25.9 mm in fork length) and photographed underwater in February 2006 (middle) and 2007 (bottom), respectively. 6 Û( Û( Û(ޗ Û(ޗ Ûޗ1 Fukuoka Pref. Oita Prefecture Sea of Japan Seto Inland Sea Ono River Japan Kumamoto Pref. Û1 Pacific Ocean Ûޗ1 Mt. Sobo 1 Ogata River Tosumi River 1 km Kohbaru Stream Hakiai Stream Hiibachi Falls Manyrou-dani Mennotsura-dani Yoroibuchi Falls waterfalls man-made dam spring Ichigome Falls Gogome Falls Fig. 1-3. Map and landscapes of the Ogata River and its tributaries, the Kohbaru and the Hakiai streams, northeastern Kyushu Island, Japan. 7 ➨ 2 ❶ ὶ⛬ἢࡗࡓ㨶㢮⩌㞟ࡢኚ Ἑᕝୖὶᇦࢆྵࡴὶ⛬ἢࡗࡓ㨶㢮⩌㞟ࡢኚࡘ࠸࡚ࡣከࡃࡢ◊✲ࡀ࠶ࡾ 㸦࠼ࡤ Rahel and Hubert, 1991㸹Aarts and Nienhuis, 2003㸹Torgersen et al., 2006㸹 Lasne et al., 2007㸹Mercado-Silva et al., 2012㸧㸪࠸ࡎࢀࡶୖὶഃࡢࢧࢣ⛉㨶㢮ࢆ୰ ᚰࡍࡿ⩌㞟ࡽୗὶഃࡢࢥ⛉㨶㢮➼ࢆ୰ᚰࡍࡿ⩌㞟ࡢ⛣⾜㸦transition㸧 ࢆሗ࿌ࡋ࡚࠸ࡿ㸬㨶㢮⩌㞟ࡢ⛣⾜ࡣ㸪Ỉ ࡸᆅᙧ࠸ࡗࡓẚ㍑ⓗࡁ࡞✵㛫ࢫ ࢣ࣮ࣝ࠾ࡅࡿ≀⌮⎔ቃࡢኚᑐᛂࡋ࡚⏕ࡌࡿ㸦Naiman et al., 1988㸹Rahel and Hubert, 1991㸹Jackson et al., 2001㸹Torgersen et al., 2006㸧 㸬ࡑࡢࡓࡵ㨶㢮⩌㞟ࡢ⛣ ⾜ࡣ㸪⎔ቃ໙㓄ࡢࡁ࡞ᒣᆅἙᕝ࠾ࡅࡿ୍⯡ⓗ࡞⩌㞟ኚࡢᵝᘧࡋ࡚▱ࡽ ࢀ࡚࠸ࡿ㸦Rahel and Hubert, 1991㸹Torgersen et al., 2006㸧 㸬᪥ᮏࡢἙᕝࡣ㝣ࡢ Ἑᕝẚ࡚ᛴ໙㓄࡛࠶ࡿࡇࡽ㸦Ỉ㔝࣭ᚚໃ㸪2000㸧㸪⩌㞟ࡢ⛣⾜ࡣᅜෆ ࠾࠸࡚ࡶ୍⯡ⓗ࡞⩌㞟ኚࡢᵝᘧ࡛࠶ࡿ⪃࠼ࡽࢀࡿ㸬୍᪉㸪Ἑᕝ⎔ቃࡢேⅭ ᨵኚࡼࡗ࡚ὶἣࡀኚࡍࡿ㸪⩌㞟ᵓᡂ✀ࡢὶ⛬ศᕸࡀኚࡋ㸪㨶㢮⩌㞟ࡢ ⛣⾜ᵝᘧࡀኻࢃࢀࡿࡇࡀሗ࿌ࡉࢀ࡚࠸ࡿ㸦Aarts and Nienhuis, 2003㸹Lasne et al., 2007㸧㸬୍⯡ⓗ㸪Ἑᕝࡢୖὶᇦୗὶᇦ࡛ࡣ✀㛫㛵ಀࡸ⩌㞟ࡢᙧᡂ࣭⥔ᣢᶵᵓ ࡀ␗࡞ࡗ࡚࠾ࡾ㸦Fagan et al., 1999㸹ୖ㸪2013㸧 㸪≉ࢧࢣ⛉㨶㢮ࢆ୰ᚰࡍ ࡿ⩌㞟ࡽࢥ⛉㨶㢮➼ࢆ୰ᚰࡍࡿ⩌㞟ࡢ⛣⾜ࡀ⏕ࡌࡿỈᇦ㸦⛣⾜ᖏ㸸 transition zone㸪Rahel and Hubert, 1991㸧࡛ࡣ㸪ᵓᡂ✀ࡀྛࠎࡢศᕸࡢ࿘⦕㒊 ⏕ᜥࡋ࡚࠸ࡿࡓࡵ㸪⎔ቃᨵኚᑐࡍࡿឤཷᛶࡀ㧗࠸ᣦࡉࢀ࡚࠸ࡿ㸦Naiman et al., 1988㸹Aarts and Nienhuis, 2003㸹Lasne et al., 2007㸧 㸬ࡋࡓࡀࡗ࡚㸪ὶ⛬ἢ ࡗࡓ㨶㢮⩌㞟ࡢኚࢆㄪグ㍕䛩䜛䛣䛸䛿㸪ᚋࡢேⅭᨵኚࡢᙳ㡪ࢆホ౯ࡍࡿࡓ ࡵࡶ㸦ᾷ㸪2001㸧㸪ᆅᇦࡢ⏕ែ⣔ಖࡸ≉ᐃ✀ࡢᣢ⥆ⓗ⏝㸦∦㔝㸪 2004㸧ࢆᅗࡿࡓࡵࡶᙺ❧ࡘ⪃࠼ࡽࢀࡿ㸬 ᮏ◊✲ࡣ㸪࣐ࢦࡀ⮬↛ศᕸ㸦ᓥ㸪1957㸧ࡍࡿᕞᮾ㒊ࡢἙᕝୖὶᇦ ࠾࠸࡚㸪ὶ⛬ἢࡗࡓ㨶㢮⩌㞟ࡢኚᵝᘧࢆ᫂ࡽࡍࡿࡇࢆ┠ⓗᐇࡋ ࡓ㸬ࡑࡢࡓࡵ㸪₯Ỉ┠どほᐹࡼࡿ㨶㢮⩌㞟ࢹ࣮ࢱࢆ⏝࠸࡚ከኚ㔞ゎᯒࡼ ࡿ⩌㞟ࡢศ㢮࣭ᗎิࢆ⾜࠺ࡶ㸪ὶ⛬ἢࡗࡓ㨶㢮⩌㞟ࡢኚᙳ㡪ࢆ 8 ࠼ࡿ⎔ቃせᅉࢆேⅭᨵኚࡢᙳ㡪ࡶ᥎ᐃ࣭⪃ᐹࡋࡓ㸬 ᮦᩱ᪉ἲ ㄪᰝሙᡤㄪᰝⅬ ᮏ◊✲࡛ࡣ㸪㔝ᕝỈ⣔⥴᪉ᕝ༑ゅᕝࡢྜὶⅬࡽ㸪⚄ཎᕝࡢ㨶㢮⏕ᜥୖ 㝈࡛࠶ࡿྜ┠ࡢ㸦ᮌᮧ㸪1959㸧ࡲ࡛ࡢ⣙ 16.4 km ࡢ⠊ᅖࢆㄪᰝỈᇦࡋ㸪 ୗὶࡽୖὶྥࡅ࡚ 15 ࡢㄪᰝⅬ㸦St. 1–15㸧ࢆタࡅࡓ㸦Fig. 2-1㸧㸬ࡇࡢ࠺ࡕ St. 11 ࡣ㸪⚄ཎᕝࡢྜὶⅬࡽ⣙ 820 m ୖὶࡢἼᮌྜᕝ㸦ሖሐࡢ┤ୗ㸧ࡢㄪ ᰝⅬࡋࡓ㸬ྛㄪᰝⅬ㛫ࡢ㊥㞳ࡣ 0.7–2.1 km㸦ᖹᆒ್ ± ᶆ‽೫ᕪ = 1.2 ± 0.4 km㸧 ࡛㸪ྛㄪᰝⅬࡢὶ㊰㛗ࡣ℩࣭ῡᵓ㐀ࢆᑡ࡞ࡃࡶ୍ࡘࡎࡘྵࡴ 42–146 m㸦94 ± 37 m㸧࡛࠶ࡿ㸬ᮏㄪᰝỈᇦࡢୗὶഃ࠾ࡼࡧỈᇦෆ࠶ࡿࢲ࣒㸦ⴠᕪ 1 m ௨ ୖࡢࡳᢤ⢋㸧ࡢ⨨≉ᚩࢆ Fig. 2-1 Table 2-1 ♧ࡋࡓ㸬ᮏㄪᰝỈᇦࡢୗὶ 23㸪19 ࠾ࡼࡧ 9 km ᆅⅬࡣࡑࢀࡒࢀࢲ࣒ A B ࠾ࡼࡧ C ࡀ࠶ࡾ㸪㏻ࡋᅇ㐟 㨶ࡣ㐍ධ࡛ࡁ࡞࠸ࡀ㸪㔝ᕝ⁺ᴗ༠ྠ⤌ྜࡼࡾࣘ Plecoglossus altivelis altivelis ࣐ࢦࡢ⛶㨶ࡀࡑࢀࡒࢀ St. 1–4 St. 1–10 ࢆ୰ᚰẖᖺᨺὶࡉࢀ࡚ ࠸ࡿ㸬ࡲࡓ㸪⫋ᴗⓗ࡞⁺ᴗࡣ⾜ࢃࢀ࡚࠸࡞࠸ࡀ㸪⁺༠⤌ྜဨࡼࡿࣘ⁺ࡢ 㸪㐟⁺ࢆྵࡴ࣐ࢦ㔮ࡾࡀᨺὶᇦࢆ୰ᚰ⾜ࢃࢀ࡚࠸ࡿ㸦㔝ᕝ⁺༠㸪 ⚾ಙ㸧 㸬 ྛㄪᰝⅬࡢ≉ᚩࢆグࡍ㸪St. 1–4 ࡢἙᕝᙧែᆺ㸦ྍඣ㸪1978㸧ࡣ Bb ᆺ࡛㸪࿘ ᅖࡣ㛤ࡅ࡚Ỉ⏣⏝ࡉࢀ࡚࠸ࡓ㸬㏆ 5 ࡘࡢỈ⩌㸦Ỉ ⣙ 14–17ºC㸧ࡀ ࠶ࡾ㸦Fig. 1㸧㸪⣙ 0.35–0.46 m3/s ࡀ⥴᪉ᕝὶධࡋ࡚࠸ࡿ㸦ᓥ㔝࣭ᡞḟ㸪1994㸧㸬 St. 5–10 ࡣ Aa-Bb ⛣⾜ᆺࢆ♧ࡋ㸪St. 6–9 ࡛ࡣ୧ᓊᩳ㠃ࡀ㏕ࡗ࡚㇂ࡢᬒほࢆ♧ ࡋࡓ㸬St. 11–15 ࡣ Aa ᆺ࡛㸪St. 13 ࡽୖὶ࡛Ἑᗋ໙㓄ࡀࡁ࠸㸬ἙᕝࡢேⅭᨵ ኚࡣ࠾ࡶ St. 6–13 ࡛ぢࡽࢀࡓ㸬ࡇࡢ࠺ࡕ St. 6–10 ࡢᨵኚࡣᕥᓊ㐨㊰ἢ࠸ࡢㆤ ᓊ࡛࠶ࡾ㸪ὶ㊰ෆᕤࡀ࠾ࡼࡪሙᡤࡣ㝈ࡽࢀࡓࡀ㸪St. 10 ┤ୖࡢሖሐ H–J ࡢ ⚄ཎᕝࡢ༊㛫࡛ࡣ㸪ὶ㊰ෆ㐩ࡍࡿㆤᓊࡀ୧ᓊࡳࡽࢀࡓ㸬≉ St. 12 ࡽ J ࡲ࡛ࡢ༊㛫ࡣᑠつᶍ࡞ሖሐ⩌ I ࡀタ⨨ࡉࢀ㸦Fig. 2-1㸧㸪ᮏㄪᰝỈᇦࡢ୰࡛᭱ࡶ ேⅭᨵኚࡢࡁ࠸ሙᡤ࡛࠶ࡗࡓ㸬୍᪉㸪Ἴᮌྜᕝࡢ St. 10–11 ࡛ࡣᶆ㧗㸪Ἑᗋ໙ 9 㓄㸪Ỉ ➼ࡢ⎔ቃኚ㔞ࡀ⚄ཎᕝࡢ St. 10–12 ࡰྠࡌ࡛࠶ࡗࡓࡀ㸦ᚋ㏙㸹Table 2-2㸧㸪ᨵኚࡣྑᓊ㐨㊰ἢ࠸ࡢ୍㒊ࡢㆤᓊᕤ㝈ࡽࢀ㸪㨶㢮ࡢ⛣ືࢆ㜼ᐖࡍࡿ ᵓ㐀ࡣ࡞ࡗࡓ㸦Fig. 2-1㸧㸬 ⎔ቃㄪᰝ ⩌㞟ኚ⎔ቃせᅉࡢ㛵ಀࢆㄪࡿࡓࡵ㸪௨ୗࡢ 6 ࡘࡢ⎔ቃኚ㔞ࢆồࡵࡓ 㸦Table 2-2㸧 㸬ྛㄪᰝⅬࡢᶆ㧗ࢆ⦰ᑻ 1/5000 ࡢᆅᅗࡽㄞࡳྲྀࡿࡶ㸪㞄 ᥋ㄪᰝⅬࡢᶆ㧗ᕪࢆㄪᰝⅬ㛫ࡢ㊥㞳࡛㝖ࡋ࡚㸪 ୖὶഃㄪᰝⅬࡢἙᗋ໙㓄ࡋࡓ㸬 ࡲࡓ㸪2004 ᖺ 3 ᭶ㄪᰝỈᇦࡢὶ㊰㛗⣙ 100 m ẖỈ㠃ᖜࢆ ᐃࡋ㸪㞄᥋ㄪ ᰝⅬ㛫ࡢᖹᆒ್ࢆୖὶഃㄪᰝⅬࡢᖹᆒỈ㠃ᖜ㸦௨ୗ㸪Ỉ㠃ᖜ㸧ࡋࡓ㸬St. 1 ࡛ ࡣ୧ኚ㔞ࡀᚓࡽࢀ࡞࠸ࡓࡵ㸪௨ୗࡢゎᯒ࡛ࡣ St. 2 ྠࡌ್ࢆ⏝࠸ࡓ㸬ࡲࡓ㸪ྛ ㄪᰝⅬ࡛ὶࢀࡢᏳᐃࡋࡓሙᡤ ᐃ᩿㠃ࢆタࡅ㸪ᦠᖏࣉࣟ࣌ࣛᘧὶ㏿ィ㸦3631 ᆺ㸪ᶓἙ㟁ᶵ㸧ࢆ⏝࠸࡚ 1 Ⅼἲ㸦୰ᮧ㸪1997㸧ࡼࡾὶ㔞ࢆ ᐃࡋࡓ㸬2003 ᖺ 8 ᭶–2004 ᖺ 10 ᭶ࡣ㸪ྛㄪᰝⅬࡢᒾࡸᶫ⬮Ỉ ࢹ࣮ࢱ࣮ࣟ࢞㸦StowAway TidbiT Temp Logger, Onset Computer ♫㸧ࢆタ⨨ࡋ࡚ 1 㛫ẖࡢỈ ࢆグ㘓ࡋ㸪 Ỉ グ㘓ࡽ᪥ᖹᆒỈ ࢆồࡵ࡚㸪ᮇ㛫ෆࡢ᪥ᖹᆒ᭱㧗Ỉ 㸪ྠ᭱పỈ 㸦௨ ୗ㸪᭱㧗࣭᭱పỈ 㸧ࢆᢳฟࡋࡓ㸬 ₯Ỉ┠どほᐹ Ἑᕝࡢ㨶㢮⩌㞟⤌ᡂࡣ㸪㨶㢮ࡢᏘ⠇ⓗ࡞⛣ືࡸᕼᑡ✀ࡢⓎぢ⋡ࡀప࠸ࡇ➼ ࡼࡾኚືࡍࡿྍ⬟ᛶࡀᣦࡉࢀ࡚࠸ࡿ㸦ᾷ㸪2001㸧 㸬ᮏ◊✲࡛ࡣṇ☜࡞⩌ 㞟⤌ᡂࢆᢕᥱࡍࡿࡓࡵ㸪⛅ᮇ㸦2003 ᖺ 10 ᭶ 21–29 ᪥㸧㸪ᮇ㸦2004 ᖺ 2 ᭶ 2–10 ᪥㸧㸪ኟᮇ㸦2004 ᖺ 8 ᭶ 6–24 ᪥㸪ࡓࡔࡋ㝆㞵ࡼࡿ⃮ࡾࡢࡓࡵ St. 15 ࡢ ࡳ 10 ᭶ 6 ᪥㸧㸪ྛㄪᰝⅬ⏕ᜥࡍࡿࡍ࡚ࡢ㨶✀ࢆᑐ㇟₯Ỉ┠どほᐹࢆ ⾜ࡗࡓ㸬₯Ỉ┠どほᐹࢆ⏝࠸ࡓ⌮⏤ࡣ㸪ከࡃࡢㄪᰝⅬ㌟㛗ࢆ㉸࠼ࡿỈ῝ࡢῡ ࡀ࠶ࡾ㸪ࡢ᪉ἲ࡛ࡣㄪᰝࡀᅔ㞴࡛࠶ࡗࡓࡓࡵ࡛࠶ࡿ㸦Thurow, 1994㸹Torgersen et al., 2006㸧 㸬Ἑᕝᕤ➼ࡢᩓⓎⓗ࡞⃮ࡾࢆ㝖ࡃᮏㄪᰝỈᇦࡣΎ࡛࠶ࡾ㸪₯ Ỉ┠どほᐹ㐺ࡋࡓỈ㉁ࢆ᭷ࡋ࡚࠸ࡓ 㸦ᮌᮏ㸪2008㸧㸬 ほᐹ᪉ἲࡣᮌᮏ 㸦2013㸧 10 ࡢ࠾ࡾ࡛࠶ࡾ㸪☜ㄆࡉࢀࡓ㨶✀ูಶయᩘࢆ 5 cm ᖜࡢ㛗㝵⣭ู㔝ᖒグ㘓 ࡋࡓ㸬࡞࠾ 㸪࣐࢝ ࢶ ࢝ Pseudogobio esocinus esocinus ࡸ ࣚ ࢩ ࣀ ࣎ࣜᒓ㨶㢮 Rhinogobius sp.➼ࡢᗏ⏕ᛶ㨶㢮ࡣἙᗋࡢ◁ࡸ♟୰㞃ࢀࡿᛶ㉁ࡀ࠶ࡿࡀ㸪Ἑᗋࢆ ᥀ࡾ㏉ࡋࡓሙྜࡣ✀ࡢⓎぢ⋡ࡀపୗࡍࡿ⪃࠼ࡽࢀࡓࡓࡵ㸪㞃ࢀ࡚࠸࡞࠸ ಶయࡔࡅࢆィᩘࡋࡓ㸬ࣚࢩࣀ࣎ࣜᒓ㨶㢮㸪ࢩ࣐ࢻࢪࣙ࢘ᒓ㨶㢮 Cobitis sp.㸪ࣇ ࢼᒓ㨶㢮 Carassius sp.ࡣ㸪₯Ỉ┠どほᐹࡼࡿṇ☜࡞✀ྠᐃࡀᅔ㞴࡞ࡓࡵᒓࡲ ࡛ࡢศ㢮ࡋ㸪ࡑࢀࡒࢀ 1 ✀ࡋ࡚ᢅࡗࡓ㸬ࡲࡓ㸪࣐ࢦ࣐࣓ࣖࡢᾋୖ⛶㨶 ࡣ㸪୧✀ࢆእ㒊ᙧែࡽ༊ู࡛ࡁ࡞ࡗࡓࡓࡵ㸪௨ୗࡢゎᯒࡽ㝖እࡋࡓ㸬ᙧ ែࡀ㢮ఝࡍࡿࢱ࢝ࣁࣖ Phoxinus oxycephalus㸪࢘ࢢ Tribolodon hakonensis㸪࢝ ࣒࣡ࢶ Candidia temminckii㸪࢜࢝࣡ Opsariichthys platypus ࡢ⛶㨶ࡘ࠸࡚ࡶ㸪 ṇ☜ࢆᮇࡍࡓࡵ㸪㛗 5 cm ௨ୗࡢࢹ࣮ࢱࢆゎᯒࡽ㝖࠸ࡓ㸬ᚓࡽࢀࡓಶయᩘ ࢹ࣮ࢱࡣ㸪ྛㄪᰝⅬࡢỈ⾲㠃✚࡛㝖ࡋ࡚ 100 m2 ࠶ࡓࡾࡢ㨶✀ูಶయ⩌ᐦᗘኚ ࡋ࡚ẚ㍑࣭ศᯒ౪ࡋࡓ㸬Ꮫྡᶆ‽ྡࡣ୰ᆓ㸦2013㸧ᚑࡗࡓ㸬 ⩌㞟ゎᯒ ὶ⛬ἢࡗࡓ㨶㢮⩌㞟ᵓ㐀ࢆᢕᥱࡍࡿࡓࡵ㸪ྛㄪᰝⅬࡢ㨶㢮⩌㞟ࡘ࠸࡚㸪 ㄪᰝᮇูࢡࣛࢫࢱ࣮ศᯒ㠀ィ㔞ከḟඖᑻᗘᵓᡂἲ㸦NMDS㸧ࡼࡿᗎิ ࢆ⾜ࡗࡓ㸬㨶✀ูಶయ⩌ᐦᗘᑐࡋ࡚ Oksanen et al.㸦2013㸧ࡀ᥎ዡࡍࡿᖹ᪉ ᰿ኚ Wisconsin double standardization ࢆࡋࡓࡢࡕ㸪ㄪᰝⅬ㛫࡛ Bray-Curtis 㠀㢮ఝᗘ⾜ิࢆ⟬ฟࡋ࡚୧ศᯒ౪ࡋࡓ㸬ࢡࣛࢫࢱ࣮ศᯒ࡛ࡣ㸪⤫ィࢯࣇࢺ R 㸦R Development Core Team, 2005㸧ࡢ㛵ᩘ hclust ࢆ⏝࠸࡚㸪㠀㢮ఝᗘ⾜ิࡽ⩌ ᖹᆒἲࡼࡾඹ⾲ᙧ㊥㞳⾜ิ㸦cophenetic distance matrix㸧ࢆ⟬ฟࡋ࡚ᶞ≧ᅗࢆ సᡂࡋࡓ㸬 NMDS ࡼࡿᗎิࡣ R ࡢ vegan ࣃࢵࢣ࣮ࢪ㸦Oksanen et al., 2013㸧 ࡽ㛵ᩘ metaMDS ࢆ⏝࠸࡚ྛㄪᰝⅬࢆḟඖᖹ㠃ୖ㓄⨨ࡋࡓ㸬NMDS ࡣ㸪 ྛㄪᰝⅬࢆ㠀㢮ఝᗘ⾜ิෆࡢ㡰ᇶ࡙࠸࡚㠀⥺ᙧⓗᖹ㠃ୖ㓄⨨ࡍࡿࡓࡵ㸪 ⥺ᙧᛶࡢ௬ᐃࡀᅔ㞴࡞⏕ែᏛࢹ࣮ࢱࡢᗎิ㐺ࡋ࡚࠸ࡿ㸦Oksanen, 2013㸧㸬 㠀㢮ఝᗘ⾜ิᑐࡍࡿ୧ศᯒࡢ㐺ྜᗘࡢᣦᶆࡋ࡚ඹ⾲ᙧ┦㛵ಀᩘࢫࢺࣞࢫ ್ࢆồࡵࡓ㸬๓⪅ࡣ㠀㢮ఝᗘ⾜ิඹ⾲ᙧ㊥㞳⾜ิࡢ┦㛵ࢆ⾲ࡋ㸪0–1 ࡢ⠊ᅖ 11 ࡛ࡁ࠸㠀㢮ఝᗘᑐࡍࡿᶞ≧ᅗࡢ㐺ྜᗘࡀ㧗࠸㸦Oksanen, 2014㸧㸬ᚋ⪅ ࡣ㠀㢮ఝᗘࡢ㡰 ᗎิࡉࢀࡓⅬ㛫㊥㞳ࡢ㡰ࡢ┦㛵ࢆ⾲ࡋ㸦Oksanen, 2013㸧㸪0–1 ࡢ⠊ᅖ࡛ᑠࡉ࠸㠀㢮ఝᗘᑐࡍࡿ NMDS ࡢ㐺ྜᗘࡀ㧗࠸㸦㔠㸪 2007㸧㸬ࡲࡓ㸪⩌㞟ᵓ㐀ྛ㨶✀ࡢ㛵ಀࢆㄪࡿࡓࡵ㸪㛵ᩘ wascore ࢆ⏝࠸࡚㸪 ᗎิࡉࢀࡓྛㄪᰝⅬࡢᗙᶆᑐࡍࡿྛ㨶✀ࡢ㔜ᚰࢆ⟬ฟࡋ NMDS ᖹ㠃ୖ 㓄⨨ࡋࡓ㸬ࡉࡽ㸪㛵ᩘ envfit ࢆ⏝࠸࡚㸪ྛㄪᰝⅬୖグ 6 ࡘࡢ⎔ቃኚ㔞࠾ࡼ ࡧἙᕝᙧែᆺࡢ㛵ಀࢆㄪࡓ㸬㔞ⓗኚᩘ࡛࠶ࡿ 6 ࡘࡢ⎔ቃኚ㔞ࡣ㸪ᗎิࡉ ࢀࡓྛㄪᰝⅬࡢᗙᶆࡢ⥺ᙧ⤖ྜࡋ࡚⾲ࡋ㸪NMDS ᖹ㠃ୖ࣋ࢡࢺࣝࡋ࡚ᑕ ᙳࡋࡓ㸬࣋ࢡࢺࣝࡢ᪉ྥࡣྛ⎔ቃኚ㔞ࡢቑຍ㏿ᗘࡀ᭱ࡶࡁ࡞᪉ྥࢆ♧ࡋ㸪㛗 ࡉࡣྛㄪᰝⅬࡢᗙᶆࡢ┦㛵ࡢ㧗ࡉࢆ⾲ࡍ㸬୍᪉㸪㉁ⓗኚᩘ࡛࠶ࡿἙᕝᙧែᆺ ࡣ㸪ྛㄪᰝⅬࡢᗙᶆᑐࡍࡿ㔜ᚰࡋ࡚ NMDS ᖹ㠃ୖ♧ࡋࡓ㸦Oksanen, 2013㸧 㸬 ᗎิࡉࢀࡓྛㄪᰝⅬᑐࡍࡿྛ⎔ቃኚ㔞ࡢ㐺ྜᗘࡘ࠸࡚㸪⎔ቃኚ㔞ࡢ୪ ᭰࠼᳨ᐃ㸦10,000 ᅇ㸧ࢆᐇࡋ㸪᭷ពᛶࢆุᐃࡋࡓ㸦Oksanen, 2013㸧㸬ୖグࡢ 㸪ࡍ࡚ࡢ⤫ィゎᯒࡣ⤫ィࢯࣇࢺ R㸪࠾ࡼࡧࡇࢀࢆ࣮࣋ࢫࡋࡓࣇ࣮ࣜ⤫ ィࢯࣇࢺ EZR㸦⚄⏣㸪2012㸧࡛ᐇࡋ㸪⤫ィ᳨ᐃࡢ᭷ពỈ‽ࡣ 0.05 ࡋࡓ㸬 ⤖ ᯝ ㄪᰝỈᇦࡢ⎔ቃ ᐃࡋࡓ 6 ࡘࡢ⎔ቃኚ㔞ࡢ࠺ࡕ㸪5 ኚ㔞ࡀὶ⛬ἢࡗࡓ໙㓄ࢆ♧ࡋࡓ㸬ࡍ࡞ ࢃࡕ㸪ᶆ㧗Ἑᗋ໙㓄ࡣୖὶ᪉ྥ㸪Ỉ㠃ᖜ㸪ὶ㔞㸪᭱పỈ ࡣୗὶ᪉ྥ᭷ព ቑຍࡋࡓ㸦Spearman ࡢ㡰┦㛵ಀᩘ㸪ᑐㄪᰝⅬ␒ྕ㸪㡰 rs = 1.0㸪0.93㸪-0.84㸪 -0.98㸪-0.90㸪ࡍ࡚ P < 0.001㸹Table 2-2㸧㸬୍᪉㸪᭱㧗Ỉ ࡣỈࡢὶධࡼ ࡾ St. 1–4 ࡛పୗࡋ㸪ㄪᰝⅬ␒ྕ᭷ព࡞┦㛵ࢆ♧ࡉ࡞ࡗࡓ㸦rs = -0.36㸪P = 0.18㸧㸬௨ୖࡢࡇࡽ㸪ᶆ㧗㸪Ἑᗋ໙㓄ࡣୖὶ᪉ྥ㸪Ỉ㠃ᖜ㸪ὶ㔞㸪᭱పỈ ࡣୗὶ᪉ྥ༢ㄪቑຍࡍࡿ⎔ቃኚ㔞࡛࠶ࡾ㸪᭱㧗Ỉ ࡣὶ⛬ἢࡗࡓኚࢆ♧ ࡉ࡞࠸ኚ㔞࠸࠼ࡿ㸬 12 㨶✀ูὶ⛬ศᕸ ࡍ࡚ࡢㄪᰝ࡛㸪࢘ࢼࢠ┠࢘ࢼࢠ⛉ 1 ✀㸪ࢥ┠ࢥ⛉ 7 ✀㸪ࢻࢪࣙ࢘⛉ 1 ✀㸪ࢧࢣ┠ࣘ⛉ 1 ✀㸪ࢧࢣ⛉ 2 ள✀㸪ࢫࢬ࢟┠ࢻࣥࢥ⛉ 1 ✀㸪ࣁࢮ⛉ 1 ✀ࡢ 4 ┠ 7 ⛉ 14 ✀㸦ள✀ࢆྵࡴ㸧ࡀ☜ㄆࡉࢀ㸪⥲☜ㄆಶయᩘࡣ 18,015 ಶయ࡛࠶ࡗ ࡓ㸦Tables 2-3-1, 2-3-2, 2-3-3㸧 㸬☜ㄆಶయᩘࡣࢱ࢝ࣁࣖ㸦9,368 ಶయ㸪52.0%㸧㸪࢝ ࣒࣡ࢶ㸦3,010 ಶయ㸪16.7%㸧㸪࢘ࢢ㸦2,274 ಶయ㸪12.6%㸧㸪ࣚࢩࣀ࣎ࣜ㢮㸦1,677 ಶయ㸪9.3%㸧㸪࣐ࢦ㸦633 ಶయ㸪3.5%㸧ࡢ㡰ከࡗࡓ㸬㨶✀ูὶ⛬ศᕸࡢ㐪 ࠸ࡼࡾ㸪ྛㄪᰝⅬࡢ㨶㢮⩌㞟ࡣ㸪ୖὶഃࡢࢧࢣ⛉㨶㢮ࢆ୰ᚰࡋࡓ⩌㞟ࡽ ୗὶഃࡢࢥ⛉㨶㢮୰ᚰࡢ⩌㞟⛣⾜ࡋࡓ㸦Tables 2-3-1, 2-3-2, 2-3-3㸧 㸬ࡍ࡞ ࢃࡕ㸪࣐ࢦࡣ St. 2–15 ࡛☜ㄆࡉࢀ㸪ୖὶഃ࡛ࡣࢱ࢝ࣁࣖඹᏑࡋࡓࡀ㸪ୗὶ ഃ࡛ࡣࡼࡾከࡃࡢࢥ⛉㨶㢮➼ඹᏑࡋ࡚࠸ࡓ㸬࣐ࢦḟ࠸࡛ୖὶ࡛☜ㄆࡉ ࢀࡓ㨶✀ࡣࢱ࢝ࣁ࡛ࣖ࠶ࡾ St. 1–14 ࡛ぢࡽࢀࡓ㸬௨ୗ࢘ࢢࡀ St. 1–13㸪࣒࢝࣡ ࢶࡀ St. 1–12㸪ࢻࣥࢥ Odontobutis obscura ࡀ St. 1–9㸪࣐࢝ࢶ࢝ࡀ St. 1–8㸪ࣚࢩ ࣀ࣎ࣜᒓ㨶㢮ࡀ St. 1–4㸪࢜࢝࣡ࢩ࣐ࢻࢪࣙ࢘ᒓ㨶㢮ࡀ St. 1–3 ࡛☜ㄆࡉࢀࡓ㸬 ኟᮇࡣᨺὶࡉࢀࡓࣘࡀ St. 1–4 ࡛ぢࡽࢀࡓ㸬࣐࣓ࣖ㸪ࢥ Cyprinus carpio㸪 ࣇࢼᒓ㨶㢮㸪ࢽ࣍ࣥ࢘ࢼࢠ Anguilla japonica ࡣ㞳ᩓࡋࡓㄪᰝⅬ࡛ 1–9 ಶయࡀぢ ࡽࢀࡓ㸬 ✀ᩘಶయ⩌ᐦᗘࡢὶ⛬ኚ ྛㄪᰝⅬࡢ☜ㄆ✀ᩘ⥲ಶయ⩌ᐦᗘࡣ⛅ᮇኟᮇୗὶ᪉ྥ༢ㄪቑຍࡋ ࡓ㸦Spearman ࡢ㡰┦㛵ಀᩘ㸪✀ᩘ㸸rs = 0.86㸪0.95㸪ࡶ P < 0.001㸹ಶయ ⩌ᐦᗘ㸸rs = 0.61㸪0.60㸪ࡶ P < 0.05㸧㸬୍᪉㸪⏕ᜥ⠊ᅖࡀᗈࡃ⏕ᜥ㔞ࡀከ ࡗࡓ࣐ࢦ㸪ࢱ࢝ࣁࣖ㸪࣒࢝࣡ࢶ㸪࢘ࢢ㸪࠾ࡼࡧࡑࡢࡢ㨶✀ูಶయ⩌ ᐦᗘࢆࡳࡿ㸦Fig. 2-2㸧㸪ࡍ࡚ࡢᏘ⠇࡛࣐ࢦࡀୗὶ᪉ྥ༢ㄪῶᑡ㸦rs = -0.89–-0.94㸪ࡍ࡚ P < 0.001㸧㸪࣒࢝࣡ࢶࡀ༢ㄪቑຍ㸦rs = 0.56–0.60㸪ࡍ࡚ P < 0.05㸧 㸪ࡑࡢࡢ㨶✀ࡀ༢ㄪቑຍࡋ㸦⛅ᮇኟᮇ㸸rs = 0.89–0.92㸪ࡶ P < 0.001㸹ᮇ㸸rs = 0.60㸪P < 0.05㸧㸪࢘ࢢࡣ⛅ᮇኟᮇ༢ㄪቑຍࡋࡓࡀ㸦⛅ ᮇ㸸rs = 0.63㸪P < 0.05㸹ኟᮇ㸸rs = 0.69㸪P < 0.001㸧㸪ࢱ࢝ࣁࣖࡢಶయ⩌ᐦᗘࡣ 13 ࡍ࡚ࡢᏘ⠇࡛ὶ⛬ἢࡗࡓኚࢆ♧ࡉ࡞ࡗࡓ㸦rs = 0.16–0.50㸪P = 0.06– 0.51㸧㸬ࢱ࢝ࣁࣖࡢಶయ⩌ᐦᗘࡣᓠᆺࢆ♧ࡋ㸦Fig. 2-2㸧㸪⛅ᮇᮇ St. 6–11 ࡛పࡃ㸪ኟᮇ St. 7–10 ࡛పࡗࡓ㸬ࡲࡓ㸪St. 4–5 ࡛ࡣ࢘ࢢ࣒࢝࣡ࢶࡢಶ య⩌ᐦᗘࡀᒁᡤⓗపࡃ㸪ࢱ࢝ࣁࣖࡀඃ༨ࡋ࡚࠸ࡓ㸦Fig. 2-2㸧㸬 ⩌㞟ࡢศ㢮ᗎิ ᮏゎᯒ࡛ࡣ㸪ᩓⓎⓗ☜ㄆࡉࢀࡓ࣐࣓ࣖ㸪ࢥ㸪ࣇࢼᒓ㨶㢮㸪ࢽ࣍ࣥ࢘ࢼࢠ㸪 ࠾ࡼࡧ᫂ࡽ࡞ᨺὶ㨶࡛࠶ࡿࣘࡢࢹ࣮ࢱࢆ㝖࠸ࡓ㸬ࢡࣛࢫࢱ࣮ศᯒ࡛ࡣ㸪㨶 ✀ูὶ⛬ศᕸಶయ⩌ᐦᗘࢆᫎࡋࡓ⤖ᯝࡀᚓࡽࢀࡓ㸬ࡍ࡞ࢃࡕ㸪⛅ᮇኟᮇ ඹ⾲ᙧ㊥㞳 0.5㸪ᮇ 0.7 ࡛ᶞ≧ᅗࢆษ᩿ࡍࡿ㸪ྛㄪᰝⅬࡢ㨶㢮⩌㞟ࡣ ࡰ St. 1–3㸪6–10㸪11–15 ࡢ 3 ⩌ศ㢮ࡉࢀ㸪St. 4–5 ࡣㄪᰝᮇࡼࡾ␗࡞ࡿࢡ ࣛࢫࢱ࣮ᒓࡋࡓ㸦Fig. 2-3㸧 㸬ඹ⾲ᙧ┦㛵ಀᩘࡣ⛅ᮇ㸪ᮇ㸪ኟᮇࡢ㡰 0.87㸪 0.76㸪0.82 1 ㏆࠸್ࢆ♧ࡋࡓ㸬NMDS ࡛ࡶࢡࣛࢫࢱ࣮ศᯒྠᵝ㸪St. 1– 3㸪4–5㸪6–10㸪11–15 ࡀࡑࢀࡒࢀ㏆࠸⨨㓄⨨ࡉࢀࡓ㸦Fig. 2-3㸧㸬ࢫࢺࣞࢫ್ ࡣ⛅ᮇ㸪ᮇ㸪ኟᮇࡢ㡰 0.09㸪0.12㸪0.09 㸪㧗࠸㐺ྜᗘࢆ♧ࡋࡓ㸬NMDS ᖹ㠃ୖࡢྛ㨶✀ࡢ㔜ᚰࡣ㸪㨶✀ูὶ⛬ศᕸಶయ⩌ᐦᗘࢆᫎࡋ㸪ࡍ࡚ࡢᏘ ⠇࡛࣐ࢦࡀ St. 11–15㸪ࢱ࢝ࣁࣖࡀ St. 11–15 4–5 ࡢ㛫㸪࢘ࢢࡀ St. 6–10㸪 ࣒࢝࣡ࢶࡀ St. 1–3 St. 6–10 ࡢ㛫㸪ࣚࢩࣀ࣎ࣜ㢮㸪࢜࢝࣡㸪ࢩ࣐ࢻࢪࣙ࢘ᒓ 㨶㢮ࡀ St. 1–3 ㏆㓄⨨ࡉࢀࡓ㸬࣐࢝ࢶ࢝ࢻࣥࢥࡢ㔜ᚰࡣ㸪⛅ᮇኟᮇ St. 6–10㸪ᮇ St. 1–3 ㏆⨨ࡋࡓ㸬௨ୖࢆࡲࡵࡿ㸪ᮏㄪᰝỈᇦࡢ㨶 㢮⩌㞟ࡣ࣐ࢦࢱ࢝ࣁࣖࢆ୰ᚰࡋࡓ⩌㞟㸦St. 11–15㸧㸪࢘ࢢ࣒࢝࣡ࢶࢆ ୰ᚰࡋࡓ⩌㞟㸦St. 6–10㸧 㸪ࢱ࢝ࣁࣖࢆ୰ᚰࡋࡓ⩌㞟㸦St. 4–5㸧㸪ከᩘࡢࢥ ⛉㨶㢮ࣚࢩࣀ࣎ࣜᒓ㨶㢮ࢆ୰ᚰࡋࡓ⩌㞟㸦St. 1–3㸧ࡢ 4 ⩌ศ㢮࣭ᗎิ ࡉࢀࡓ࠸࠼ࡿ㸬ࡉࡽ St. 4–5 6–10 ࡣ㸪ୗὶ᪉ྥ༢ㄪῶᑡࡍࡿ࣐ࢦࡢ ศᕸᇦୗὶ㒊࠾ࡼࡧୗὶഃ࡛ቑຍࡍࡿ」ᩘࡢ㨶✀ࡢศᕸୖ㝈ࢆྵࡴࡇࡽ 㸦Naiman et al., 1988㸹Lasne et al., 2007㸧㸪ࡶ St. 1–3 11–15 ࡢ⩌㞟ࡢ⛣⾜ ᖏ┦ᙜࡍࡿ࠸࠼ࡿ㸬 14 㨶㢮⩌㞟⎔ቃኚ㔞 ᭱㧗Ỉ ௨እࡢ⎔ቃኚ㔞ࡣ㸪ࡍ࡚ࡢᏘ⠇࡛ᗎิࡉࢀࡓྛㄪᰝⅬ᭷ព࡞ 㐺ྜࢆ♧ࡋࡓ㸦Fig. 2-3㸧㸬Ἑᕝᙧែᆺࡢ㔜ᚰࡣࡍ࡚ࡢᏘ⠇ࢆ㏻ࡌ࡚ Aa ᆺࡀ St. 11–15㸪Aa-Bb ⛣⾜ᆺࡀ St. 6–10㸪Bb ᆺࡀ St. 1–3 ㏆⨨ࡋࡓ㸬ᶆ㧗Ἑ ᗋ໙㓄ࡢ࣋ࢡࢺࣝࡣࡍ࡚ࡢᏘ⠇࡛ St. 6–10 11–15 ࡢ㛫㸪ὶ㔞Ỉ㠃ᖜࡣࡍ ࡚ࡢᏘ⠇࡛ St. 1–3 ࢆᣦࡋࡓ㸬᭱పỈ ࡣ㸪⛅ᮇᮇ St. 1–3㸪ኟᮇ St. 6– 10 ࢆᣦࡋࡓ㸬୍᪉㸪᭱㧗Ỉ ࡣ⛅ᮇ᭷ព࡞㐺ྜࢆ♧ࡋ㸪࣋ࢡࢺࣝࡣ St. 6–10 ࢆᣦࡋࡓ㸬࡞࠾㸪ᮇኟᮇࡢ㐺ྜࡣ᭷ព࡛ࡣ࡞ࡗࡓࡀ㸦P = 0.083, 0.052㸧 㸪 ྠᵝ St. 6–10 ࢆᣦࡋࡓ㸬 ⪃ ᐹ 㨶✀ูὶ⛬ศᕸ⛣ື㞀ቨ ࢡࣛࢫࢱ࣮ศᯒ࠾ࡼࡧ㠀ィ㔞ከḟඖᑻᗘᵓᡂἲࡼࡿゎᯒࡽ㸪ᮏㄪᰝỈᇦ ࡢ㨶㢮⩌㞟ࡣ 4 ⩌ศ㢮࣭ᗎิࡉࢀࡓ㸬ࡓࡔࡋ㸪ᮏㄪᰝỈᇦࡣ㨶㢮ࡢ⛣ື 㞀ቨ࡞ࡾᚓࡿሖሐࡸࡀከᩘᏑᅾࡋ࡚࠸ࡓ㸦Fig. 2-1㸧㸬ྛ㨶✀ࡢὶ⛬ศᕸ ⩌㞟ࡢศ㢮࣭ᗎิࡀࡶ⛣ື㞀ቨ࡛Ỵᐃࡉࢀࡿሙྜࡣ㸪⩌㞟ᵓ㐀⎔ቃ せᅉࡢ㛵㐃ࢆ⪃ᐹࡍࡿពࡣ࡞ࡃ࡞ࡿ㸬ࡑࡇ࡛㸪ࡲࡎྛ㨶✀ࡢὶ⛬ศᕸ⩌㞟 ࡢศ㢮࣭ᗎิ㸪࠾ࡼࡧ⛣ື㞀ቨࡢ㛵ಀࡘ࠸࡚⪃ᐹࡍࡿ㸬ὶ⛬ศᕸࡢୖ㝈ࡀ ࢲ࣒ࡸࡢ⨨୍⮴ࡋࡓ㨶✀ࡣࢱ࢝ࣁࣖ㸪࢘ࢢ㸪࣒࢝࣡ࢶ㸪ࣚࢩࣀ࣎ࣜᒓ 㨶㢮࡛࠶ࡗࡓ㸬ࢱ࢝ࣁࣖࡣ St. 1–14 ࡛ぢࡽࢀࡓࡀ㸦Tables 2-3-1, 2-3-2, 2-3-3㸧 㸪 M ࡢୖὶഃศᕸࡋ࡞࠸ࡇࡽ㸦ᮌᮏ㸪ᮍⓎ⾲㸧 㸪 M ࡀ⛣ື㞀ቨ ࠸࠼ࡿ㸦Fig. 2-1㸧㸬ࡋࡋ㸪 M ࡢୖὶഃࡢ St. 15 ࡣ㸪ᮇኟᮇ St. 11–14 ྠࡌࢡࣛࢫࢱ࣮ᒓࡋ㸪⛅ᮇࡶྠࢡࣛࢫࢱ࣮᭱ࡶ㏆࠸ᯞࢆᵓᡂࡋࡓࡇ ࡽ㸦Fig. 2-3㸧㸪 M ࡼࡿศ㢮࣭ᗎิᑐࡍࡿᙳ㡪ࡣᑡ࡞࠸⪃࠼ࡽࢀࡓ㸬 ࢘ࢢ࣒࢝࣡ࢶࡢὶ⛬ศᕸࡢୖ㝈ࡣࡑࢀࡒࢀ St. 13㸪12 ࡛࠶ࡗࡓ㸦Tables 2-3-1, 2-3-2, 2-3-3㸧㸬⚄ཎᕝࡢ St. 12 ࡽሖሐ J ࡲ࡛ࡢ༊㛫ࡣ㸪 」ᩘࡢሖሐࡀᐦ㞟ࡋࡓ㸪 ᮏㄪᰝỈᇦࡢ୰࡛᭱ࡶேⅭᨵኚࡀ㐍ࢇࡔሙᡤ࡛࠶ࡾ㸦Fig. 2-1㸧㸪ࡇࢀࡽࡢሖሐ 15 ⩌ࡀ୧✀ࡢ⛣ື㞀ቨ࡞ࡗࡓྍ⬟ᛶࡀ࠶ࡿ㸬ࡋࡋ㸪⚄ཎᕝࡢ St. 12 ࡰྠ ࡌ⎔ቃኚ㔞ࢆࡶࡘἼᮌྜᕝࡢ St. 11 ࡛ࡣ㸦Table 2-2㸧 㸪ୗὶഃ㨶㢮ࡢ⛣ື㞀ቨ ࡀ࡞࠸ࡶࢃࡽࡎ㸪୧✀ࡢಶయ⩌ᐦᗘࡣ St. 12 ྠᵝపࡃ㸦Fig. 2-2㸧 㸪 ୧ㄪᰝⅬࡣࡍ࡚ࡢᏘ⠇࡛ྠࡌࢡࣛࢫࢱ࣮㸦St. 11–15㸧ྵࡲࢀࡓ㸬ࡇࡢࡇ ࡣ㸪୧✀ࡢศᕸୖ㝈ࢆ St. 12–13 ไ㝈ࡍࡿせᅉࡀ㸪ᑡ࡞ࡃࡶࢲ࣒ࡸሖሐࡼ ࡿ⛣ື㞀ቨࡔࡅ࡛ࡣ࡞࠸ࡇࢆ♧ࡋ࡚࠸ࡿ㸬Ἑᕝ㨶㢮⩌㞟࠾ࡅࡿୗὶ᪉ྥ ࡢ✀ᩘࡢቑຍせᅉࡣ㸪୍ḟ⏕⏘ࡢቑຍ㸦Vannote et al., 1980㸧㸪⏕ᜥሙᡤࡢከᵝ ᛶࡢቑຍ㸦Schlosser, 1982㸧 㸪⏕ᜥሙᡤࢧࢬࡢቑ㸦Rahel and Hubert, 1991㸧➼ ࡀ▱ࡽࢀ࡚࠸ࡿ㸬Ἴᮌྜᕝࡢ St. 11 ⚄ཎᕝࡢ St. 12 ࡛ࡣከࡃࡢ⎔ቃኚ㔞ࡀ㢮 ఝࡋ࡚࠸ࡓࡀ㸪୧ㄪᰝⅬ St. 10 ࡢ㛫࡛ࡣ㸪Ἑᕝᙧែᆺࡀ Aa ᆺࡽ Aa-Bb ⛣ ⾜ᆺኚࡋ㸪Ἑᗋ໙㓄ࡀῶᑡ㸪ᖹᆒỈ㠃ᖜࡣቑຍࡋࡓ㸦Table 2-2㸧 㸬Inoue and Nunokawa㸦2002㸧ࡣ㸪࢚ࢰ࢘ࢢ Tribolodon sachalinensis ࡢ⏕ᜥ㔞ࡀ㸪ୗὶ ቑຍࡍࡿᓊᐤࡾࡢὶ㏿ࡢ㐜࠸㒊ศࡢ㠃✚ẚṇࡢ┦㛵ࢆ♧ࡍሗ࿌ࡋ࡚࠸ࡿ㸬 ࡋࡓࡀࡗ࡚㸪ᮏㄪᰝỈᇦ࡛࣒࢝࣡ࢶ࢘ࢢࡢศᕸࢆไ㝈ࡍࡿせᅉࡣ㸪⏕ᜥሙ ᡤࡢࢧࢬከᵝᛶ࡛࠶ࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡓ㸬ࣚࢩࣀ࣎ࣜᒓ㨶㢮ࡣ St. 1–4 ࡛ ぢࡽࢀ St. 5 ࡛☜ㄆࡉࢀ࡞ࡗࡓࡓࡵ㸦Tables 2-3-1, 2-3-2, 2-3-3㸧 㸪 D㸪ሖሐ E㸪 F ࡢ࠸ࡎࢀࡀ⛣ື㞀ቨ⪃࠼ࡽࢀࡓ㸦Fig. 2-1㸧㸬ࡋࡋ㸪ࡇࢀࡽࢆᣳࢇ࡛㐃⥆ ࡍࡿ St. 4㸪5 ࡣᖖྠࡌࢡࣛࢫࢱ࣮ྵࡲࢀࡓ㸬௨ୖࡢࡇࡽ㸪ᮏ◊✲࠾ ࡅࡿ㨶㢮⩌㞟ࡢศ㢮࣭ᗎิࡣ㸪⛣ື㞀ቨ௨እࡢせᅉࢆᫎࡋ࡚࠸ࡿ⪃࠼ࡽ ࢀࡿ㸬 㨶㢮⩌㞟ኚࡢ⎔ቃせᅉ ୖ࣭ୗὶ᪉ྥ༢ㄪኚࡍࡿࡍ࡚ࡢ⎔ቃኚ㔞ࡀ㸪ᗎิࡉࢀࡓㄪᰝⅬᑐ ࡋ࡚᭷ព࡞㐺ྜࢆ♧ࡋࡓࡇࡽ㸦Fig. 2-3㸧㸪ᮏㄪᰝỈᇦࡢ㨶㢮⩌㞟ᵓ㐀ࡣὶ ⛬ἢࡗࡓ⎔ቃ໙㓄ᙉࡃつᐃࡉࢀ࡚࠸ࡿ⪃࠼ࡽࢀࡓ㸬ࡇࡢ⤖ᯝࡣ㸪ࢧࢣ⛉ 㨶㢮ࢆ୰ᚰࡍࡿ⩌㞟ࡽࢥ⛉㨶㢮ࢆ୰ᚰࡍࡿ⩌㞟ࡢ⛣⾜ࡀ㸪Ỉ ࡸᆅ ᙧ࠸ࡗࡓࡁ࡞✵㛫ࢫࢣ࣮ࣝࡢ⎔ቃ໙㓄つᐃࡉࢀࡿࡋࡓሗ࿌㸦Naiman et al., 1988㸹Rahel and Hubert, 1991㸹Jackson et al., 2001㸧୍⮴ࡋ࡚࠸ࡿ㸬୍᪉㸪 16 ᭱㧗Ỉ ࡣ㸪ࢧࢣ⛉㨶㢮ࢆ୰ᚰࡍࡿ⩌㞟ࡽࢥ⛉㨶㢮ࢆ୰ᚰࡍࡿ⩌㞟 ࡢ⛣⾜࠾ࡅࡿ㔜せ࡞せᅉ࡛࠶ࡿࡀ㸦Rahel and Hubert, 1991㸹Taniguchi et al., 1998㸹Ỉ㔝࣭ᚚໃ㸪2000㸹Lasne et al., 2007㸹Mercado-Silva et al., 2012㸧 㸪ᮏ◊✲ ࡛ࡣ⛅ᮇ௨እ᭷ព࡞㐺ྜࢆ♧ࡉ࡞ࡗࡓ㸬ἙᕝỈ ࡣ୍⯡ୗὶ᪉ྥୖ᪼ ࡍࡿࡀ㸦Vannote et al., 1980㸧㸪ᮏㄪᰝỈᇦ࡛ࡣỈࡢὶධࡼࡾୗὶ㒊࡛పࡃ୰ ὶ㒊࡛㧗ࡗࡓ㸦Table 2-2㸧㸬ࡲࡓ㸪ᮏゎᯒ⏝࠸ࡓ envfit 㛵ᩘࡣ㸪⎔ቃኚ㔞 ᗎิࡉࢀࡓㄪᰝⅬࡢ㛫⥺ᙧ㛵ಀࢆ௬ᐃࡋ࡚࠸ࡿ㸦Oksanen, 2013㸧㸬ࡇࡢࡓ ࡵ㸪᭱㧗Ỉ ࡣㄪᰝⅬ⩌㐺ྜࡋࡃࡗࡓ⪃࠼ࡽࢀࡿ㸬ࡓࡔࡋ㸪᭱㧗Ỉ ࡢ࣋ࢡࢺࣝࡣ㸪ࡍ࡚ࡢᏘ⠇࡛ St. 11–15 ࡽ St. 6–10 ࡢ᪉ྥࢆᣦࡋ࡚࠾ࡾ㸦Fig. 2-3㸧 㸪࣐ࢦࡢಶయ⩌ᐦᗘࡣࡇࡢ᪉ྥῶᑡࡋ࡚࠸ࡓ㸦Fig. 2-2㸧 㸬ࡋࡓࡀࡗ࡚㸪 ᮏㄪᰝỈᇦ࠾࠸࡚ࡶ㸪᭱㧗Ỉ ࡣࢧࢣ⛉㨶㢮ࢆ୰ᚰࡋࡓ⩌㞟ࡽࢥ⛉㨶 㢮ࢆ୰ᚰࡋࡓ⩌㞟ࡢኚ㛵ࡋ࡚࠸ࡿ⪃࠼ࡽࢀࡓ㸬 㨶✀ูಶయ⩌ᐦᗘ⎔ቃせᅉ ᮏ◊✲࡛ࡣ㸪⎔ቃኚ㔞ࡢ㸪☜ㄆ✀ᩘࡸከࡃࡢ㨶✀ูಶయ⩌ᐦᗘࡶὶ⛬ ἢࡗࡓ༢ㄪ࡞ኚࢆ♧ࡋࡓࡀ㸪ࢱ࢝ࣁࣖࡢಶయ⩌ᐦᗘࡣὶ⛬ἢࡗࡓኚࢆ♧ ࡉࡎ㸪St. 6–11 ࡛ప࠸ᓠᆺࡢὶ⛬ศᕸࢆ♧ࡋࡓ㸬ࡲࡓ㸪࢘ࢢ࣒࢝࣡ࢶࡢ ಶయ⩌ᐦᗘࡣୗὶ᪉ྥ༢ㄪቑຍࡋࡓࡀ㸪St. 4–5 ࡛ᒁᡤⓗప࠸್ࢆ♧ࡋࡓ 㸦Fig. 2-2㸧㸬ᮏ◊✲࡛ࡣ㸪ࡇࢀࡽࡢᐦᗘኚᑐᛂࡍࡿ⎔ቃኚ㔞ࡣᚓࡽࢀ࡞ ࡗࡓ㸬ὶ⛬ἢࡗࡓ㨶㢮⩌㞟ࡢኚࡢ࠺ࡕ㸪ࢧࢣ⛉㨶㢮ࢆ୰ᚰࡍࡿ⩌㞟ࡽ ࢥ⛉㨶㢮ࢆ୰ᚰࡍࡿ⩌㞟ࡢ⛣⾜ࡣ㸪Ỉ ࡸᆅᙧ࠸ࡗࡓࡁ࡞✵㛫ࢫ ࢣ࣮ࣝࡢ⎔ቃ໙㓄ࡀ㔜せ࡞ാࡁࢆ♧ࡍࡀ㸪ࡼࡾᑠࡉ࡞✵㛫ࢫࢣ࣮࡛ࣝࡣ㸪ࡑࢀ ࡽࡣูࡢ≀⌮࣭⏕≀ⓗせᅉࡀാࡃࡇࡀ▱ࡽࢀ࡚࠸ࡿ㸦Jackson et al., 2001㸹 Inoue and Nunokawa, 2002㸹Torgersen et al., 2006㸧 㸬ࡋࡓࡀࡗ࡚㸪ࡇࢀࡽࡢ㨶✀ࡢ ಶయ⩌ᐦᗘࡢኚࡣ㸪ᮏ◊✲࡛ ᐃࡋࡓ⎔ቃኚ㔞௨እࡢせᅉࡀ㛵ࡋ࡚࠸ࡿ ⪃࠼ࡽࢀࡓ㸬୍᪉㸪ࡇࢀࡽࡢㄪᰝⅬࡣ㸪㨶㢮⩌㞟ᵓ㐀ࡽ࠸ࡎࢀࡶ⛣⾜ᖏ ᥎ᐃࡉࢀࡓ㸬⛣⾜ᖏ࡛ࡣ㸪ᵓᡂ✀ࡀศᕸࡢ࿘⦕㒊⏕ᜥࡍࡿࡓࡵ㸪⎔ቃᨵኚ ᑐࡍࡿឤཷᛶࡀ㧗࠸ᣦࡉࢀ࡚࠸ࡿ㸦Naiman et al., 1988㸹Aarts and Nienhuis, 17 2003㸹Lasne et al., 2007㸧 㸬ᮏㄪᰝỈᇦ࡛᥎ᐃࡉࢀࡓ⛣⾜ᖏࡢୖὶഃࡣሖሐ⩌ ࡀᏑᅾࡋ࡚࠸ࡓࡇࡽ㸦Fig. 2-1㸧㸪ࢱ࢝ࣁࣖ㸪࢘ࢢ㸪࣒࢝࣡ࢶࡢಶయ⩌ᐦ ᗘࡣேⅭᨵኚࡢᙳ㡪ࢆཷࡅ࡚࠸ࡿྍ⬟ᛶࡶ⪃࠼ࡽࢀࡿ㸬ࡲࡓ㸪Aarts and Nienhuis 㸦2003㸧ࡣ㸪ሖሐࡼࡿேⅭᨵኚࡀ㐍ࢇࡔ⛣⾜ᖏ࡛ࡣ㸪≉ࢽࢵࢳᖜࡢ⊃࠸ࢧ ࢣ⛉㨶㢮➼ࡢಖࡀᅔ㞴࡛ษ㏕ࡋࡓၥ㢟࡛࠶ࡿᣦࡋ࡚࠸ࡿ㸬ᮏㄪᰝỈᇦ䛷 䛿䜰䝬䝂䛿ୗὶ᪉ྥ䛻༢ㄪῶᑡ䛧䠈☜ㄆ䛥䜜䛯ὶ⛬䠄St. 2–15䠖15.2 km䠅䛾 7 ௨ୖ 㸦St. 2–10㸸11.2 km㸧䛜⛣⾜ᖏࢆྵࡴୗὶഃỈᇦᏑᅾࡋࡓ㸬ࡲࡓ㸪㨶✀ ␗࡞ࡾ㸪ᮏㄪᰝỈᇦࡼࡾୗὶ䛾㔝ᕝ࡛䛿☜ㄆࡉࢀ࡚࠸䛺䛔䛣䛸䛛䜙㸦⎔ቃᗇ㸪 1987㸧䠈ᮏㄪᰝỈᇦ䛾⛣⾜ᖏ䛷䛿䜰䝬䝂䛾ಖ䛻䜒ὀព䜢ᡶ䛖ᚲせ䛜䛒䜛䛸⪃䛘䜙䜜 䜛䠊ᮏㄪᰝỈᇦ䛻䛚䛡䜛࣐ࢦಶయ⩌ᐦᗘࡢไ㝈せᅉࡣ㸪᭱㧗Ỉ 㸪Ἑᕝᙧែᆺ ➼ࡢᆅᙧ᮲௳㸪㐟⁺ࡼࡿ㔮⋓➼᥎ᐃࡉࢀࡿ㸦ᮌᮏ㸪2013㸧㸬ࡋࡋ㸪ሖ ሐ䛾ୗὶഃ䛷⏕䛨䜛ἙᕝᆅᙧࡢኚࡸἙᗋ䛾పୗ䠄బᕝ䞉୰ᮧ䠈2010䠅䠈Ἑᗋ䛾㟢 ┙䠄୰ᮧ䠈2011䠅➼䛿䠈Ỉ῝䠈ὶ㏿䠈Ἑᗋᮦᩱ➼䛾ኚ䜢క䛳䛶㨶㢮䛾⏕ᜥሙᡤ䜢 ຎ䛥䛫䜛䛸⪃䛘䜙䜜䜛䛣䛸䛛䜙䠈ேⅭᨵኚ䛿࣐ࢦࡢୗὶ᪉ྥࡢῶᑡࡶ㛵ࡋ ࡚࠸ࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡓ㸬௨ୖࡢࡇࡽ㸪Ἑᕝୖὶᇦ࠾ࡅࡿ㨶㢮⩌㞟ࡢ ኚᵝᘧࢆ᫂ࡽࡋ㸪 ≉⛣⾜ᖏ࠾࠸࡚ᑠࡉ࡞✵㛫ࢫࢣ࣮࡛ࣝỈ῝㸪 ὶ㏿㸪 Ἑᗋᮦᩱ➼ࡢ⎔ቃኚ㔞ࢆㄪࡿࡇࡣ㸪ᚋࡢேⅭᨵኚࡢᙳ㡪ホ౯㸪ᆅᇦࡢ⏕ ែ⣔ಖ㸪≉ᐃ✀ࡢᣢ⥆ⓗ⏝➼ࢆᅗࡿୖ࡛㔜せ⪃࠼ࡽࢀࡿ㸬 18 Û(ޗ N Û(ޗ 1 Ûޗ1 3 D Fukuoka Prefecture E 2 4 F Oita Prefecture 5 Ogata River Seto Inland Sea G 6 Tosumi River Ono River 7 Kumamoto Prefecture 8 waterfalls Ûޗ1 C man-made dam 9 Mt. Sobo B spring A H Kohbaru Stream I K J N O survey station 11 Hakiai Stream 13 L M 12 10 14 15 1 km Fig. 2-1. Stations established in the Ogata River and its tributaries (Kohbaru and Hakiai streams) (right) in northeastern Kyushu Island (left). Numerals and capital letters (A–O) indicate survey stations and dams/falls, respectively. 19 Density (Number of individuals/100 m2) 60 50 40 30 20 10 0 Oct. 2003 10 Feb. 2004 8 6 4 2 0 140 120 100 80 60 40 20 0 Aug. 2004 1 2 3 O. masou ishikawae 4 5 6 7 P. oxycephalus 8 9 10 Station T. hakonensis 11 12 13 C. temminckii 14 15 Others Fig. 2-2. Longitudinal changes in population densities of four dominant fishes (Oncorhynchus masou ishikawae, Phoxinus oxycephalus, Candidia temminckii and Tribolodon hakonensis) and other fish species in the Ogata River and its tributaries (Kohbaru and Hakiai streams) in October 2003 (top), February 2004 (middle) and August 2004 (bottom). 20 0.0 -0.5 2 Bb 3 CspOp -0.5 1.0 -1.0 0.0 Max -0.5 Axis 2 0.0 0.5 Ct 1.0 15 9 10 Th 6 8 15 4 Alt Gra Om AaBb 12 Aa 14 13 11 Po 7 -1.0 7 9 8 6 10 3 1 2 4 5 14 15 11 12 13 -1.5 -1.0 4 5 Bb 1 2 Wid 3 Dis PeOo Min Rsp Op -0.5 0.0 0.5 1.0 1.0 Aug. 2004 Om 11 Aa13 14 Po 12 5 Dis Wid Min Rsp 0.5 0.8 0.6 0.4 7 8 Th Oo AaBb Ct 910 1 -1.5 Feb. 2004 Alt Gra 6 Max -1.0 2 1 3 6 9 10 7 8 15 4 11 13 14 5 12 0.2 -0.5 0.0 0.5 Min -1.0 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.0 Cophenetic distance 1.0 0.5 Pe 1.0 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 Oct. 2003 15 11 12 13 14 2 1 3 9 10 6 7 8 4 5 Dis 1 Wid OpRsp 2 Pe 5 4 Oo 8 Po 6 Th AaBb 12 11 13 7 Aa Bb 14 3 Om 10 Ct 9 Csp -1.5 Station Max Gra Alt -1.0 -0.5 0.0 Axis 1 Fig. 2-3. See the next page for figure caption because of its length. 21 15 0.5 1.0 1.5 Fig. 2-3. Dendrograms and ordination plots resulting from cluster analysis and non-metric multidimensional scaling (NMDS) of fish assemblage structures among 15 stations in the Ogata River and its tributaries (Kohbaru and Hakiai streams) in October 2003 (top), February 2004 (middle) and August 2004 (bottom). In dendrograms, dotted lines indicate cophenetic distances classifying 15 fish assemblages into three or four clusters. In NMDS plots, numerals, italic and roman letters represent survey stations, fish species (Om: Oncorhynchus masou ishikawae, Po: Phoxinus oxycephalus, Th: Tribolodon hakonensis, Ct: Candidia temminckii, Pe: Pseudogobius esocinus esocinus, Oo: Odontobutis obscura, Rsp: Rhinogonius sp., Csp: Cobitis sp.) and environmental variables (Aa, AaBb, and Bb: topographic types [based on Kani (1978)], Alt: altitude, Gra: gradient, Wid: mean wetted width, Dis: discharge, Min: minimum DMWT [daily mean water temperature], Max: maximum DMWT), respectively. Fish species and topographic types are plotted as centroids with respect to site scores. Vectors indicate directions of increasing environmental gradients and strength of their correlations with site scores. Solid and dotted lines of vectors indicate significant (P < 0.05) or non-significant relationships with site scores based on random permutation tests (10, 000 iterations). 22 23 A B C D E F G H I J K L M N O Kawabe Tinda Harajiri Ichigome Sangome Gogome Dam Falls Falls Falls Dam Dam Falls Dam Dam (5) Dam Dam Dam Falls Falls Falls 47 51 61 74.6 74.7 75.4 76.6 82.3 82.9-83.4 83.8 84.4 84.7 85.6 86 86.3 -23 -19 -9 4.6 4.7 5.4 6.6 12.3 12.9-13.4 13.8 14.4 14.7 15.6 16 16.3 8.4 20 20 5.6 1.1 1.4 11 1.4 1.0-1.4 2.9 5.2 8.9 16 17 11 + - *** Presence (+) or absence (-) of fish way. ** DF St. 1 (km): Distance from St.1 (km). * DFRM: Distance from river mouth. Symbol Name Falls / Dam DFRM (km)* DF St. 1 (km)** Height (m) Fish way*** Hakiai streams). Only dams of height > 1 m indicated. Table 2-1. Dams and falls present below (A–C) and within (D–O) the survey area of the Ogata River and its tributaries (Kohbaru and 24 *** DMWT: Daily mean water temperature. ** Topographic types follow Kani (1978). * DF St. 1 (km): Distance from St.1 (km). Station 1 2 3 4 5 6 7 8 9 10 11 12 DF St. 1 (km)* 0.0 1.0 2.3 3.8 5.9 7.0 8.8 10.0 11.4 12.2 13.0 13 Survey area (m2) 2,263 2,567 1,573 650 999 688 929 751 883 1,090 491 745 Topographic type** Bb Bb Bb Bb Aa-Bb Aa-Bb Aa-Bb Aa-Bb Aa-Bb Aa-Bb Aa Aa Altitude (m) 232 238 255 260 284 296 325 342 384 397 414 416 Gradient (%) 0.6 1.35 0.3 1.14 1.14 1.62 1.43 2.86 1.69 2.11 2.7 Mean wetted width (m) 21.2 20.6 23 12.2 10.4 15.2 13.4 12.2 16.7 7.8 8.9 Discharge (m3/s) 2.81 2.13 2.33 2 1.04 0.75 0.32 0.35 0.19 0.13 0.09 0.1 DMWT (°C)*** Minimum 10.2 10.4 10.1 11 7.1 5.4 5.6 5.6 3.9 4.1 4.4 5.4 Maximum 20.3 20.6 20.8 21 21.5 23.6 21.6 21.4 22.2 22 19.8 21 the Hakiai Stream 820 m upstream from its confluence with the Kohbaru Stream. 13 14 473 Aa 486 5.4 10 0.1 5.1 19 14 15 479 Aa 604 11 9.1 0.1 2.6 19 15 16 411 Aa 820 21 7.1 0 0.2 18 Table 2-2. Environmental data for 15 stations in the Ogata River and its tributaries (Kohbaru and Hakiai streams). Station 11 was located in 25 October 2003 Oncorhynchus masou ishikawae Phoxinus oxycephalus Tribolodon hakonensis Candidia temminckii Odontobutis obscura Pseudogobio esocinus esocinus Rhinogobius sp. Opsariichthys platypus Cobitis sp. Oncorhynchus masou masou Anguilla japonica Number of species Number of individuals Station 1 17 226 22 3 675 405 170 2 60 25 812 186 523 2 3 1 7 8 7 1,111 1,519 1,609 344 267 450 1 1 23 25 1 3 95 7 4 84 4 5 375 11 345 3 15 1 5 included. Fishes arranged in order of upper limits of distribution ranges. 6 172 4 50 106 4 3 5 6 8 4 1 9 45 24 52 9 1 18 58 64 60 10 6 5 5 5 185 496 131 201 1 1 23 11 27 112 105 219 28 150 7 3 38 8 21 9 11 3 261 14 17 230 12 2 76 36 40 13 2 87 24 63 14 1 35 35 15 203 2,906 1,388 1,466 7 12 107 276 22 3 1 11 6,391 Total individuals < 5 cm in total length of non-benthic cyprinid fish species (P. oxycephalus, T. hakonensis, C. temminckii, and O. platypus) not and Hakiai streams) in October 2003. Newly emerged fry of amago O. masou ishikawae and masu salmon O. masou masou, and Table 2-3-1. Number of fish species and individuals observed by snorkelers at 15 stations in the Ogata River and its tributaries (Kohbaru 26 February 2004 Oncorhynchus masou ishikawae Phoxinus oxycephalus Tribolodon hakonensis Candidia temminckii Odontobutis obscura Pseudogobio esocinus esocinus Rhinogobius sp. Opsariichthys platypus Oncorhynchus masou masou Cyprinus carpio Number of species Number of individuals Station 6 264 5 201 55 1 1 10 44 11 50 2 4 153 2 134 1 6 202 112 5 61 11 12 1 3 1 2 2 4 1 14 14 5 included. Fishes arranged in order of upper limits of distribution ranges. 2 18 1 17 6 1 3 3 7 1 3 27 24 2 8 2 5 3 2 9 3 38 1 36 1 10 3 4 1 1 2 11 4 68 6 59 2 1 12 3 60 21 38 1 13 2 18 13 5 14 1 9 9 15 53 360 105 124 4 5 275 5 1 1 10 933 Total individuals < 5 cm in total length of non-benthic cyprinid fish species (P. oxycephalus, T. hakonensis, C. temminckii, and O. platypus) not and Hakiai streams) in February 2004. Newly emerged fry of amago O. masou ishikawae and masu salmon O. masou masou, and Table 2-3-2. Number of fish species and individuals observed by snorkelers at 15 stations in the Ogata River and its tributaries (Kohbaru 27 2 4 5 13 6 2 1114 566 1,000 154 128 25 489 7 2 5 4 2 31 391 3 12 1 20 13 1 1 1 3 8 820 8 50 1 116 6 8 38 18 272 195 58 133 62 85 23 76 131 7 10 58 56 62 1 9 14 54 25 17 10 12 13 23 51 68 114 306 325 5 5 2 11 58 53 14 68 15 Total 377 575 650 6,102 164 20 781 38 615 1,420 11 11 61 9 2 371 441 460 1,295 18 5 35 10 172 183 2 23 58 3 5 1 2 1 1 9 10 12 7 6 6 5 7 5 5 3 4 2 2 1 13 1,268 1,959 2,207 720 1,064 1,003 506 586 187 113 142 364 393 111 68 10,691 1 *Survey at St. 15 conducted in October 2004 due to low water visibility from late August to late September in 2004. August 2004 Oncorhynchus masou ishikawae Phoxinus oxycephalus Tribolodon hakonensis Candidia temminckii Odontobutis obscura Pseudogobio esocinus esocinus Rhinogobius sp. Opsariichthys platypus Cobitis sp. Plecoglossus altivelis altivelis Oncorhynchus masou masou Cyprinus carpio Carassius sp. Number of species Number of individuals Station included. Fishes arranged in order of upper limits of distribution ranges. individuals < 5 cm in total length of non-benthic cyprinid fish species (P. oxycephalus, T. hakonensis, C. temminckii, and O. platypus) not and Hakiai streams) in August 2004*. Newly emerged fry of amago O. masou ishikawae and masu salmon O. masou masou, and Table 2-3-3. Number of fish species and individuals observed by snorkelers at 15 stations in the Ogata River and its tributaries (Kohbaru ➨ 3 ❶ ࣐ࢦᾋୖ⛶㨶ࡢὶ⛬ศᕸ ㏆ᖺ㸪ࢧࢣ⛉㨶㢮ࡢᅾ᮶ಶయ⩌ࡢಖ⁺ᴗ࣭㐟⁺ࡼࡿ⏝ࡢ୧❧ࢆᅗࡿ ࡓࡵ㸪ὶ㨶ሙࡢࢰ࣮ࢽࣥࢢ⟶⌮ࡀ᪥ᮏྛᆅ࡛ヨࡳࡽࢀ࡚࠸ࡿ㸦ຍ⸨㸪1991㸹 ᮌᮧ㸪2004㸹୰ᮧ㸪2009㸧㸬ࢰ࣮ࢽࣥࢢ⟶⌮࡛ࡣ㸪ᅾ᮶ಶయ⩌ࡢศᕸࣃࢱ࣮ࣥ ⮬↛⏕⏘ࡢᐇែᇶ࡙࠸࡚⁺ሙู␗࡞ࡿ⟶⌮ᡭἲࡀ㐺⏝ࡉࢀࡿࡓࡵ 㸦୰ᮧ㸪 2009㸧㸪ᑐ㇟㨶✀ࡢ⏕άྐࢆ㏻ࡌࡓ⛣ື࠾ࡼࡧ⮬↛⏕⏘ࡢᐇែ➼ࡢ⏕ែᏛⓗ▱ ぢࢆᢕᥱࡍࡿᚲせࡀ࠶ࡿ㸬⮬↛⏕⏘ࡢᐇែࡣ㸪⏘༸ᗋࡸᾋୖ⛶㨶ࡢศᕸ➼ ᇶ࡙࠸࡚ᢕᥱࡍࡿࡇࡀྍ⬟࡛࠶ࡾ㸪࣡ࢼ Salvelinus leucomaenis㸦୰ᮧ㸪 2009㸧㸪ࢧࢡ࣐ࣛࢫ㸦ᮡⱝ㸪1999㸧 㸪ࣅ࣐࣡ࢫ Oncorhynchus masou subsp.㸦ᑿ ⏣㸪2011㸧㸪ࢱࢭࣚ࢘ࢧࢣ Salmo salar ࣈࣛ࢘ࣥࢺࣛ࢘ࢺ Salmo trutta 㸦Foldvik et al., 2010 ཧ↷㸧➼࡛ሗ࿌ࡀ࠶ࡿ㸬࣐ࢦࡸ࣐࣓ࣖࡢἙᕝᆺಶయ⩌ࡣ ࡁ࡞Ἑᕝෆ⛣ືࢆࡏࡎ㸪⏘༸ሙᡤࡑࢀ௨እࡢ⏕άሙᡤࡀ㏆᥋ࡍࡿྍ⬟ᛶࡀ ᣦࡉࢀ࡚࠸ࡿࡇࡽ㸦Nakano et al., 1990㸹Sakata et al., 2005㸧㸪ࡑࡢ⏕⏘ ࡢᐇែࢆᢕᥱࡍࡿࡓࡵࡣ⏕ᜥᇦయࢆྵࡴࢫࢣ࣮ࣝࡢㄪᰝࡀᚲせ⪃࠼ࡽࢀ ࡿ㸬ࡲࡓ㸪࣐ࢦࡢ⏘༸ሙᡤࡀ⏕άሙᡤ㏆᥋ࡍࡿ࡞ࡽࡤ㸪⏘༸ᗋ㸪ᾋୖ⛶㨶 ࠾ࡼࡧᡂ㨶ࡢศᕸࣃࢱ࣮ࣥࡢ㛫ࡣఱࡽࡢ㛵ಀࡀ࠶ࡿணࡉࢀࡿࡀ㸪ࡑࢀ ࡽࡢ㛵ಀࡣ࡛᫂࠶ࡿ㸬ࡉࡽ㸪࣐ࢦࡢᾋୖ⛶㨶ࡣᡂ㛗కࡗ࡚ᚤ⏕ᜥሙᡤ ࢆኚࡉࡏࡿࡇࡀ▱ࡽࢀࡿࡀ㸦ྡ㉺㸪1988㸧 㸪ᾋୖ┤ᚋࡢ⛣ື㛵ࡍࡿ▱ ぢࡣ࡞࠸㸬 ᮏ◊✲ࡣ㸪㔝ᕝỈ⣔⥴᪉ᕝᨭὶ⚄ཎᕝ࠾࠸࡚㸪࣐ࢦࡢ⮬↛⏕⏘ࡢ ᐇែࢆ᫂ࡽࡍࡿࡇࢆ┠ⓗࡋࡓ㸬ࡑࡢࡓࡵ㸪ὶᇦࢫࢣ࣮ࣝ࠾ࡅࡿᾋ ୖ⛶㨶ࡢὶ⛬ศᕸࢆ₯Ỉ┠どほᐹࡼࡾ᫂ࡽࡍࡿࡶ㸪ᗂ㨶ᮇ௨㝆ࡢ ಶయࡢὶ⛬ศᕸࡢẚ㍑ࢆ⾜࠸㸪ࡑࢀࡽࡢὶ⛬ศᕸࡢ㛵ಀᇶ࡙࠸࡚㸪≉ᮏ ✀ࡢ⏕άྐึᮇ࠾ࡅࡿ⛣ືᵝᘧࡘ࠸࡚㆟ㄽࢆ⾜࠺㸬 ᮦᩱ᪉ἲ 28 ㄪᰝሙᡤ ᮏ◊✲࡛ࡣ㸪⥴᪉ᕝ༑ゅᕝࡢྜὶⅬ㸦ᇶ‽Ⅼ㸧ࡽ⚄ཎᕝࡢ࣐ࢦ⏕ᜥୖ 㝈࡛࠶ࡿྜ┠ࡢࡲ࡛ࡢ⣙ 16.4 km ࢆㄪᰝỈᇦࡋ࡚㸪࠾ࡶὶἣࡀኚࡍ ࡿᆅⅬࢆቃ 14 ㄪᰝ༊ศࡅࡓ㸦Fig. 3-1㸧㸬ྛㄪᰝ༊ࡢ㛗ࡉࡣ㸪࣐ࢦᾋୖ⛶ 㨶ࡢ₯Ỉ┠どせࡍࡿ㛫ࢆ⪃៖ࡋ࡚ 1 km ๓ᚋ㸦0.73–1.80 km㸹ᖹᆒ ± ᶆ‽ ೫ᕪ = 1.17 ± 0.28 km㸧ࡋࡓ㸬ࢧࢣ⛉㨶㢮ᡂ㨶ࡢ㐳ୖࡣⴠᕪ 1 m ௨ୖࡢࡸሖ ሐ➼࡛㜼ᐖࡉࢀࡿࡇࡀ▱ࡽࢀ࡚࠸ࡿࡀ㸦㧗ᶫ㸪1988㸧㸪ᮏㄪᰝỈᇦࡣⴠᕪ 11–17 m ࡢ⮬↛ࡀ Sec. 6 1 ࣨᡤ㸪Sec. 14 3 ࣨᡤᏑᅾࡋ㸪ⴠᕪ 1 m ௨ୖࡢ ሖሐ➼ࡀ Sec. 5㸪11㸪12㸪13 ࡑࢀࡒࢀ 2 ᇶ㸪1 ᇶ㸪6 ᇶ㸪2 ᇶタ⨨ࡉࢀ࡚࠸ࡿ 㸦Fig. 3-1㸧㸬࣐ࢦࡢᨺὶࡣ Sec. 11 ࢆ୰ᚰẖᖺ⾜ࢃࢀ࡚࠸ࡿࡀ㸪᭱ୖὶ㒊ࡢ Sec. 14 ࡣබⓗ࡞ᨺὶグ㘓ࡣ࡞࠸㸦㔝ᕝ⁺ᴗ༠ྠ⤌ྜ㸪⚾ಙ㸧㸬 ⏕ᜥ⎔ቃ ྛㄪᰝ༊ࡢἙᕝᙧែᆺ㸦ྍඣ㸪1978㸧ࡣ Sec. 1–4 ࡛ Bb ᆺ㸪Sec. 5–10 ࡛ Aa-Bb ⛣⾜ᆺ㸪Sec. 11–14 ࡛ Aa ᆺ࡛࠶ࡗࡓ㸦Table 3-1㸧㸬⦰ᑻ 1/5000 ࡢᆅᅗࡽ⟬ฟ ࡋࡓἙᗋ໙㓄ࡣ Sec. 1–11 ࡛ 3%ᮍ‶㸪Sec. 12–14 ࡛ 5.3–21.1%࡛࠶ࡗࡓ㸬ᇶ‽Ⅼ ࡽ⣙ 100 m 㛫㝸࡛ ᐃࡋࡓỈ㠃ᖜࡽồࡵࡓᖹᆒỈ㠃ᖜࡣ㸪 Sec. 1–3 ࡛⣙ 20 m㸪 Sec. 14 ࡛⣙ 7 m ࡛࠶ࡗࡓ㸬2003 ᖺ 8 ᭶–2005 ᖺ 4 ᭶Ỉ ࢹ࣮ࢱ࣮ࣟ࢞ 㸦StowAway TidbiT Temp Logger, Onset Computer㸧ࡼࡾグ㘓ࡉࢀࡓ 1 㛫ẖࡢ Ỉ ࢹ࣮ࢱᇶ࡙ࡃ᪥ᖹᆒ᭱㧗Ỉ ࡣ Sec. 1–3 ࡛ 20°C ྎ㸪 Sec. 4–10 ࡛ 21–24°C㸪 Sec. 12–14 ࡛ 20°C ᮍ‶㸪 ྠ᭱పỈ ࡣ Sec. 1–3 ࡛ 9.5°C ๓ᚋ㸪Sec. 4–13 ࡛ 3–7°C㸪 Sec. 14 ࡛ࡣ 0.2°C ࡛࠶ࡗࡓ㸦Table 1㸧 㸬Sec. 1–5 ࡣỈ⩌㸦⣙ 15°C㸧ࡀᏑᅾࡍ ࡿࡓࡵ㸪Sec. 1–3 ࡢ᭱㧗–᭱పỈ ᕪࡣᑠࡉࡗࡓ㸬ྛㄪᰝ༊ෆࡢὶࢀࡢᏳᐃࡋ ࡓሙᡤ ᐃ᩿㠃ࢆタᐃࡋ㸪ᦠᖏࣉࣟ࣌ࣛᘧὶ㏿ィ㸦3631 ᆺ㸪ᶓἙ㟁ᶵ㸧ࢆ⏝ ࠸ࡓ 1 Ⅼἲ㸦Ỉ㠃ᖜࡢ 10–15%ศࡋࡓ ᐃ᩿㠃ࡢ㠃✚ 60%Ỉ῝ࡢὶ㏿ࢆ ࡌࡿ㸹୰ᮧ㸪1997㸧࡛ồࡵࡓὶ㔞ࡣ㸪Sec. 1–4 ࡛⣙ 2.0 m3/s㸪Sec. 14 ࡛ࡣ 0.02 m3/s ࡛࠶ࡗࡓ㸦Table 3-1㸧㸬 29 ࣐ࢦᾋୖ⛶㨶ࡢศᕸㄪᰝ ᮏㄽᩥ࡛ࡣ㸪ஂಖ㸦1980㸧ࡼࡿࢧࢡ࣐ࣛࢫࡢⓎ⫱ẁ㝵༊ศࢆཧ⪃㸪ὸࡃ ὶ㏿ࡀ㐜࠸ᓊ㏆ࡃࡢሙᡤࢆ⏝ࡍࡿ㛗⣙ 5 cm ࡲ࡛ࡢಶయࢆᾋୖ⛶㨶㸦newly emerged fry㸧㸪Ỉ῝ࡀ῝ࡃὶ㏿ࡀࡁ࡞ὶᚰ㒊⛣ືࡍࡿ㛗⣙ 5 cm ௨㝆ࡢಶ యࢆࡍ࡚ᗂ㨶ᮇ௨㝆ࡢಶయ㸦post-fry㸧ࡋ࡚༊ูࡋࡓ㸬 ᮏㄪᰝ࡛ࡣ㸪2004 ᖺ⛅⏘༸ࡉࢀࡓᖺ⣭⩌ࡢ࣐ࢦᾋୖ⛶㨶ࢆᑐ㇟㸪ㄪᰝ Ỉᇦࡢὶ⛬㸦16.4 km㸧࠾࠸࡚₯Ỉ┠どㄪᰝࢆ⾜ࡗࡓ㸬ࡓࡔࡋ㸪ㄪᰝỈᇦࡢ ୖ࣭ୗὶ࡛Ỉ ᕪࡀࡁ࠸ࡓࡵ㸦Table 3-1㸧㸪⏘༸ᗋࡽࡢ࣐ࢦ⛶㨶ࡢᾋୖࡢ ┠Ᏻ࡛࠶ࡿ✚⟬ ᗘࡀ⣙ 800°C 㐩ࡍࡿᮇ㸦ᮏⲮ㸪1977㸹௨ୗ㸪ᾋୖ᪥㸧ࡀ ㄪᰝ༊ࡈ␗࡞ࡿணࡉࢀࡓ㸬ࡑࡇ࡛㸪ㄪᰝ๓ᖺࡢỈ ࢹ࣮ࢱ 2004 ᖺ 2–3 ᭶⾜ࡗࡓணഛⓗ࡞₯Ỉ┠どㄪᰝࡢ⤖ᯝ㸦ᮌᮏ㸪ᮍⓎ⾲㸧ࡽᾋୖ ᮇࢆ᥎ᐃࡋࡓ㸬࣐ࢦࡢ⏘༸᪥ࢆ 2003 ᖺ 11 ᭶ 1 ᪥௬ᐃࡍࡿ㸪ྛㄪᰝ༊ࡢ ᾋୖ᪥ࡣୗὶ᪩ࡃ㸦Sec. 1㸸ྠᖺ 12 ᭶ᮎ㸧 㸪ୖὶ㐜࠸㸦Sec. 14㸸2004 ᖺ 4 ᭶ึ᪪㸧᥎ᐃࡉࢀࡓ㸬ᐇ㝿ணഛⓗ₯Ỉ┠どㄪᰝ࡛ࡣ㸪᥎ᐃᾋୖ᪥ྠ ㄪࡋ࡚ᾋୖ⛶㨶ࡀࡳࡽࢀ㸪Ỉ ࡢ㧗࠸ Sec. 1–4 ࡛ࡣᾋୖ᪥ࡽ 1 ࣨ᭶௨ෆ⛶ 㨶ࡀᗂ㨶ࡲ࡛Ⓨ⫱ࡍࡿྍ⬟ᛶࡀ♧ࡉࢀࡓ㸦ᮌᮏ㸪ᮍⓎ⾲㸧 㸬௨ୖࡢࡇࡽ㸪 ㄪᰝࡣ᥎ᐃᾋୖ᪥ࡽ 1 ࣨ᭶௨ෆୗὶഃࡽୖὶഃ㐍ࡵࡿࡇࡋ㸪2005 ᖺ 1 ᭶ 7 ᪥–3 ᭶ 28 ᪥➨ 1 ᅇㄪᰝࢆ⾜ࡗࡓ㸬ࡲࡓ㸪࣐ࢦ⛶㨶ࡢᾋୖ᪥ࡣ⏘ ༸᪥ࡢ㐪࠸ࡸⓎ⏕ࡢಶయᕪࡼࡾኚືࡍࡿࡶ㸪ᗂ㨶ᮇ㐩ࡋࡓಶయࡽ 㡰ὶᚰ㒊⛣ືࡍࡿࡓࡵ㸦ྡ㉺㸪1988㸧 㸪ྛㄪᰝ༊ࡢᾋୖ⛶㨶ᩘࡣᾋୖ᪥ ௨㝆ቑຍࡋࡓࡢࡕῶᑡࡍࡿணࡉࢀࡓ㸬ࡉࡽ㸪ᾋୖ┤ᚋࡢ࣐ࢦ⛶㨶ࡀ ࡢ⛬ᗘ⛣ືࡍࡿࡢࡣ࡛᫂࠶ࡗࡓ㸬ࡋࡓࡀࡗ࡚㸪⮬↛⏕⏘ࡢᐇែ࠾ࡼࡧᾋ ୖ⛶㨶ࡢ⛣ືᵝᘧࢆᢕᥱࡍࡿࡓࡵࡣ㸪ᾋୖ⛶㨶ᩘࡢከ࠸ᮇྜࢃࡏࡓ」ᩘ ᅇࡢㄪᰝࡀᚲせ࡛࠶ࡿࡇࡽ㸪1 ᅇ┠ࡢㄪᰝࡽ 11–32 ᪥ࡢ㛫㝸ࢆ࠾࠸࡚㸪 㸬ㄪᰝࡀ 1 ᪥࡛ ྠᖺ 1 ᭶ 21 ᪥–4 ᭶ 15 ᪥ 2 ᅇ┠ࡢㄪᰝࢆᐇࡋࡓ㸦Fig. 3-2㸧 ⤊ࢃࡽ࡞ࡗࡓሙྜࡣ 3–5 ᪥ᚋᮍ㒊ศࡢㄪᰝࢆ⾜ࡗࡓ㸬 ୖグ᪥⛬ࡢ᪥୰㸦9–17 㸧㸪Ỉ୰࣓࢞ࢿ㸪ࢩࣗࣀ࣮ࢣࣝ㸪ࢻࣛࢫ࣮ࢶ➼ ࢆ╔⏝ࡋࡓㄪᰝ⪅ 2 ྡࡀྛㄪᰝ༊ࡢୗὶഃࡽ㟼₯Ỉࡋ㸪୧ᓊἢ࠸ୖὶ 30 ᪉ྥ㐍ࡳ࡞ࡀࡽᾋୖ⛶㨶ࢆ┠どィᩘࡋࡓ㸬Ỉ㠃ᖜࡀ⣙ 10 m ௨ୖࡢㄪᰝ༊㸦Sec. 1–10㸧࡛ࡣἙ㐨୰ኸ㒊ࡶ࠺ 1 ྡࡢㄪᰝ⪅ࢆ㓄ࡋࡓ㸬₯Ỉ┠どࡣ㸪ᓊἢ࠸ࡢὶ ㏿ࡀ㐜࠸ሙᡤ㸪Ỉ῝ࡀ⣙ 1 m ௨ὸࡢሙᡤ㸪᳜≀ࡸᒾࡢୗ➼㸪࣐ࢦࡸࢧࢡ࣐ࣛ ࢫࡢᾋୖ⛶㨶ࡀศᕸࡍࡿሙᡤ㸦ྡ㉺㸪1988㸹Nagata and Yanai, 2002㸧ࢆ୰ ᚰ⾜࠸㸪ὶ㏿ࡢ㏿࠸Ἑ㐨୰ኸ㒊ࡸ⣙ 1 m ௨῝ࡢᒙ࡛ࡣᐇࡋ࡞ࡗࡓ㸬Ỉ῝ ⣙ 20 cm ࢆୗᅇࡿὸ࠸ሙᡤ࡛ࡣ㸪῝࠸ሙᡤࡽὸ࠸ሙᡤࢆぢ㏻ࡍ࡞ࡋ࡚Ỉ୰ ࡢど⏺ࢆ☜ಖࡋࡓ㸬▼ࡸ᳜≀➼࡛ど⏺ࡀ㐽ࡽࢀࡿሙྜࡣ㸪ᑠᯞ➼࡛ྍど㒊 ࢆ่⃭ࡋ࡚ᾋୖ⛶㨶ࢆ㏨㑊ࡉࡏ㸪どㄆࡋࡓ㸬௨ୖࡢ᪉ἲ࡛㸪1 ࡲࡓࡣ」ᩘಶయ ࡀᐃࡍࡿᆅⅬࡈ㸪࡛ࡁࡿ㝈ࡾṇ☜ᾋୖ⛶㨶ࡢಶయᩘࢆィᩘࡋࡓ㸬 ࣐ࢦᾋୖ⛶㨶ࡢ㐟Ὃຊࡢᙅࡉࡸ㧗࠸どㄆᛶࡢࡓࡵ㸪 ┠どࡢຠ⋡ࡣ㧗࠸ࡳ࡞ࡋ㸪 ᮏ◊✲࡛ࡣ┠どィᩘࡢㄗᕪࢆ↓どࡍࡿࡇࡋࡓ㸬 ㄪᰝ⪅ࡣ㸪ᾋୖ⛶㨶ࡢᐃⅬࡽ㝣ୖࡢグ㘓⪅ಶయᩘࢆ▱ࡽࡏ㸪グ㘓⪅ࡣ ⦰ᑻ 1/2500 ࡢᆅᅗୖࡑࢀࡽࢆグ㘓ࡋࡓ㸬ㄪᰝ⤊ᚋ㸪ᆅᅗୖࡢᇶ‽Ⅼࡽ ᐃⅬࡲ࡛ࡢ㊥㞳ࢆ࢟ࣝࣅ࣓࣮ࢱ࣮࡛ ᐃࡍࡿࡶ㸪☜ㄆࡉࢀࡓᾋୖ⛶㨶 ᩘࢆྛㄪᰝ༊ࡢỈ⾲㠃✚㸦Table 3-1㸧࡛㝖ࡋ࡚㸪100 m2 ࠶ࡓࡾࡢಶయᩘᐦᗘࢆ ⟬ฟࡋࡓ㸬࡞࠾㸪㔝ᕝ⁺ᴗ༠ྠ⤌ྜ」ᩘࡢ㔮ࡾᅋయࡣ㸪ᮏㄪᰝᮇ㛫ᙜヱ Ỉᇦ࡛⛶㨶ࢆᨺὶࡋ࡚࠾ࡽࡎ㸦㔝ᕝ⁺ᴗ༠ྠ⤌ྜ㸪⚾ಙ㸧㸪ࡲࡓ☜ㄆࡉࢀࡓᾋ ୖ⛶㨶ࡣ୍⯡ⓗᨺὶࡉࢀࡿᗂ㨶㸦㛗 5 cm ௨ୖ㸧ẚᑠࡉ࠸㸦2.5–5 cm㸧 ࡇࡽ㸪ࡍ࡚⮬↛⏕⏘ࡉࢀࡓ࣐ࢦࡳ࡞ࡋࡓ㸬 ᗂ㨶ᮇ௨㝆ࡢ࣐ࢦࡢศᕸㄪᰝ 2003 ᖺ 10 ᭶ 21–29 ᪥㸦⛅ᮇ㸧 㸪2004 ᖺ 2 ᭶ 2–10 ᪥㸦ᮇ㸧㸪2004 ᖺ 8 ᭶ 6– 24 ᪥㸦ኟᮇ㸹㝆㞵ࡢࡓࡵ Sec. 14 ࡣ 10 ᭶ 6 ᪥㸧ࡢ᪥୰㸦9–17 㸧㸪ྛㄪᰝ༊ ෆᑡ࡞ࡃࡶ℩ῡࢆ୍ࡘࡎࡘྵࡴὶ㊰㛗 42–146 m㸦93.8 ± 36.7 m㸧ࡢㄪᰝ ⠊ᅖࢆタࡅ㸪ᗂ㨶ᮇ௨㝆ࡢ࣐ࢦࡢ₯Ỉ┠どㄪᰝࢆ⾜ࡗࡓ㸬ࡇࢀࡽࡢㄪᰝ⠊ᅖ ࡣ㸪➨ 2 ❶㸦ᮌᮏ㸪2015㸧ࡢㄪᰝⅬࡢ࠺ࡕ St. 1–10 St. 12–15 ྠࡌሙᡤ ࡛࠶ࡿ㸬ὶ㊰㛗㸪ୖ࣭ୗὶ➃୰㛫Ⅼࡢ 3 ࣧᡤࡢỈ㠃ᖜࡢᖹᆒ್ࡽ⟬ฟࡋࡓ ྛㄪᰝ⠊ᅖࡢỈ⾲㠃✚ࡣ 411–2,567 m2㸦1,035 ± 658 m2㸧࡛࠶ࡗࡓ㸦Table 3-1㸧㸬 31 ὶἣᕪࡼࡿ┠ど⋡ࡢᕪ␗ࢆ᭱ᑠࡍࡿࡓࡵ㸪Ỉ㠃ᖜࡀ⣙ 5 m ௨ୗࡢሙᡤ࡛ࡣ 1 ྡ㸪5–10 m ࡛ࡣ 2 ྡ㸪⣙ 10 m ௨ୖ࡛ࡣ 3 ྡࡀ㸪Ỉ୰࣓࢞ࢿ㸪ࢩࣗࣀ࣮ࢣࣝ㸪 ࢻࣛࢫ࣮ࢶ➼ࢆ╔⏝ࡋ࡚ほᐹࡋࡓ㸬1 ྡࡢㄪᰝ࡛ࡣ㸪ὶ㊰୰ᒾ▼➼ࡀ࠶ࡿ ሙྜࡣ㌟ࢆ㞃ࡋ࡞ࡀࡽ㸪࡞࠸ሙྜࡣᓊἢ࠸ୖὶ᪉ྥ⛣ືࡋ࡞ࡀࡽᑐᓊࢆぢ Ώࡋ࡚ὶ㊰యࢆほᐹࡋࡓ㸬2 ྡ௨ୖ࡛ࡣ㸪Ỉ㠃ᖜࢆࡰᆒ➼ㄪᰝேᩘ ࡾᙜ࡚㸪ဨࡀୖὶ᪉ྥ⛣ືࡋ࡞ࡀࡽ㐍⾜᪉ྥࡢྑഃࢆ┠どࡋ㸪ྑᓊࡢㄪᰝ ⪅ࡣᕥഃࡶほᐹࡋࡓ㸦Thurow, 1994㸧 㸬࠸ࡎࢀࡢሙྜࡶ㸪ୗὶഃࡽ㟼₯Ỉ ࡋ㸪ほᐹ୰ࡣᛴ⃭࡞ືࡁࢆ㑊ࡅ㸪࣐ࢦᐹ▱ࡉࢀ࡞࠸ࡼ࠺ὀពࢆᡶࡗࡓ㸬2 ྡ௨ୖࡢほᐹࡢ⤊ࡣ㸪ྛ⮬ࡢほᐹỈᇦෆࡢ࣐ࢦࡢ⛣ฟධࢆ☜ㄆࡋࡓ㸬 ࣐ࢦࡢಶయᩘࢆ 5 cm ᖜࡢ㛗㝵⣭ูグ㘓ࡋ㸪ྛㄪᰝ⠊ᅖ࡛ᚓࡽࢀࡓಶయᩘ ࢆỈ⾲㠃✚࡛㝖ࡋ࡚ 100 m2 ࠶ࡓࡾࡢಶయᩘᐦᗘࢆ⟬ฟࡋࡓ㸬ࡲࡓ㸪ྛ㛗㝵⣭ ࡢ୰ኸ್ಶయᩘࡽྛㄪᰝ༊ࡢᖹᆒ㛗ศᩓࢆồࡵࡓ㸬ᮇࡣ୍㒊ࡢㄪ ᰝ༊࡛ᾋୖ⛶㨶ࢆㄆࡵࡓࡀ㸪ୖグࡢᾋୖ⛶㨶ศᕸㄪᰝࡢΰྠࢆ㑊ࡅࡿࡓࡵࢹ ࣮ࢱྵࡵ࡞ࡗࡓ㸬 ⤫ィゎᯒ ྛㄪᰝ༊ࡢ࣐ࢦᾋୖ⛶㨶ࡢಶయᩘᐦᗘ㸪ᗂ㨶ᮇ௨㝆ࡢಶయᩘᐦᗘ࠾ࡼࡧ 㛗ศᕸࡘ࠸࡚㸪Jonckheere-Terpstra ᳨ᐃࡼࡾὶ⛬ἢࡗࡓഴྥࢆㄪࡓ㸬ࡲ ࡓ㸪2 ᅇࡢㄪᰝ࡛ᚓࡓᾋୖ⛶㨶ࡢὶ⛬ศᕸ㸪3 Ꮨ⠇ࡢᗂ㨶ᮇ௨㝆ࡢಶయࡢὶ⛬ศ ᕸ㸪࠾ࡼࡧᾋୖ⛶㨶ᗂ㨶ᮇ௨㝆ࡢಶయࡢὶ⛬ศᕸࡢ┦㛵ࢆ Spearman ࡢ㡰┦ 㛵ಀᩘ࡛ホ౯ࡋࡓ㸬᭷ពỈ‽ࡣ 0.05 ࡋ㸪ᗂ㨶ᮇ௨㝆ࡢಶయࡢὶ⛬ศᕸ㛵ࡍ ࡿ 3 ᅇࡢ┦㛵ศᯒ࡛ࡣ㸪Bonferroni ⿵ṇࡋࡓ᭷ពỈ‽㸦0.017㸧ࢆ⏝࠸ࡓ㸬ࡍ ࡚ࡢゎᯒࡣ⤫ィࢯࣇࢺ R㸦R Development Core Team, 2005㸧ࢆ࣮࣋ࢫࡋࡓࣇࣜ ࣮⤫ィࢯࣇࢺ EZR㸦⚄⏣㸪2012㸧࡛⾜ࡗࡓ㸬 ⤖ ᯝ ࣐ࢦᾋୖ⛶㨶ࡢὶ⛬ศᕸ 32 ᾋୖ⛶㨶ࡣᓊἢ࠸ࡢὸ࠸⦆ὶ㒊㸪᳜≀ࡸᒾࡢୗ➼ከࡃࡳࡽࢀ㸪2 ᅇࡢㄪᰝ ࡛☜ㄆࡉࢀࡓ࣐ࢦᾋୖ⛶㨶ࡢ⥲ಶయᩘࡣ㸪ࡑࢀࡒࢀ 1,374 ಶయ 1,476 ಶయ ࡛࠶ࡗࡓ㸦Table 3-2㸪Fig. 3-3㸧㸬୧ㄪᰝ࠾ࡅࡿྛㄪᰝ༊ࡢಶయᩘࡣ㸪Sec. 2㸪3㸪 13 ࠾ࡼࡧ Sec. 11㸪12 ࠾࠸࡚ࡸࡸ␗࡞ࡗ࡚࠸ࡓࡀ㸦Table 3-2㸧 㸪ྛㄪᰝ༊ࡢಶ యᩘᐦᗘࡣ➨ 1 ᅇㄪᰝ࡛ 0–4.59 ಶయ/100 m2㸦1.0 ± 1.5 ಶయ/100 m2㸧㸪➨ 2 ᅇ࡛ 0–4.70 ಶయ/100 m2㸦1.0 ± 1.6 ಶయ/100 m2㸧ࡢ⠊ᅖ࠶ࡾ㸪୧ㄪᰝ㛫࡛᭷ព࡞ᕪ ࡣࡳࡽࢀ࡞ࡗࡓ㸦Wilcoxon ࡢ➢ྜࡁ㡰᳨ᐃ㸪 V = 32㸪 P = 0.61㸹Table 3-2㸧 㸬 ࡲࡓ㸪ྛㄪᰝ༊ࡢಶయᩘᐦᗘࡣ୧ㄪᰝ㛫࡛᭷ព࡞ṇࡢ┦㛵ࢆ♧ࡋࡓ㸦Spearman ࡢ㡰┦㛵ಀᩘ㸪rs = 0.95㸪P < 0.001㸧 㸬୧ㄪᰝࡶୖὶ㒊ࡢ Sec. 11–14 ࡛ಶయ 㸪ࡘ㐃⥆ⓗ ᩘᐦᗘࡀ㧗ࡃ㸦➨ 1 ᅇ➨ 2 ᅇࡢᖹᆒ್㸸1.88–4.64 ಶయ/100 m2㸧 ᾋୖ⛶㨶ࡀ☜ㄆࡉࢀࡓ㸦Table 3-2㸪Fig. 3-3㸧 㸬୍᪉㸪Sec. 1㸪Sec. 4 ࡢୖὶഃ 㸦⣙ 1.1 km㸧 㸪Sec. 5 ࡢୗὶഃ㸦⣙ 0.6 km㸧࠾ࡼࡧ Sec. 6 ࡛ࡣ㸪2 ᅇࡢㄪᰝࡶ ᾋୖ⛶㨶ࡀࢇ☜ㄆࡉࢀ࡞ࡗࡓ㸦Fig. 3-3㸧 㸬Sec. 7–10 ࡢಶయᩘᐦᗘࡣୖ ὶ㒊ࡼࡾࡸࡸపࡃ㸦0.21–0.42 ಶయ/100 m2㸧㸪Sec. 5 ࡛ࡣ⊃࠸⠊ᅖ㸦⣙ 0.3 km㸧 㸬Sec. 2–4 ࡢಶయᩘᐦ ࡛ࡢࡳᒁᡤⓗᾋୖ⛶㨶ࡀ☜ㄆࡉࢀࡓ㸦0.29 ಶయ/100 m2㸧 㸬ྛㄪᰝ༊ࡢಶయᩘᐦᗘࡣୖὶ ᗘࡣᴟࡵ࡚పࡗࡓ㸦0.04–0.08 ಶయ/100 m2㸧 㧗࠸ഴྥࢆ♧ࡋࡓ㸦Jonckheere-Terpstra ᳨ᐃ㸪➨ 1 ᅇ㸸J = 81.5㸪P < 0.001㸹 ➨ 2 ᅇ㸸J = 80㸪P < 0.001㸧 㸬 ᗂ㨶ᮇ௨㝆ࡢ࣐ࢦࡢὶ⛬ศᕸ ᗂ㨶ᮇ௨㝆ࡢ࣐ࢦࡣ Sec. 1 ࢆ㝖ࡃ 13 ㄪᰝ༊࡛☜ㄆࡉࢀ㸪ಶయᩘࡣኟᮇከ ࡃ㸪ᮇᑡ࡞ࡗࡓ㸦Tables 3-3-1㸪3-3-2㸪3-3-3㸧㸬ྛㄪᰝ༊ࡢಶయᩘᐦᗘࡣ ୖὶഃࡢ Sec. 11–14 ࡛㧗ࡃ㸦⛅ᮇ㸸2.3–8.5 ಶయ/100 m2㸹ᮇ㸸0.8–4.4 ಶయ/100 m2 㸹 ኟ ᮇ 㸸 6.2–16.5 ಶ య /100 m2 㸹 Fig. 3-4 㸧㸪 ୖ ὶ 㧗 ࠸ ഴ ྥ ࢆ ♧ ࡋ ࡓ 㸦Jonckheere-Terpstra ᳨ᐃ㸪⛅ᮇ㸸J = 80.5㸪P < 0.001㸹ᮇ㸸J = 73㸪P < 0.01㸹 ኟᮇ㸸J = 81.5㸪P < 0.001㸧㸬ࡓࡔࡋ㸪Sec. 7 ࡛ࡣ⛅ᮇኟᮇẚ㍑ⓗ㧗࠸ᐦᗘࡀ 㸬ࡲࡓ㸪 ほᐹࡉࢀࡓ㸦⛅ᮇ㸸2.5 ಶయ/100 m2㸹ኟᮇ㸸4.1 ಶయ/100 m2㸹Fig. 3-4㸧 ྛㄪᰝ༊ࡢಶయᩘᐦᗘࡣ␗࡞ࡿᏘ⠇㛫࡛᭷ព࡞ṇࡢ┦㛵ࢆ♧ࡋࡓ㸦⛅ᮇ 33 ᮇ㸸rs = 0.75㸪P = 0.002㸹ᮇኟᮇ㸸rs = 0.76㸪P = 0.002㸹⛅ᮇኟᮇ㸸rs = 0.92㸪 P < 0.001㸹Fig. 3-4㸧㸬ㄪᰝ༊㛫ࡢ㛗ࡣ⛅ᮇኟᮇ᭷ព࡞ᕪࡀࡳࡽࢀ 㸦Kruskal-Wallis ᳨ᐃ㸪⛅ᮇ㸸df = 11㸪Ȥ2 = 32.7㸪P < 0.001㸹ᮇ㸸df = 6㸪Ȥ2 = 4.3㸪 㸪ᖹᆒ㛗ࡣୖὶഃᑠࡉ࠸ഴ P = 0.63㸹ኟᮇ㸸df = 11㸪Ȥ2 = 85.6㸪P < 0.001㸧 ྥࢆ♧ࡋࡓ㸦Jonckheere-Terpstra ᳨ᐃ㸪⛅ᮇ㸸J = 17.5㸪P = 0.03㸹ኟᮇ㸸J = 10.5㸪 P < 0.01㸹Tables 3-3-1㸪3-3-2㸪3-3-3㸧 㸬ࡲࡓ㸪ྛㄪᰝ༊࠾ࡅࡿ㛗ࡢศᩓࡣ⛅ ᮇ࡛᭷ព␗࡞ࡗ࡚࠾ࡾ㸦Fligner-Killeen ᳨ᐃ㸪⛅ᮇ㸸df = 11㸪Ȥ2 = 42.3㸪P < 㸪 0.001㸹ᮇ㸸df = 6㸪Ȥ2 = 10.4㸪P = 0.11㸹⛅ᮇ㸸df = 11㸪Ȥ2 = 17.1㸪P = 0.11㸧 ࡍ࡚ࡢㄪᰝ࡛ୖὶࡁ࠸ഴྥࡀ࠶ࡗࡓ㸦Jonckheere-Terpstra ᳨ᐃ㸪⛅ᮇ㸸 J = 185.5㸪P < 0.001㸹ᮇ㸸 J = 118.5㸪P < 0.001㸹ኟᮇ㸸J = 177㸪P < 0.001㸧㸬 ᗂ㨶ᮇ௨㝆ࡢಶయᾋୖ⛶㨶ࡢὶ⛬ศᕸࡢ㛵ಀ 2004 ᖺ 8 ᭶࠾ࡅࡿྛㄪᰝ༊ࡢᗂ㨶ᮇ௨㝆ࡢ࣐ࢦಶయᩘᐦᗘ㸪 ⩣ᖺ 1㸫4 ᭶࠾ࡅࡿᾋୖ⛶㨶ࡢಶయᩘᐦᗘࡢ㛫ࡣ㸪᭷ព࡞ṇࡢ┦㛵ࡀㄆࡵࡽࢀࡓ㸦Fig. 3-5㸹ᗂ㨶ᮇ௨㝆ࡢ࣐ࢦ➨ 1 ᅇㄪᰝࡢᾋୖ⛶㨶㸸rs = 0.82㸪P < 0.001㸹ᗂ㨶 ᮇ௨㝆ࡢ࣐ࢦ➨ 2 ᅇㄪᰝࡢᾋୖ⛶㨶㸸rs = 0.80㸪P < 0.001㸧 㸬 ⪃ ᐹ ᮏ◊✲ࡣ㸪࣐ࢦࡢ༡㝈㏆ࡢಶయ⩌࠾࠸࡚㸪ᾋୖ⛶㨶࠾ࡼࡧᗂ㨶ᮇ௨㝆 ࡢಶయᩘᐦᗘࡀㄪᰝᮇ㛫ࢆ㏻ࡌ࡚ୖὶഃ㧗ࡃ㸪୧⪅ࡢὶ⛬ศᕸࣃࢱ࣮ࣥࡣ ୍⮴ࡋࡓഴྥࢆ♧ࡍࡇࢆ᫂ࡽࡋࡓ㸬ࡲࡓ㸪ᾋୖ⛶㨶ࡢᐦᗘࡀ 2 ᅇࡢㄪᰝ 㛫࡛ࡁ࡞ᕪࢆ♧ࡉ࡞ࡗࡓࡇࡽ㸪ᮏㄪᰝࡣ⛶㨶ࡢᾋୖ┒ᮇᐇࡉࢀ㸪 ࡑࡢὶ⛬ศᕸࡢᐇែࢆᴫࡡᢕᥱࡍࡿࡇࡀ࡛ࡁࡓ⪃࠼ࡽࢀࡿ㸬௨ୗ࡛ࡣ㸪୧ ⪅ࡢὶ⛬ศᕸࡢᡂ❧せᅉࡑࢀࡽࡀ୍⮴ࡍࡿ⌮⏤ࡘ࠸࡚⪃ᐹࡋࡓࡢࡕ㸪ྛㄪ ᰝ༊࠾ࡅࡿ⮬↛⏕⏘ࡢホ౯࠾ࡼࡧࡑࡢຓ㛗ࡢ᪉ἲࡘ࠸᳨࡚ウࡍࡿ㸬 ᗂ㨶ᮇ௨㝆ࡢ࣐ࢦࡢὶ⛬ศᕸᡂ❧せᅉ 34 㜰⏣㸦1993㸧ࡣ㸪ᕞࡢᒣᆅὶ࠾ࡅࡿ࣐࣓ࣖࡢ⏕ᜥᐦᗘࡀ㸪᪩℩ῡࡢ 㐃⥆ᙧែࡢྜ࠾ࡼࡧἙᗋ໙㓄ṇࡢ┦㛵ࢆ♧ࡍࡇࢆሗ࿌ࡋ㸪ࡑࡢ⌮⏤ࡋ ࡚㸪᪩℩ῡࡢ㐃⥆ᙧែࡀࡶࡓࡽࡍỈ୰ࡢⓑἻࡸἙᗋࡢᒾࡸὶᮌࡀእᩛࡽࡢ ࡃࢀሙᡤࡋ࡚ᶵ⬟ࡍࡿࡇ㸪ῡ㢌ࡀ㣵࡞ࡿὶୗື≀ࡢ᭱ࡶຠ⋡ⓗ࡞౪⤥ Ⅼ࡞ࡿࡇゝཬࡋ࡚࠸ࡿ㸬ࡲࡓ㸪Inoue et al.㸦1997㸧ࡣ㸪ᾏ㐨࠾ࡅࡿࢧ ࢡ࣐ࣛࢫᗂ㨶ࡢ⏕ᜥࢆไ㝈ࡍࡿせᅉࡀኟᮇࡢ᭱㧗Ỉ ࠾ࡼࡧỈ୰ࡢಽὶᮌ㸪Ἑ ᓊ㒊ࡢ࠼ࡄࢀࡸ᳜⏕➼ࡢ࢝ࣂ࣮ࡢ㔞࡛࠶ࡿࡇࢆሗ࿌ࡋ࡚࠾ࡾ㸪ᚋ⪅ࡀࢧࢡࣛ ࣐ࢫࡢ⏕ᜥᐦᗘࢆつᐃࡍࡿ⌮⏤ࡋ࡚㸪࢝ࣂ࣮ࡀዲ㐺࡞᥇㣵ሙᡤ㸪⿕㣗ࣜࢫࢡ ࡢ㍍ῶ㸪ಶయ㛫ᖸ΅ࡢ⦆స⏝➼ࢆࡶࡓࡽࡋ㸪⎔ቃᐜຊࢆ㧗ࡵࡿࡇࢆᣦ ࡋ࡚࠸ࡿ㸬ᮏㄪᰝᆅ࠾࠸࡚ಶయᩘᐦᗘࡀ㧗ࡗࡓ Sec. 11–14 ࡢ≀⌮⎔ቃ᮲௳ ࡣ㸪Ἑᕝᙧែᆺࡀ Aa ᆺ࡛Ἑᗋ໙㓄ࡀࡁࡃ㸦2.6–21.1%㸧 㸪᭱㧗Ỉ ࡀ 20°C ௨ ୗ࡛࠶ࡾ㸪ୖグࡢሗ࿌ࡀ♧ࡍ⎔ቃ᮲௳୍⮴ࡋ࡚࠸ࡓ㸦Table 3-1㸧㸬ᮏ◊✲࡛ࡣ ࢝ࣂ࣮ࡸὶୗື≀㛵ࡍࡿᐃ㔞ⓗㄪᰝࢆᐇࡋ࡚࠸࡞࠸ࡀ㸪Sec. 11–14 ࡣ Aa ᆺ ࡢἙᕝᙧែᆺࢆ♧ࡍࡶୗὶഃࡢㄪᰝ༊ẚ㇏࡞Ἑ␁ᯘࢆకࡗ࡚࠾ࡾ 㸦ᮌᮏ㸪ᮍⓎ⾲㸧㸪ࡃࢀሙᡤ࡞ࡿỈ୰ࡢᒾࡸ࠼ࡄࢀ㸪࠾ࡼࡧ࣐ࢦࡢ㔜 せ࡞㣵⏕≀࡛࠶ࡿ㝣⏕᪻㢮㸦ྡ㉺㸪1988㸧ࡀ┦ᑐⓗ㇏ᐩ࡛࠶ࡿ⪃࠼ ࡽࢀࡿ㸬ࡇࢀࡽࡢࡇࡽ㸪ᮏㄪᰝỈᇦ࠾ࡅࡿᗂ㨶ᮇ௨㝆ࡢ࣐ࢦࡢὶ⛬ศ ᕸࡣ㸪᭱㧗Ỉ ࡢ㸪ἙᕝᙧែᆺࡸἙᗋ໙㓄࠾ࡼࡧἙ␁ᯘ࠸ࡗࡓᆅᙧ᮲௳ ࡀࡶࡓࡽࡍ⎔ቃᐜຊࡼࡾつᐃࡉࢀ࡚࠸ࡿ⪃࠼ࡽࢀࡿ㸬 ࣐ࢦᾋୖ⛶㨶ࡢὶ⛬ศᕸࡢỴᐃせᅉ ࢧࢣ⛉㨶㢮ࡢᾋୖ⛶㨶ࡢὶ⛬ศᕸࡣ㸪⏘༸ᗋࡢ⨨ᾋୖ⛶㨶ࡢ⛣ືᵝᘧࡀ ┦ᙳ㡪ࡋ࡚ᙧᡂࡉࢀࡿ⪃࠼ࡽࢀࡿ㸦Foldvik et al., 2010㸧㸬ᮏㄪᰝỈᇦ࠾ ࡅࡿ⏘༸ᗋࡢὶ⛬ศᕸࡣ᫂ࡽ࡛ࡣ࡞࠸ࡀ㸪ᮏᕞࡢ࣐ࢦᕞࡢ࣐࣓࡛ࣖࡣ ᶆ㆑ᤕㄪᰝࡼࡾᙉ࠸ᐃఫᛶࡀㄆࡵࡽࢀ㸪⏘༸ሙᡤ⏕άሙᡤࡀ㏆᥋ࡍࡿྍ ⬟ᛶࡀ♧ࡉࢀ࡚࠸ࡿ㸦Nakano et al., 1990㸹Sakata et al., 2005㸧 㸬ࡲࡓ㸪ᕞࡢ ࣐ࢦ࣓࡛࣡ࡶྠᵝࡢㄪᰝࡽᐃఫᛶࡀ☜ㄆࡉࢀ࡚࠸ࡿ㸦ᚨග㸪2004㸹㏆⸨࣭ ➉ୗ㸪2005㸧 㸬ࡇࡢ࠺ࡕ㸪㏆⸨࣭➉ୗ㸦2005㸧ࡢㄪᰝᆅࡣ㸪ᮏㄪᰝỈᇦὶධࡍ 35 ࡿἼᮌྜᕝࡢୖὶ㒊࡛࠶ࡾ㸦Fig. 3-1㸧 㸪Ἑᕝᙧែᆺ㸦Aa ᆺ㸧 㸪Ἑᗋ໙㓄㸦9–15%㸧㸪 ᭱㧗Ỉ 㸦20°C ௨ୗ㸧ࡣ Sec. 11–14 ࡢ≀⌮⎔ቃ᮲௳㢮ఝࡋ࡚࠸ࡿ㸬ࡉࡽ㸪 ᚨග㸦2004㸧ࡣ Aa-Bb ⛣⾜ᆺ࡛Ἑᗋ໙㓄ࡀᑠࡉ࠸㸦2.9–3.2%㸧ㄪᰝᆅ࡛ࡶ࣐ ࢦࡢ㧗࠸ᐃఫᛶࢆሗ࿌ࡋ࡚࠸ࡿ㸬ᮏㄪᰝ࡛ぢࡽࢀࡓᗂ㨶ᮇ௨㝆ࡢ࣐ࢦࡢὶ⛬ ศᕸࡢᏳᐃᛶ㸦Fig. 3-4㸧ࡣ㸪ࡑࢀࡽࡢ⛣ືࡀᑡ࡞࠸ࡇࢆᫎࡍࡿࡶࡢ⪃࠼ ࡽࢀࡿ㸬ࡇࢀࡽࡢࡇࡽ㸪ᮏㄪᰝỈᇦࡢ࣐ࢦࡶᙉ࠸ᐃఫᛶࢆ᭷ࡋ㸪ࡑࡢ⏘ ༸ᗋࡣᗂ㨶ᮇ௨㝆ࡢಶయࡢ⏕άሙᡤ㏆ᙧᡂࡉࢀ㸪⏘༸ᗋࡢὶ⛬ศᕸࡣᗂ㨶 ᮇ௨㝆ࡢಶయࡢὶ⛬ศᕸ㢮ఝࡍࡿྍ⬟ᛶࡀ㧗࠸᥎ᐹࡉࢀࡿ㸬 ᾋୖ⛶㨶ࡢ⛣ືᵝᘧ㛵ࡋ࡚㸪ࢧࢡ࣐ࣛࢫࢱࢭࣚ࢘ࢧࢣࡢⓎ║༸ࡢᨺ ὶᐇ㦂࡛ࡣ㸪ᾋୖ⛶㨶ࡀୖὶࡣࢇ⛣ືࡏࡎ㸪⏘༸ᗋ㏆␃ࡲࡿୗ ὶ᪉ྥ⛣ືࡍࡿሗ࿌ࡉࢀ࡚࠸ࡿ㸦Nagata, 2002㸹Einum et al., 2011, 2012㸧 㸬 ࡋࡋ㸪ᮏㄪᰝᆅ࡛ࡣ࣐ࢦࡢᾋୖ⛶㨶ᗂ㨶ᮇ௨㝆ࡢಶయࡢὶ⛬ศᕸࡀ୍⮴ ࡍࡿഴྥࢆ♧ࡋࡓ㸬ୖ࡛⪃ᐹࡋࡓࡼ࠺㸪ࡶࡋ࣐ࢦࡢ⏘༸ᗋᗂ㨶ᮇ௨㝆ࡢ ಶయࡢὶ⛬ศᕸࡀ୍⮴ࡍࡿ࡞ࡽࡤ㸪ࡇࡢᾋୖ⛶㨶ᗂ㨶ᮇ௨㝆ࡢಶయࡢὶ⛬ศ ᕸࡢ୍⮴ࡣ㸪ᾋୖ┤ᚋࡢ࣐ࢦ⛶㨶ࡀ⏘༸ᗋ㏆␃ࡲࡿࡇ࡛ᡂ❧ࡋࡓࡶࡢ ᥎ᐹࡉࢀࡿ㸬ࡲࡓ㸪11–32 ᪥ࡢ㛫㝸ࢆ࠾࠸ࡓ 2 ᅇࡢㄪᰝࢆ㏻ࡌ࡚ᾋୖ⛶㨶ࡢ ὶ⛬ศᕸࡀኚࡋ࡞ࡗࡓࡇ㸪≉ࢇ⛶㨶ࡀぢࡽࢀ࡞࠸⣙ 1 km ࢃ ࡓࡿὶ⛬ࡀ」ᩘ㸪Ᏻᐃࡋ࡚Ꮡᅾࡋࡓࡇࡣ㸪ᾋୖ⛶㨶ࡀ⣙ 1 ࣨ᭶㛫㸪ྛㄪᰝ༊ ࡢ㛗ࡉ㸦⣙ 1 km㸧ࢆ㉸࠼ࡿࡼ࠺࡞ࡁ࡞⛣ືࢆࡋ࡞࠸ྍ⬟ᛶࢆ♧၀ࡍࡿ㸬 ⮬↛⏕⏘ࡢホ౯ࢰ࣮ࢽࣥࢢ⟶⌮ ࢰ࣮ࢽࣥࢢ⟶⌮࡛ࡣ㸪ᅾ᮶ಶయ⩌ࡢಖࢆᅗࡿࡓࡵ㸪ᅾ᮶ಶయ⩌ࡢศᕸࣃ ࢱ࣮ࣥ⮬↛⏕⏘ࡢᐇែᇶ࡙࠸࡚⁺ሙู␗࡞ࡿ㐟⁺⟶⌮࠾ࡼࡧᨺὶᡭἲ ࡀ㐺⏝ࡉࢀࡿ㸦ᮌᮧ㸪2004㸹୰ᮧ㸪2009㸧 㸬୰ᮧ㸦2009㸧ࡢࢰ࣮ࢽࣥࢢ⟶⌮ࡢᣦ 㔪ࡼࢀࡤ㸪ᅾ᮶ಶయ⩌ࡀศᕸࡋ㸪⮬↛⏕⏘ࡀⰋዲ࡞Ỉᇦ࡛ࡣ㸪ᅾ᮶ಶయ⩌ ࡢಖࡢࡓࡵ✀ⱑᨺὶࢆᐇࡏࡎ㸪⚗⁺ࡍࡿ㸬ࡲࡓ㸪ᅾ᮶ಶయ⩌ࡀᏑᅾࡋ ࡞࠸㸪ࡲࡓࡣ㣴Ṫ✀ⱑ➼ࡢ㠀ᅾ᮶ಶయΰྜࡋ࡚࠸ࡿỈᇦ࡛ࡣ㐟⁺ࢆ⾜࠸㸪⮬ ↛⏕⏘ࡀⰋዲ࡛࠶ࢀࡤ㸪ᙧែࡸ⏕ែࡀࡼࡾኳ↛㨶㏆࠸ಶయࡢቑṪࢆᮇᚅࡋ 36 ࡚Ⓨ║༸ࡸ⛶㨶ࡢᨺὶࡀ᥎ዡࡉࢀࡿࡀ㸪⮬↛⏕⏘ࡀపㄪ࡞ሙᡤ࡛ࡣ⛶㨶ࡲࡓ ࡣᡂ㨶ࡢᨺὶࡀ㐺⏝ࡉࢀࡿ㸬ୖ࡛᥎ᐹࡉࢀࡓࡼ࠺㸪࣐ࢦࡢ⏘༸ᗋᾋୖ⛶ 㨶ࡢὶ⛬ศᕸࡀ㢮ఝࡋ㸪ࡲࡓሙᡤ㛫࡛Ꮴ⋡ࡸึᮇ⏕ṧ⋡ࡁ࡞㐪࠸ࡀ࡞࠸ ࡍࢀࡤ㸪ᮏㄪᰝỈᇦ࠾ࡅࡿ࣐ࢦࡢ⮬↛⏕⏘ࡣᾋୖ⛶㨶ࡢὶ⛬ศᕸ 㸦Table 3-2㸧ࡽホ౯ࡍࡿࡇࡀྍ⬟࡛࠶ࡾ㸪ୖὶഃࡢ Sec. 11–14 ࡛Ⰻዲ㸪ࡑ ࢀࡼࡾୗὶഃ࡛పㄪࡔホ౯ࡉࢀࡿ㸬ࡲࡓ㸪ୖὶഃㄪᰝ༊ᗂ㨶ᮇ௨㝆ࡢ ࣐ࢦࡢಶయᩘᐦᗘࡀ㧗ࡃ㸪ࡲࡓᖹᆒ㛗ࡀᑠࡉࡃ㸪ࡑࡢศᩓࡀࡁ࠸ഴྥࡣ㸪 ୖὶഃỈ ࡀప࠸ࡓࡵᙜṓ㨶ࡢᡂ㛗ࡀ㐜࠸ࡇࡼࡿྍ⬟ᛶࡀ࠶ࡿ㸪 ୖὶഃࡢಶయ⩌ࡀ㸪┦ᑐⓗከᩘࡢⱝ㱋㨶࠾ࡼࡧከᵝ࡞ᖺ⣭⩌ࢆྵࡴࡇࢆ♧ ࡍྍ⬟ᛶࡀ࠶ࡾ㸪ୖὶഃ࡛Ᏻᐃࡋࡓ⮬↛⏕⏘ࡀ⾜ࢃࢀ࡚࠸ࡿࡇࢆᫎࡋ࡚ ࠸ࡿࡢࡶࡋࢀ࡞࠸㸬ᮏㄪᰝᆅ࠾ࡅࡿᅾ᮶ಶయ⩌ࡢ⏕ᜥ༊㛫ࡣ᫂ࡽ࡛࡞࠸ ࡀ㸪⏕⏘ࡢὶ⛬ࣃࢱ࣮ࣥᇶ࡙࠸࡚ࢰ࣮ࢽࣥࢢ⟶⌮ࢆ⾜࠺ࡍࡿ㸪ୖὶഃ ࡢ Sec. 11–14 ࡛ࡣ⏕⏘ࡀⰋዲ⪃࠼ࡽࢀࡿࡓࡵ㸪Ⓨ║༸ᨺὶࡲࡓࡣᨺὶࢆࡋ ࡞࠸㑅ᢥࡀ㸪୍᪉㸪Sec. 11 ࡼࡾୗὶഃ࡛ࡣㄪホ౯ࡉࢀࡿࡓࡵ㸪㐟⁺ࢆ⥔ ᣢࡋࡼ࠺ࡍࢀࡤ㸪⛶㨶ࡲࡓࡣᡂ㨶ࡢᨺὶࡀ᥎ዡࡉࢀࡿ㸦୰ᮧ㸪2009㸧㸬 ⮬↛⏕⏘ࡢຓ㛗 Sec. 11 ࡼࡾୗὶഃ࡛⏕⏘ࡀపㄪ࡛࠶ࡗࡓཎᅉࡢ୍ࡘࡣ㸪ᗂ㨶ᮇ௨㝆ࡢಶయ ࡢὶ⛬ศᕸࡽุ᩿ࡋ࡚㸪࣐ࢦࡢぶ㨶ᩘࡀࡑࡶࡑࡶᑡ࡞ࡗࡓࡓࡵ⪃࠼ࡽ ࢀࡿ㸬ᮏㄪᰝᆅ࠾ࡅࡿᗂ㨶ᮇ௨㝆ࡢ࣐ࢦࡢ⏕ᜥࡣ㸪Ỉ ࠾ࡼࡧἙᕝᙧែ➼ ࡢ≀⌮⎔ቃ᮲௳ࡀࡶࡓࡽࡍ⎔ቃᐜຊつᐃࡉࢀࡿ⪃࠼ࡽࢀࡿࡀ㸪ୗὶഃ ࡶᒁᡤⓗಶయᩘᐦᗘࡢ㧗࠸ㄪᰝ༊㸦Sec. 7㸧ࡀᏑᅾࡋࡓࡇࡽ㸪ୗὶഃỈᇦ ࡢಶయᩘᐦᗘࡢపࡉࡣ㸪ࡇࢀࡽࡢ⎔ቃせᅉࡔࡅ࡛ࡣㄝ࡛᫂ࡁ࡞࠸㸬୍᪉㸪ୗὶ ഃ㸦≉ Sec. 1–6㸧࡛ࡣ㐟⁺⪅ࡀධࡋࡸࡍ࠸ࡓࡵ㸪㔮⋓ᅽࡀ㧗࠸ࡇࡀᣦ ࡉࢀ࡚࠸ࡿ㸦ྜྷ㔝ဴஓẶ㹙㐟⁺ᅋయ 33club㹛㸪⚾ಙ㸧 㸬⁺⋓ไ㝈ࡼࡾぶ㨶ᩘࢆ ቑຍࡉࡏࡿࡇࡣ㸪⮬↛⏕⏘ࡢຓ㛗ࢆᅗࡿ୍ࡘࡢ㑅ᢥ⫥࡛࠶ࡿ࠸࠼ࡼ࠺㸬 ୗὶഃࡢㄪᰝ༊ࡣ㸪ᾋୖ⛶㨶ࡢಶయᩘᐦᗘࡀᒁᡤⓗ㧗࠸Ỉᇦࡀ」ᩘᏑᅾ ࡋࡓ㸬≉ Sec. 5 ࡢୖὶഃ㸦⣙ 0.3 km㸧࡛ࡣᾋୖ⛶㨶ࡀ㧗ᐦᗘ☜ㄆࡉࢀࡓࡀ㸪 37 ࡑࡢୖ࣭ୗὶ⣙ 1 km ࡢ⠊ᅖࡣᾋୖ⛶㨶ࡀࢇぢࡽࢀ࡞ࡗࡓ㸦Fig. 3-3㸧 㸬 ᾋୖ⛶㨶ࡀ☜ㄆࡉࢀ࡞࠸Ỉᇦࡣᒾ┙ࡢἙᗋ㸦Sec. 4 ࡢୖὶഃ㸧㸪ࢺࣟ㸦Sec. 6㸧 㸪 ሖሐ┤ୗ࠾ࡅࡿ࣮࣐࣮࠾ࡼࡧ㟢┙㸦୰ᮧ㸪2011㸧ࡋࡓἙᗋ㸦Sec. 5 ࡢ ୗὶഃ㸧ࡀぢࡽࢀࡓࡀ㸪㧗ᐦᗘỈᇦࡣ♟ࡢሁ✚ࡋࡓᖹ℩ࡀぢࡽࢀࡓ㸦ᮌᮏ㸪 ಶேⓗほᐹ㸧 㸬ࡇࡢࡼ࠺࡞ሙᡤࡣ࣐ࢦࡢ⏘༸㐺ᆅ࡛ࡶ࠶ࡾ㸪⏘༸㐺ᆅࡢ㊊ࡀ ୗὶഃỈᇦయࡢ⮬↛⏕⏘ࢆไ㝈ࡋ࡚࠸ࡿྍ⬟ᛶࡀ࠶ࡿ㸬㏆ᖺ㸪࣡ࢼ࡛ࡣ ேᕤ⏘༸ሙࢆ㐀ᡂࡋ࡚⮬↛⏕⏘ࢆຓ㛗ࡍࡿヨࡳࡀ⾜ࢃࢀ࡚࠸ࡿ㸦୰ᮧ㸪1999b㸧 㸬 ࡇࡢ᪉ἲࡣ㸪≉Ἑᕝᕤస≀ࡢᘓタ➼࡛⏘༸㐺ᆅࡀῶᑡࡋࡓᑠᨭὶ࠾࠸࡚ಶ య㛫ࡢ㔜」⏘༸ࡼࡿ⏕⏘ࡢ㜼ᐖࢆᅇ㑊ࡍࡿ┠ⓗ࡛ᐇࡉࢀ࡚࠸ࡿࡀ㸪⏘༸ ሙ♟ࢆᩜࡃࡇࡼࡾ㸪Ἑᗋᮦᩱࡀ㊊ࡋࡓỈᇦ࡛ࡶ⮬↛⏕⏘ࢆຓ㛗࡛ࡁ ࡿࡶࡋࢀ࡞࠸㸬ࡓࡔࡋ㸪ศᕸ༡㝈㏆ࡢ࣐ࢦ࡛⏘༸ሙᡤᾋୖ⛶㨶ࡢ⏕ᜥ ሙᡤࡀ㏆᥋ࡍࡿྍ⬟ᛶࢆ⪃៖ࡍࡿ㸪ேᕤ⏘༸ሙࡢ㐀ᡂຍ࠼㸪30 cm ௨ୗࡢ Ỉ῝ 20 cm/s ௨ୗࡢ⾲ᒙὶ㏿㸦ྡ㉺㸪1988㸧㸪20 cm/s ௨ୗࡢᗏ㠃ὶ㏿ ከ㔞ࡢỈ୰᳜⏕ࡸಽὶᮌ㸦Nagata and Yanai㸪2002㸧࠸ࡗࡓ࣐ࢦࡸࢧࢡ࣐ࣛ ࢫࡢᾋୖ⛶㨶ࡀዲࡴ≀⌮ⓗ⎔ቃࢆྠ౪⤥ࡍࡿᚲせࡀ࠶ࡿ㸬 ᮏㄪᰝ࡛ࡣ㸪ከᩘࡢሖሐ➼࡛ศ᩿ࡉࢀࡓ Sec. 12–13 ࠾࠸࡚㸪ᾋୖ⛶㨶ࡢಶ యᩘᐦᗘࡀ㧗࠸್ࢆ♧ࡋࡓ㸬Ἑᕝᕤస≀ࡣ㸪▷ᮇⓗࡣಶయࡢ⛣ືࢆไ㝈ࡋ࡚ ୖὶഃ࠾ࡅࡿᅇ㐟ᛶ㨶㢮ࡢ⤯⁛ࢆࡶࡓࡽࡍ㸦୰㔝㸪1995㸧㸪୰࣭㛗ᮇ ⓗࡣ◁♟ࡢ⛣ືࡢไ㝈ࡼࡾ㸪ୖὶഃ࡛ࡣ◁♟ࡢሁ✚㸪ୗὶഃ࡛ࡣὶኻࡀ㐍 ⾜ࡋ㸪⏘༸ሙᡤࡸ⛶㨶ࡢ⏕ᜥሙᡤ➼ࢆࡶࡓࡽࡍἙᕝ⎔ቃࡢከᵝᛶࡀኻࢃࢀࡿࡇ ࡀ▱ࡽࢀࡿ㸦๓ᕝ㸪2000㸹୰ᮧ㸪2011㸧 㸬୍᪉㸪ᐃఫᛶࢆ♧ࡍ㨶㢮ᑐࡋ࡚ࡣ ࡇࢀࡽࡢᝏᙳ㡪ࡀẚ㍑ⓗᑠࡉ࠸㸦୰㔝㸪1995㸧 㸪ᕤస≀ୗὶഃࡢἙᗋࡢ 㟢┙ࡣᕤస≀タ⨨ᚋࡢ㛫⤒㐣ᚑࡗ࡚㐍⾜ࡍࡿࡇࡀ▱ࡽࢀ࡚࠸ࡿ 㸦୰ᮧ㸪 2011㸧 㸬ᮏㄪᰝỈᇦࡢ࣐ࢦࡣἙᕝᆺಶయ⩌࡛࠶ࡾᐃఫᛶࡀ㧗࠸⪃࠼ࡽࢀࡿࡇ 㸪ࡲࡓ Sec. 12–13 Ꮡᅾࡍࡿ 2 ࡘࡢࡁ࡞◁㜵ࢲ࣒ࡀẚ㍑ⓗ᭱㏆㸦1972 ᖺ 1995 ᖺ㸧❹ᕤࡋࡓࡇࡽ㸪ㄪᰝᐇⅬ࡛ࡣ㸪ᙜヱỈᇦࡢ࣐ࢦࡢ⮬↛⏕ ⏘ᑐࡍࡿศ᩿ࡢᙳ㡪ࡣᑡ࡞ࡗࡓࡶࡢ⪃࠼ࡽࢀࡿ㸬ࡋࡋ㸪Ἑᕝᕤస≀ ࡼࡾᑠ㞟ᅋࡉࢀࡓ㝸㞳ಶయ⩌࠾࠸࡚㸪㛗ᮇⓗࡣ㸪ಶయᩘኚືࡢ☜⋡ᛶ 38 ࡸ㑇ఏⓗຎࡼࡗ࡚⤯⁛☜⋡ࡀ㧗ࡲࡿࡇࡀᣦࡉࢀ࡚࠾ࡾ㸦᳃⏣࣭ᒣᮏ㸪 2004㸧 㸪ᐃఫᛶࡀ㧗࠸ࡉࢀࡿἙᕝᆺࡢ࣡ࢼ࣐ࢦ࠾࠸࡚ࡶ㸪◁㜵ሖሐ࡛ ศ᩿ࡉࢀࡓ㝸㞳ಶయ⩌ࡀ⤯⁛ࡍࡿࡀሗ࿌ࡉࢀ࡚࠸ࡿ㸦㐲⸨㸪2006㸧 㸬ࡲ ࡓ㸪㏆ᖺ㸪ேᕤ✀ⱑࡢᨺὶࡼࡾᅜⓗῶᑡࡋ࡚࠸ࡿࢧࢣ⛉㨶㢮ࡢᅾ᮶ಶయ ⩌ࡣ㸪ሖሐ➼ࡢୖὶഃ㝸㞳ಶయ⩌ࡢ≧ែ࡛⏕ᜥࡋ࡚࠸ࡿሙྜࡀከࡃ㸦୰ᮧ㸪 2009㸧㸪ྠᵝ⤯⁛ࡢྍ⬟ᛶࡀ༴ࡉࢀ࡚࠸ࡿ㸦Kawamura et al., 2007㸹Kubota et al., 2007㸧 㸬ᅾ᮶ಶయ⩌ࡢ⤯⁛ࢆᅇ㑊ࡍࡿࡓࡵࡣ㸪ࡑࡢ㑇ఏⓗከᵝᛶ⊂⮬ᛶ ࡀྠ☜ಖࡉࢀࡿᚲせࡀ࠶ࡿࡇࡽ㸪ᅾ᮶ಶయ⩌㛫࠾ࡅࡿಶయࡢ⛣᳜ࡸ 㠀ᅾ᮶ಶయࡢ㝖ཤ➼㸪㑇ఏᏛⓗሗᇶ࡙࠸ࡓಶయ⩌⟶⌮ࢆᐇࡍࡿࡇࡀᥦ ࡉࢀ࡚࠸ࡿ㸦Kawamura et al., 2007㸹Sato et al., 2010㸧 㸬ᮏㄪᰝỈᇦ࠾࠸࡚ࡶ㸪 ሖሐ➼࡛ศ᩿ࡉࢀࡓ㝸㞳ಶయ⩌ࡀᏑᅾࡍࡿࡇ㸪≉᭱ୖὶ㒊ࡢ Sec. 14 ࡣබ ⓗ࡞ᨺὶグ㘓ࡀ࡞ࡃ㸪ᅾ᮶ಶయ⩌ࡀ⏕ᜥࡍࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡿࡇࡽ㸦 㔝ᕝ⁺ᴗ༠ྠ⤌ྜࡽࡢሗ㸧 㸪ᚋࡣ㸪ᅾ᮶ಶయ⩌ࡢ᥈⣴ࢆྵࡴ㑇ఏⓗᵓ㐀ࡢ ⌧≧ᢕᥱࡸࣔࢽࢱࣜࣥࢢࢆ⾜࠸㸪㑇ఏᏛⓗሗࢆ✚ࡍࡿᚲせࡀ࠶ࡿ㸬 ᚋࡢㄢ㢟 ᮏ◊✲࡛ࡣ㸪ᾋୖ┤ᚋࡢ࣐ࢦ⛶㨶ࡀ⏘༸ᗋ㏆␃ࡲࡿྍ⬟ᛶࢆ♧၀ࡋࡓ ࡀ㸪ᩘ✀ࡢࢧࢣ⛉㨶㢮࡛ࡣ㸪ᾋୖ⛶㨶ࡢ୍㒊ࡀᵝࠎ࡞⛬ᗘ㸦0.1–2.0 km㸧࡛ᐦ ᗘ౫Ꮡⓗὶୗࡍࡿࡇࡀ▱ࡽࢀ㸦Nagata, 2002㸹Foldvik et al., 2010㸧㸪⛣ື ࡼࡿ⿕㣗ࣜࢫࢡ⛣ືඛ࡛ᚓࡽࢀࡿ㧗ᡂ㛗ࡣࢺ࣮ࣞࢻ࢜ࣇࡢ㛵ಀ࠶ࡿ࠸࠺ 㸦Einum et al., 2011, 2012㸧 㸬ᮏ◊✲࡛ࡣ㸪Ἑᕝᆺ࣐ࢦಶయ⩌ࡢ⏕ᜥሙᡤయࢆ ᑐ㇟ὶᇦࢫࢣ࣮ࣝࡢㄪᰝࢆ⾜࠸㸪ྛㄪᰝ༊ࡢ㛗ࡉࢆ⣙ 1 km ࡋࡓࡓࡵ㸪ࡑ ࢀ௨ୗࡢࢫࢣ࣮ࣝ࠾ࡅࡿᾋୖ⛶㨶ࡢ⛣ືࢆ᳨ฟࡋ࡚࠸࡞࠸㸬ᾋୖ⛶㨶ᮇᐦ ᗘ౫Ꮡⓗ࡞⛣ືࡀ⏕ࡌࡿࡍࢀࡤ㸪⎔ቃᐜຊẚࡋ࡚㐣࡞Ⓨ║༸ࡸ⛶㨶ࡀ ᨺὶࡉࢀࡓሙྜࡣ㸪ࡑࢀࡽࡢᨺὶຠᯝࡀపୗࡍࡿࡔࡅ࡛ࡣ࡞ࡃ㸪ኳ↛ࡢᾋୖ ⛶㨶ࡢ⏕ṧࡶᝏᙳ㡪ࢆ࠼ࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡿ㸬ᚋࡣ㸪ࡼࡾ⣽࠸ࢫࢣ ࣮ࣝࡢㄪᰝࢆ⾜࠸㸪࣐ࢦᾋୖ⛶㨶࠾ࡅࡿࡼࡾヲ⣽࡞⛣ືᵝᘧࢆᢕᥱࡍࡿᚲ せࡀ࠶ࡿ㸬 39 Û( Û( Sec. 2 N Sec. 1 Sec. 3 Sec. 4 Sec. 5 Sea of Japan Sec. 6 Ogata River Japan Tosumi River Sec. 7 Û1 Pacific Ocean Û(ޗ Sec. 8 Sec. 9 Û(ޗ Ûޗ1 Fukuoka Prefecture Sec. 10 Kohbaru Stream waterfalls man-made dam spring Sec. 11 Oita Prefecture Seto Inland Sea Sec. 12 Ichigome Falls Hakiai Stream Sec. 13 Ono River Sec. 14 Kumamoto Prefecture Gogome Falls Ûޗ1 Mt. Sobo Mennotsura-dani Stream 1 km Fig. 3-1. Sections established in the Ogata River and its tributaries (Kohbaru and Hakiai streams), northeastern Kyushu Island, Japan. 40 Section 1 2 3 4 5 6 7 8 9 10 11 12 13 14 䕰 䕰 䕰 䕿 䕿 䕧 䕧 䕿 䕿 䕰 䕰 䕧 䕧 䕿 䕿 䕰 䕧 䕧 䕰 䕿 䕰 䕰 䕰 䕰 䕰 䕧 䕿 䕧䕧 䕿䕿 䕧 䕿 䕿 䕿 䕿䕿 䕰 䕧 䕧䕧 䕧 䕧 䕿 䕧 䕿䕰 1 Jan. 15 Jan. 1 Feb. 15 Feb. 1 Mar. 15 Mar. 1 Apr. 䕧 15 Apr. 2005 Fig. 3-2. Survey schedule to clarify the longitudinal distribution of newly emerged fry of amago in 14 sections in the Ogata River and its tributary, the Kohbaru Stream, from January to April in 2005. Closed inverted triangles: estimated dates when cumulative water temperature reached 800°C from November 1, 2004 (amago fry have been reported to emerge from the spawning redd at 800°C). Open circles: first survey dates; open triangles: second survey dates; two surveys were conducted for each section at intervals of 11–32 days. 41 Section 1 25 2 3 4 5 6 7 8 9 10 11 12 13 14 35 First survey 䡚 䡚 20 Number of newly emerged fry 15 10 5 0 0 2 1 25 2 3 4 4 6 5 6 8 7 10 8 9 12 10 11 14 12 13 16 14 44 Second survey 䡚 䡚 20 15 10 5 0 0 2 4 6 8 10 12 14 16 Distance from base point (km) Fig. 3-3. Longitudinal distribution of newly emerged fry of amago observed in a 16.4 km stretch of the Ogata River and its tributary, the Kohbaru Stream, in the first (top) and second (bottom) surveys, conducted from 7 January to 28 March 2005 and from 21 January to 17 April 2005, respectively (see Fig. 2). Amago fry counted by snorkelers moving upstream (Sections 1–14). Arrows in Sec. 6 and 14 indicate locations of water falls of height >10 m, and others, man-made dams of height > 1 m. 42 Density of post-fry amago (N/100 m2) 18 16 14 12 10 8 6 4 2 0 October 2003 February 2004 August 2004 1 2 3 4 5 6 7 8 9 10 11 12 13 14 Section Fig. 3-4. Longitudinal changes in density of post-fry amago in 14 sections established in the Ogata River and its tributary, the Kohbaru Stream, in October 2003, February 2004 and August 2004. Amago counted by snorkelers for ca. 100 m in each section. 43 Density of newly emerged fry of amago (N/100 m2) 5.0 4.0 3.0 2.0 1.0 0.0 0 5 10 15 Density of post-fry amago (N/100 20 m 2) Fig. 3-5. Relationship between density of newly emerged fry (surveyed from 7 January to 28 March, 2005) and that of post-fry amago (density assessed in August 2004) in 14 sections established in the Ogata River and its triburary, the Kohbaru Stream. 44 45 Max. Min. Difference **** DMWT: Daily mean water temperature. *** At the upper end of each section. ** Post-fry amago salmon counted in these areas. * Topographic types follow Kani (1978). Discharge (m /s) DMWT (°C)**** 3 Survey area (m )** Altitude (m)*** Gradient (%) 2 Topographic type* Section length (km) Mean wetted width (m) Section area (ha) 1 2 Bb Bb 1.01 1.29 21.2 20.6 2.14 2.67 2,263 2,567 232 238 255 0.6 1.3 2.81 2.13 20.3 20.6 20.8 9.8 9.9 9.4 10.5 10.7 11.4 Base Point Section 3 4 5 6 7 8 9 10 11 12 13 Bb Bb Aa-Bb Aa-Bb Aa-Bb Aa-Bb Aa-Bb Aa-Bb Aa Aa Aa 1.00 1.45 1.19 1.04 1.80 1.20 1.46 0.78 0.73 1.31 1.10 26.0 11.8 14.6 10.4 15.2 13.4 12.2 16.7 8.9 10.3 9.1 2.60 1.71 1.73 1.08 2.72 1.61 1.77 1.30 0.65 1.35 1.00 1,573 650 999 688 929 751 883 1,090 745 473 479 258 273 284 296 325 342 384 397 416 486 604 0.3 1 0.9 1.1 1.6 1.4 2.9 1.7 2.6 5.3 10.8 2.33 1.95 1.04 0.75 0.32 0.35 0.19 0.13 0.05 0.06 0.08 20.5 22.9 21.5 23.6 21.6 21.4 22.2 22.0 20.7 18.8 19.1 9.6 6.6 7.0 5.4 5.6 5.5 3.9 3.8 5.4 4.5 2.6 10.9 16.2 14.5 18.1 16.0 16.0 18.2 18.2 15.3 14.4 16.5 Japan, established for the investigation of longitudinal distribution of amago. 14 Aa 1.02 7.1 0.73 411 820 21.1 0.02 18.2 0.2 18.1 Table 3-1. Environmental data for 14 sections of the Ogata River and its tributary, the Kohbaru Stream, in northeastern Kyushu Island, 46 (per 100 m2 ) Number of Individuals Density 1 2 3 4 5 first 0 4 9 7 54 second 1 31 35 7 48 first 0 0.01 0.03 0.04 0.31 second 0.005 0.12 0.13 0.04 0.28 Survey second (21 January to 17 April) surveys in 2005. 6 0 0 0 0 Section 7 8 9 10 11 12 13 48 52 40 67 179 428 152 68 67 70 42 64 311 390 0.18 0.32 0.23 0.51 2.77 3.18 1.53 0.25 0.42 0.39 0.32 0.99 2.31 3.92 Total 14 334 1,374 342 1,476 4.59 0.60 4.70 0.64 surveyed) in the Ogata River and its tributary, the Kohbaru Stream, in the first (7 January to 28 March) and Table 3-2. Number of individuals and densities of newly emerged fry of amago observed in 14 sections (16.4 km 47 21–29 October 2003 5.0–9.9 10.0–14.9 15.0–19.9 20.0–24.9 25.0–29.9 Subtotal Mean Variance Total length (cm) 0 1 3 17.5 0 3 2 0 3 4 17.5 0 4 4 5 3 5 2 1 11 18 22.3 its tributary, the Kohbaru Stream, in October 2003. 4 17.5 0 4 6 23 16.2 7.3 7 15 1 11 16.6 9.1 3 7 1 Section 7 8 9 12 6.3 1 7 1 9 18 13.3 3.7 15 3 10 17 12.8 14 3 11 2 1 11 36 12.9 23.4 13 9 12 2 12 24 14.8 23.9 3 12 4 5 13 6 11 10 3 5 35 16.1 40.5 14 26 78 70 15 6 195 Total Table 3-3-1. Total length distribution of amago observed in 14 sections (ca. 100 m surveyed in each) in the Ogata River and 48 2–10 February 2004 5.0–9.9 10.0–14.9 15.0–19.9 20.0–24.9 25.0–29.9 Subtotal Mean Variance Total length (cm) 0 1 0 2 0 3 0 4 its tributary, the Kohbaru Stream, in February 2004. 0 5 0 6 0 0 17.5 0 Section 7 8 2 17.5 0 2 9 1 17.5 0 1 10 6 17 12.3 6 11 21 17.9 1.9 6 11 4 12 13 19.2 6.3 12 1 13 9 17.7 6.8 6 3 14 0 6 38 8 0 52 Total Table 3-3-2. Total length distribution of amago observed in 14 sections (ca. 100 m surveyed in each) in the Ogata River and 49 0 0 1 3 0 2 13 17.5 8.3 13 2 9 2 3 6 15.8 6.7 10 2 4 4 2 15 12.5 13 1 1 5 8 15 14.3 12 1 2 5 6 38 13.3 14.2 61 7 19 11 1 17 12.5 9.4 28 3 11 3 Section 7 8 10 11 11.4 21 4 5 1 9 14 10 10.6 33 8 5 1 10 46 8.9 6.3 69 34 11 1 11 15 31 15 6 1 68 13.6 22.3 125 12 57 11.3 19.9 94 28 18 8 3 13 46 11 6 4 1 68 10.4 24.1 112 14 594 146 118 65 16 2 347 Total 2004. *Survey of Section 14 conducted on 6 October 2004 because of water discoloration from late August to late September 6–24 August 2004* 5.0–9.9 10.0–14.9 15.0–19.9 20.0–24.9 25.0–29.9 Subtotal Mean Variance Total Total length (cm) its tributary, the Kohbaru Stream, in August 2004. Table 3-3-3. Total length distribution of amago observed in 14 sections (ca. 100 m surveyed in each) in the Ogata River and ➨ 4 ❶ ࣐ࢦᾋୖ⛶㨶ࡢ⏕ᜥሙᡤ⏝ ㏆ᖺ㸪Ἑᕝᛶࢧࢣ⛉㨶㢮ࡢಖ⏝ࡢ୧❧ࢆᅗࡿࡓࡵ㸪ࢰ࣮ࢽࣥࢢࡼ ࡿ⁺ሙ⟶⌮ࡀ᪥ᮏྛᆅ࡛ヨࡳࡽࢀ࡚࠸ࡿ㸦୰ᮧ㸪2009㸧㸬ࡇࡢᡭἲ࡛ࡣ㸪ᨺὶࡉ ࢀࡓ㣴Ṫ㨶ࡢ㞧ࢆචࢀ࡚⏕ᜥᇦࡢ᭱ୖὶ㒊⏕ᜥࡍࡿᅾ᮶ಶయ⩌ࢆ᥈⣴ࡋ㸪 ⚗⁺➼ࡼࡗ࡚✚ᴟⓗಖࡍࡿ㸦୰ᮧ㸪2001㸹Kubota et al., 2007㸹Kimoto et al., 2015㸧㸪ᅾ᮶ಶయ⩌⏕ᜥỈᇦ࡛ࡣཎ๎ࡋ࡚ᨺὶࡀ⚗Ṇࡉࢀࡿ㸬୍᪉㸪ᨺ ὶࡉࢀࡓ㣴Ṫ㨶ࡢ㞧ࡀぢࡽࢀࡿ⏕ᜥᇦࡢୗὶഃ࡛ࡣ⁺⋓㐟⁺ࡀྍ⬟࡛࠶ ࡾ㸪ᨺὶࡼࡿቑṪࡀᐜㄆࡉࢀࡿࡀ㸪Ἑᕝ⎔ቃࡸᑐ㇟㨶✀ࡢ≧ἣ࠾ࡼࡧᆅඖఫ Ẹࡸ㐟⁺⪅ࡢពྥ➼ࡼࡾ㸪↓ᨺὶࡽᡂ㨶ᨺὶࡲ࡛ᵝࠎ࡞㑅ᢥ⫥ࡀ࠶ࡿ㸦୰ ᮧ㸪2009㸧 㸬≉⮬↛⏕⏘ࡀྍ⬟࡞ሙᡤ࡛ࡣ㸪↓ᨺὶ㸪ேᕤ⏘༸ሙࡢ㐀ᡂ㸦୰ ᮧ㸪1999㸧 㸪Ⓨ║༸ᨺὶ㸦Ṋᒃ㸪2009㸧 㸪ぶ㨶ᨺὶ㸦ᚫཎ㸪2010㸧➼ࡢᡭἲ ࡀ⪃࠼ࡽࢀ㸪ࡇࢀࡽࡣᾋୖᚋࡢ⛶㨶ࡢ⏕⫱ࢆ⮬↛⎔ቃࡺࡔࡡࡿⅬ࡛ඹ㏻ࡋ࡚ ࠸ࡿ㸬ࡓࡔࡋ㸪※ὶ㒊ࡼࡾୗὶഃࡢᓙ㇂ࡽᡪ≧ᆅࡅ࡚ࡣ㸪ୖὶഃᘓタ ࡉࢀࡓ◁㜵ࢲ࣒ࡼࡾᅵ◁౪⤥ࡀไ㝈ࡉࢀ㸪ㆤᓊࡸᗋᅛᕤ➼ࡼࡿἙᕝᨵಟࡀ 㐍ࡳ㸪⮬↛⎔ቃࡀᝏࡋ࡚࠸ࡿࡇࡀ▱ࡽࢀ࡚࠸ࡿ㸦㧗ᶫ㸪2000㸹Yoshimura et al., 2005㸧㸬ࡲࡓ୍⯡ⓗ㸪ୗὶഃࡣୖὶഃࡼࡾࡶ᭱㧗Ỉ ࡀ㧗ࡃ㸦Vannote et al., 1980㸧㸪㏆ᖺࡢ ᬮഴྥࢆ⪃៖ࡍࡿ㸦㇂ཱྀ࣭୰㔝㸪2000㸧 㸪Ἑᕝᛶࢧࢣ⛉㨶 㢮ࡢ⏕ᜥᇦୗὶഃ࠾ࡅࡿಖㆤࡸቑṪࡣ㸪ୖὶ㒊ࡼࡾࡶᅔ㞴࡛ษ㏕ࡋࡓၥ㢟 ࠸࠼ࡿ㸦Aarts and Nienhuis, 2003㸧㸬ࡇࢀࡽࡢቑṪᡭἲࢆຠᯝⓗᶵ⬟ࡉࡏࡿࡓ ࡵࡣ㸪ᑐ㇟㨶✀ࡢୗὶഃỈᇦ࠾࠸࡚㸪ᾋୖ⛶㨶ࡢ⏕ᜥሙᡤ⏝㛵ࡍࡿ▱ ぢࢆ㞟ࡍࡿࡇࡀᚲせ࡛࠶ࡿ㸦Kramer et al., 1997㸧 㸬 ࢧࢣ⛉㨶㢮ࡢᾋୖ⛶㨶ࡢ⏕ᜥሙᡤ⏝ࡘ࠸࡚㸪ᅜෆ࡛ࡣ࣐ࢦ Oncorhynchus masou ishikawae㸦ྡ㉺㸪1988㸧㸪ࢧࢡ࣐ࣛࢫ Oncorhynchus masou masou㸦Nagata and Yanai, 2002㸧㛵ࡍࡿሗ࿌ࡀ࠶ࡾ㸪ࡶὸࡃὶ㏿ࡀᑠࡉ࠸ ᓊἢ࠸ࡢሙᡤࢆ⏝ࡍࡿࡇࡀ▱ࡽࢀ࡚࠸ࡿ㸬ࡓࡔࡋ㸪ࡇࢀࡽࡢ◊✲ࡣἙᕝࡢ ୖὶᇦ࡛⾜ࢃࢀ࡚࠾ࡾ㸪⏕ᜥሙᡤࡢ≉ᚩࡣ㸪ᚤ⏕ᜥሙᡤࢫࢣ࣮ࣝ㸦Frissell et al., 1986㸧࠾ࡅࡿᾋୖ⛶㨶ࡢᐃⅬࡢ⎔ቃࡋ࡚⾲ࡉࢀ࡚࠸ࡿ㸬࠶ࡿሙᡤࡸ✵㛫 50 ࢫࢣ᳨࣮࡛ࣝฟࡉࢀࡓ㨶㢮⎔ቃせᅉࡢ㛵ಀࢆ㸪ูࡢሙᡤࡸ✵㛫ࢫࢣ࣮ࣝᣑ ᙇࡍࡿࡇࡣ㐺ᙜ࡞ሙྜࡀ࠶ࡿ㸦Fausch et al., 1988㸹୰ᮧ㸪1989㸧㸬ࡲࡓ㸪ᚤ ⏕ᜥሙᡤࢫࢣ࣮ࣝࡢ◊✲࡛ࡣ㸪㨶㢮ࡢᐃⅬ࠾ࡅࡿỈ῝ࡸὶ㏿ࡀ᫂ࡽ࡞ ࡿࡀ㸦Bozek and Rahel, 1991㸹Inoue et al., 1997㸧㸪ࡑ࠺ࡋࡓᚤ⏕ᜥሙᡤࢆἙᕝෆ ࡛࠺㓄⨨ࡍࡁࡘ࠸࡚ࡣ㸪ࡼࡾୖࡢࢫࢣ࣮ࣝࡢሗࡀᚲせ࡛࠶ࡿ 㸦Inoue and Nakano, 1999㸧㸬➨ 3 ❶㸦ᮌᮏ㸪2013㸧࡛ࡣ㸪⚄ཎᕝ⥴᪉ᕝ࡛ ࣐ࢦᾋୖ⛶㨶ࡢὶ⛬ศᕸࢆㄪ㸪⏕ᜥᇦࡢୗὶഃ࡛㸪ᾋୖ⛶㨶ࡀᩘⓒ࣓࣮ࢺ ࣝつᶍ࡛㐃⥆ศᕸࡍࡿࡇࢆ♧ࡋࡓ㸬ࡇࢀࡣ㸪⏕ᜥᇦୗὶഃ࠾ࡅࡿᾋୖ ⛶㨶ࡢศᕸỴᐃ㸪ᚤ⏕ᜥሙᡤࢫࢣ࣮ࣝࡣ␗࡞ࡿ㸪ࡁ࡞✵㛫ࢫࢣ࣮ࣝࡢ⎔ ቃせᅉࡀ㛵ࡍࡿࡇࢆ♧၀ࡍࡿ㸬 ᮏ◊✲ࡣ㸪㔝ᕝỈ⣔⥴᪉ᕝࡢ࣐ࢦ⏕ᜥᇦୗὶഃ࠾࠸࡚㸪ὶ㊰༢ࢫࢣ ࣮ࣝ࠾ࡅࡿᾋୖ⛶㨶ࡢ⏕ᜥሙᡤ⏝ࢆ᫂ࡽࡍࡿࡇࢆ┠ⓗࡋࡓ㸬ࡑࡢ ࡓࡵ㸪₯Ỉ┠どほᐹࡼࡾ࣐ࢦᾋୖ⛶㨶ࡢศᕸࢆㄪ㸪⎔ቃኚ㔞ࡢ㛵ಀ ࢆ୍⯡⥺ᙧΰྜࣔࢹࣝ㸦GLMM㸧ࡼࡾゎᯒࡋࡓ㸬ࡓࡔࡋ㸪Ἑᕝᛶࢧࢣ⛉㨶 㢮ࡢຍධ㔞ࡣᖺኚືࡀࡁࡃ㸦Nakano and Nagoshi, 1985㸹ஂಖ⏣㸪2001㸧 㸪 ࢧࢡ࣐ࣛࢫࢱࢭࣚ࢘ࢧࢣ Salmo salar ࡢᾋୖ⛶㨶࡛ࡣᐦᗘ౫Ꮡⓗ࡞⛣ື ࡀሗ࿌ࡉࢀ࡚࠸ࡿࡇࡽ㸦Nagata, 2002㸹Einum et al., 2011, 2012㸧 㸪ຍධ㔞 ࡼࡗ࡚ࡣ⛶㨶ࡀ⛣ືࡍࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡿ㸬⛣ື୰ࡢ⛶㨶ࡣ㸪ᮏ᮶ࡣ⏝ࡋ ࡞࠸㐺ᙜ࡞ሙᡤᐃࡍࡿྍ⬟ᛶࡀ࠶ࡿࡓࡵ㸪1 ᅇࡢほᐹᇶ࡙ࡃ᥎ᐃࡣㄗ ࡗࡓ⤖ㄽࢆ⏕ࡴ༴㝤ᛶࡀ⪃࠼ࡽࢀࡿ㸬ࡑࡇ࡛㸪ᮏ◊✲࡛ࡣ」ᩘᖺ㸪」ᩘᅇࢃ ࡓࡗ࡚ㄪᰝࢆ⾜࠸㸪ᖺࡈᾋୖ⛶㨶ಶయᩘ┦㛵ࡍࡿ⎔ቃせᅉࢆ᥎ᐃࡋࡓࡢ ࡕ㸪యඹ㏻ࡍࡿせᅉࢆ⪃ᐹࡋࡓ㸬 ᮦᩱ᪉ἲ ㄪᰝሙᡤㄪᰝ༊⏬ 㔝ᕝỈ⣔⥴᪉ᕝࡣ㸪ศ┴➉⏣ᕷࡢ㉺ᩜᓅ㸦ᶆ㧗 1,060 m㸧ࡽⓎࡋ㸪⣙ 12 km ᮾὶࡋ࡚♽ẕᒣ㸦1,756 m㸧Ỉ※ࢆᣢࡘᨭὶ⚄ཎᕝྜὶࡋࡓࡢࡕ㸪⣙ 51 25 km ᮾ᪉ྥὶୗࡋ㸪Ἑཱྀࡽ⣙ 52 km ᆅⅬࡢ㔝ᕝὶධࡍࡿ㸬ᮏ◊✲ ࡛ࡣ㸪⚄ཎᕝࡢྜὶⅬࡽ⣙ 1,800 m ୗὶࡢྲྀỈሖሐ㸦ⴠᕪ⣙ 1.4 m㸪㨶㐨࡞ ࡋ㸧ࢆᇶⅬࡋ㸪⣙ 1,400 m ୖὶࡢ⮬↛㸦ⴠᕪ⣙ 11 m㸧ࡲ࡛ࢆㄪᰝỈᇦࡋ ࡓ㸦Fig. 4-1㸧㸬Ἑᕝᙧែᆺࡣ Aa-Bb ⛣⾜ᆺ㸦ྍඣ㸪1978㸧㸪ᶆ㧗ࡣ 280–290 m㸪 Ἑᗋ໙㓄ࡣ 0.7%㸦1/140㸧࡛࠶ࡾ㸪࣐ࢦ௨እࢱ࢝ࣁࣖ Phoxinus oxycephalus㸪 ࢘ࢢ Tribolodon hakonensis㸪࣒࢝࣡ࢶ Candidia temminckii ࡀከࡃ⏕ᜥࡍࡿ㸦ᮌ ᮏ㸪2015㸧 㸬ㄪᰝᮇ㛫୰㸪ᮏㄪᰝỈᇦ࡛࣐ࢦ⛶㨶ࡣᨺὶࡉࢀ࡞ࡗࡓ㸬2007 ᖺ 1 ᭶㸪ㄪᰝỈᇦࡢἢᓊᇶⅬࡽ 25 m ࠾ࡁࣛࢵ࣮࢝ࢫࣉ࣮࡛ࣞ␒ྕࢆ グࡋ࡚ᶆⅬࡋ㸪ᶆⅬ࡛༊ษࡽࢀࡓ 25 m ࡢὶ㊰ࢆ 1 ༊⏬㸦section㸧ࡋ࡚ 56 ಶࡢ༊⏬㸦ྜィ 1,400 m㸧ࢆタࡅࡓ㸬ྛ༊⏬ࡣ㸪ᇶⅬࡽୖὶ᪉ྥ␒ྕࢆ ࡋ࡚༊⏬ 1 ࡽ༊⏬ 56 ࡋࡓ㸬1 ༊⏬ࡢ㛗ࡉࢆ 25 m ࡋࡓ⌮⏤ࡣ㸪୍⯡ⓗ ྛὶ㊰༢㸦℩㸪ῡ➼㸧ࡢ㛗ࡉࡀᖹᆒỈ㠃ᖜ௨ୖ࡛࠶ࡾ㸦Bisson et al., 2006㸧 㸪 ᅜෆࡢἙᕝ୰ὶᇦ࡛ࡣ⣙ 1.5–3.5 ಸ࡛࠶ࡿࡇ㸦ⴛሙ㸪2003a㸧 㸪࠾ࡼࡧᮏㄪ ᰝỈᇦࡢᖹᆒỈ㠃ᖜࡀ 12.7 m ࡛࠶ࡗࡓࡇࡼࡿ㸦Table 1㸧 㸬ᮏㄪᰝỈᇦࡢᬒ ほࡣ༊⏬ 37 ㏆ࢆቃୖ࣭ୗὶഃ࡛␗࡞ࡾ㸪ୖὶഃ࡛ࡣ୧ᓊࡀษࡾ❧ࡕỈ῝ࡀ ࡁ࠸Ἑ㐨⊃✽㒊㸦ࢦࣝࢪࣗ㸧ࡢᬒほࢆ♧ࡋࡓ㸦Fig. 1㸧 㸬 ⎔ቃኚ㔞ࡢ ᐃ 2007 ᖺ 4 ᭶ྛᶆⅬ࡛Ỉ㠃ᖜ㸪᭱Ỉ῝㸪ᕥྑᓊἢ࠸ࡢỈ῝㸦௨ୗ㸪ἢᓊỈ ῝㸧ὶ㏿ࢆ ᐃࡋࡓ㸬᭱Ỉ῝ࡣ㸪ὶ㊰ࢆᶓษࡾ࡞ࡀࡽ⟽ᑻࡲࡓࡣ㗽ࡁࣟ ࣮ࣉ࡛ ᐃࡋࡓ㸬ἢᓊỈ῝ὶ㏿ࡣ㸪࣐ࢦࡸࢧࢡ࣐ࣛࢫࡢᾋୖ⛶㨶ࡀᓊἢ࠸ ࡢሙᡤࢆዲࡴ㸦ྡ㉺㸪1988㸹Nagata and Yanai, 2002㸧ࡇࡽ㸪Ỉ㝿ࡽ ⣙ 40 cm ࡢ⨨࡛ ᐃࡋࡓ㸬࡞࠾㸪῝࠸ሙᡤ࡛ࡣᦠᖏࣉࣟ࣌ࣛᘧὶ㏿ィ㸦3631 ᆺ㸪ᶓἙ㟁ᶵ㸧ࡼࡿ ᐃࡀྍ⬟࡛࠶ࡗࡓࡓࡵ㸪࣐ࢦ࠾ࡼࡧࢧࢡ࣐ࣛࢫࡢ ᾋୖ⛶㨶࡛ሗ࿌ࡉࢀ࡚࠸ࡿᐃỈ῝㸦⣙ 20 cm㸹ྡ㉺㸪 1988㸹Nagata and Yanai, 2002㸹ⱝᯘ㸪2003㸧ࢆཧ⪃㸪40 cm ᮍ‶ࡢỈ῝࡛ࡣ 60%Ỉ῝㸪40 cm ௨ ୖ࡛ࡣỈ㠃ࡽ 24 cm ࡢὶ㏿ࢆ ᐃࡋࡓ㸬ࡲࡓ㸪Ỉ㠃ࡢⓑἼ❧ࡕᇶ࡙࠸࡚㸦ྍ ඣ㸪1978㸹Bisson et al., 2006㸧㸪ㄪᰝỈᇦෆ 14 ಶࡢ᪩℩㸦riffle㸧ࢆ≉ᐃࡋ㸦Fig. 52 1㸧㸪ὶ㊰㛗୰ኸ㒊ࡢ᭱Ỉ῝ࢆ ᐃࡋࡓ㸬ࡇࢀࡽࡣㄪᰝᮇ㛫ࢆ㏻ࡌ࡚ྠࡌ ⨨Ꮡᅾࡋࡓࡀ㸪2008 ᖺ⛅༊⏬ 50 ࡢୖὶ➃㸦1,250 m ᶆⅬ㸧᪂ࡋ࠸᪩℩ࡀ 1 ࡘ⌧ࢀࡓࡓࡵ㸦Fig. 1㸧 㸪㏆ࡢ⎔ቃኚ㔞ࢆ ᐃࡋࡓ㸬ྛ༊⏬ࡢୖ࣭ୗὶഃᶆ Ⅼࡢ⎔ቃኚ㔞ࢆᖹᆒࡋ㸪༊⏬ࡢỈ㠃ᖜ㸪᭱Ỉ῝㸪ᕥྑᓊูࡢἢᓊỈ῝㸪ὶ㏿ ࡋࡓ㸬᪩℩ࢆྵࡴ༊⏬࡛ࡣ᭱Ỉ῝ࡢ⟬ฟ᪩℩୰ኸ㒊ࡢỈ῝ࢆྵࡵࡓ㸬ࡲ ࡓ㸪Ỉ୰࠾ࡼࡧỈ㠃ࡽ⣙ 20 cm ௨ෆࡢ㧗ࡉ࠶ࡿỈ㝿ࡢ㝣⏕࣭Ỉ⏕᳜≀᳜⏕ ࡢὶ㊰᪉ྥࡢ㛗ࡉࢆ༊⏬ࡈ ᐃࡋ㸪ὶ㊰㛗࡛㝖ࡋ࡚ᕥྑᓊูࡢ᳜≀⿕ᗘ ࡋࡓ㸬 ᳜≀యࡀ 1–ᩘᮏࡢሙྜࡢ᳜≀⿕ᗘࡣ 1%ࡋࡓ㸬ྛ⎔ቃኚ㔞ࡢᩘ್ࢆ Table 4-1 ♧ࡋࡓ㸬 ㄪᰝ༊⏬ࡢศ㢮 ༊⏬ࡢタᐃ㸪Ỉ㠃ࡢἼ❧ࡕ㸪ὶ㏿㸪Ỉ῝ᇶ࡙࠸࡚㸦ྍඣ㸪1978㸹ⴛሙ 㸪2002㸪2003a㸹Bisson et al., 1982, 2006㸧 㸪ㄪᰝỈᇦෆ௨ୗࡢ 3 ✀㢮ࡢὶ ㊰༢ࢆ≉ᐃࡋࡓ㸬᪩℩㸦riffle㸧㸸ⓑἼࡀ❧ࡕὶ㏿ࡀࡁࡃỈ῝ࡀᑠࡉ࠸㸹ῡ 㸦pool㸧㸸Ỉ㠃ࡀἼ❧ࡓࡎὶ㏿ࡀᑠࡉࡃỈ῝ࡀࡁ࠸㸹ᖹ℩㸦glide㸧㸸ῡࡽ᪩ ℩ࡢ⛣⾜㒊ศ࡛࠶ࡾ㸪Ἴ❧ࡕࡀᑠࡉࡃỈ῝ࡀᑠࡉ࠸㸬ࡓࡔࡋ㸪ᮏ◊✲࡛ࡣὶ ㊰༢ࡢቃ⏺↓㛵ಀᶆⅬࢆタᐃࡋࡓࡓࡵ㸪ྛ༊⏬ࡀ」ᩘࡢὶ㊰༢ࢆྵࡴ ሙྜࡀ࠶ࡗࡓ㸬ࡲࡓ㸪ὶ㊰༢㛫᫂☜࡞ቃ⏺ࢆᐃࡵࡿࡇࡣᮏ㉁ⓗᅔ㞴࡛ ࠶ࡿ㸦ྍඣ㸪1978㸹ⴛሙ㸪2002㸹Bisson et al., 2006㸧㸬ࡑࡇ࡛㸪௨ୗࡢᇶ‽ ࡼࡾྛ༊⏬ࢆᖹ℩㸦G: glide㸧㸪ᖹ℩–᪩℩㸦GR: glide-riffle㸧㸪᪩℩㸦R: riffle㸧 㸪 ᪩℩–ῡ㸦RP: riffle-pool㸧 㸪ῡ㸦P: pool㸧㸪ῡ–ᖹ℩㸦PG: pool-glide㸧ࡢ 6 ✀㢮 ศࡅࡓ㸬ࡲࡎ㸪᪩℩ࢆྵࡴ༊⏬ࡘ࠸࡚㸪᪩℩ࡀ༊⏬యࡲࡓࡣ୰ኸ࠶ࡿሙ ྜࢆ᪩℩㸦R㸧㸪ୗὶഃ೫ࡿሙྜࢆᖹ℩–᪩℩㸦GR㸧㸪ୖὶഃ೫ࡿሙྜࢆ᪩ ℩–ῡ㸦RP㸧ࡋࡓ㸬ḟ㸪ୖグࡢ࠺ࡕ 2 ✀㢮ࡢ༊⏬㸦R, GR㸧ࡢ࠸ࡎࢀୗ ὶഃ࡛᥋ࡍࡿ༊⏬ࢆ᪩℩–ῡ㸦RP㸧ࡋ㸪ୖグ 3 ✀㢮ࡢ༊⏬㸦GR, R, RP㸧ࡢ࠸ ࡎࢀୖὶഃ࡛᥋ࡍࡿ༊⏬ࡢ࠺ࡕ㸪Ỉ㠃ࡀἼ❧ࡓ࡞࠸༊⏬ࢆῡ㸦P㸧 㸪ࡋࢃࡢ ࡼ࠺࡞ᑠἼࡀ❧ࡘ㸦ྍඣ㸪1978㸸Ỉ㔝࣭ᚚໃ㸪2000㸧༊⏬ࢆᖹ℩㸦G㸧㸪ୗὶ᪉ ྥῡࡽᖹ℩ኚࡍࡿ༊⏬ࢆῡ–ᖹ℩㸦PG㸧ࡋࡓ㸬ࡉࡽ㸪ᖹ℩㸦G㸧 53 ୖὶഃ࡛᥋ࡍࡿ༊⏬ࢆྠࡌᇶ‽࡛ῡ㸦P㸧㸪ᖹ℩㸦G㸧㸪ῡ–ᖹ℩㸦PG㸧ศࡅ㸪 ࡑࢀ௨እࢆࡍ࡚ῡ㸦P㸧ࡋࡓ㸬ࡇࡢ⤖ᯝ㸪2 ༊⏬ࡀᖹ℩㸪4 ༊⏬ࡀᖹ℩–᪩ ℩㸪5 ༊⏬ࡀ᪩℩㸪11 ༊⏬ࡀ᪩℩–ῡ㸪29 ༊⏬ࡀῡ㸪5 ༊⏬ࡀῡ–ᖹ℩ศ㢮ࡉ ࢀࡓ㸬࡞࠾㸪༊⏬ 50 51 ࡣ㸪2008 ᖺ⛅ᮇ᪂ࡋ࠸᪩℩ࡀฟ⌧ࡍࡿ๓ࡣῡ㸪ࡑ ࡢᚋࡣࡑࢀࡒࢀ᪩℩–ῡ㸪ῡ–ᖹ℩ࡋࡓ㸬 ⏘༸ㄪᰝ ⛶㨶ࡢศᕸㄪᰝࢆ⾜࠺๓ᖺ㸦2006–2008 ᖺ㸧ࡢ 10 ᭶ึ᪪–12 ᭶ୖ᪪㸪ㄪᰝ Ỉᇦࢆ㋃ᰝࡋ࡚࣐ࢦࡢ⏘༸≧ἣࢆㄪࡓ㸬Ἑᕝᆺ࣐࣓ࣖࡢ㞤ಶయࡣ⏘༸ 3– 4 ᪥㛫ࢆせࡍࡿࡇࡽ㸦ᮌᮧ㸪1972㸧 㸪ㄪᰝ㢖ᗘࡣࡰ 3 ᪥ 1 ᅇࡋࡓ㸬ㄪ ᰝࡣ㝣ୖࡽࡢ⫗║ほᐹࡋ㸪ᗏ㉁⾲㠃ࡢᅵ◁ࡸ⸴㢮➼ࡀ㝖ࢀ࿘ᅖࡼࡾ᫂ࡿ ࡃኚⰍࡋࡓἙᗋ㒊ศࢆ࣐ࢦࡢ⏘༸ᗋࡳ࡞ࡋ 1/2,500 ࡢᆅᅗୖグ㘓ࡋࡓ㸬 ࣐ࢦࡢ⏘༸ሙᡤᾋୖ⛶㨶ࡢ⏕ᜥሙᡤࡢ⨨㛵ಀࢆㄪࡿࡓࡵ㸪ࡍ࡚ࡢ ༊⏬ࡢ୰ᚰࡽୖὶഃࡢ᭱ᐤࡾࡢ⏘༸ᗋࢆྵࡴ༊⏬ࡢ୰ᚰࡲ࡛ࡢ㊥㞳ࢆồࡵ㸪 ⏘༸ᗋࡽࡢ㊥㞳ࡋࡓ㸬ㄪᰝỈᇦෆ࡛ୖὶഃ⏘༸ᗋࡀ☜ㄆࡉࢀ࡞ࡗࡓ༊ ⏬ࡘ࠸࡚ࡣ㸪ㄪᰝỈᇦୖὶ➃ࡢ⮬↛ࡽ 100 m ୖὶ࠶ࡿᖹ℩ࢆ㸪᭱ࡶ㏆ ࠸⏘༸ሙᡤ௬ᐃࡋ࡚㊥㞳ࢆ⟬ฟࡋࡓ㸬ࡇࡢᖹ℩⮬↛ࡢ㛫ࡢ༊㛫ࡣ໙㓄ࡀ ࡁࡃ㸪࣐ࢦࡢ⏘༸ࡣྍ⬟࡛࠶ࡿ㸦ᮌᮏ㸪ᮍⓎ⾲㸧㸬ࡇࡢᖹ℩࡛ࡣ⏘༸ ᗋࡢ᭷↓ࢆㄪᰝࡋ࡚࠸࡞࠸ࡓࡵ㸪ୖὶഃ⏘༸ᗋࡀ☜ㄆࡉࢀ࡞ࡗࡓ༊⏬ࡢ⏘ ༸ᗋࡽࡢ㊥㞳ࡣ㐣ᑠホ౯࡞ࡿྍ⬟ᛶࡀ࠶ࡿ㸬ࡓࡔࡋ㸪ࡇࢀࡽࡢ༊⏬ࡢ⛶㨶 ಶయᩘࡣᑡ࡞ࡗࡓࡓࡵ㸦ᚋ㏙㸧 㸪ゎᯒࡢᙳ㡪ࡣᑡ࡞࠸ุ᩿ࡋࡓ㸬 ₯Ỉ┠どほᐹ 2007–2009 ᖺࡢ 1–3 ᭶」ᩘᅇࡢ₯Ỉ┠どほᐹࢆ⾜࠸㸪࣐ࢦᾋୖ⛶㨶ࡢศ ᕸࢆㄪࡓ㸬ᮏ◊✲࡛ࡣ㸪ஂಖ㸦1980㸧ࢆཧ⪃㸪ὸࡃὶ㏿ࡀᑠࡉ࠸ᓊἢ࠸ࡢ ሙᡤࢆ⏝ࡍࡿ㛗 5–6cm ࡲ࡛ࡢಶయࢆᾋୖ⛶㨶ᐃ⩏ࡋࡓ㸬2007 ᖺࡣほᐹ ྍ⬟ᮇ㛫ࢆᢕᥱࡍࡿࡓࡵ 1 ᭶ึ᪪ࡽ 3 ᭶ୗ᪪ 1 ᅇ/㐌ࡢ㢖ᗘ࡛ 11 ᅇࡢㄪ ᰝࢆ⾜࠸㸪ᾋୖ⛶㨶ࡀᙜヱᮇ㛫ෆほᐹࡉࢀࡿࡇࢆ☜ㄆࡋࡓ㸬୍᪉㸪ྠᖺ 3 54 ᭶ᮎ᪩℩ὶᚰ㒊⏕ᜥࡋ࡚࠸ࡓ⛶㨶ࡢ㛗ࡣ 4.7–8.3㸦6.5 ± 1.0㸧cm ࡛࠶ࡾ㸦n = 17㸹㛗 ᐃᚋࡍ࡚ᨺὶ㸹ᮌᮏ㸪ᮍⓎ⾲㸧 㸪ࡇࡢᮇ㸪Ỉ῝ὶ㏿ ࡀࡁ࡞ὶᚰ㒊ࡢᚤ⏕ᜥሙᡤኚ㸦ྡ㉺㸪1988㸹Nagata and Yanai, 2002㸹 ⱝᯘ㸪2003㸧ࡀ⏕ࡌࡿ᥎ᐃࡉࢀࡓ㸬ࡋࡓࡀࡗ࡚㸪2008 ᖺ 2009 ᖺࡢㄪ ᰝࡶ 1–3 ᭶ᐇࡋ㸪๓⪅࡛ࡣ 1 ᅇ/᭶ࡢ㢖ᗘ࡛ 3 ᅇ㸪ᚋ⪅࡛ࡣ 1 ᭶ 1 ᅇ㸪2– 3 ᭶ྛ 2 ᅇࡢィ 5 ᅇࡋࡓ㸬ㄪᰝࡣ᪥୰㸦9–16 㸧⾜࠸㸪Ỉ୰࣓࢞ࢿ㸪ࢩ ࣗࣀ࣮ࢣࣝ㸪ࢻࣛࢫ࣮ࢶ➼ࢆ╔⏝ࡋࡓㄪᰝ⪅ 2 ྡࡀㄪᰝỈᇦࡢୗὶ➃ࡽ୧ ᓊศࢀ࡚₯Ỉࡋ㸪ࡑࢀࡒࢀᓊἢࡗ࡚ୖὶ᪉ྥ⛣ືࡋ࡞ࡀࡽᾋୖ⛶㨶ࢆ ィᩘࡋࡓ㸬ほᐹࡣ࣐ࢦࡸࢧࢡ࣐ࣛࢫࡢᾋୖ⛶㨶ࡀศᕸࡍࡿᓊἢ࠸ࡢሙᡤ㸦ྡ ㉺㸪1988㸹Nagata and Yanai, 2002㸧ࢆ୰ᚰᐇࡋࡓ㸬Ỉ῝⣙ 20 cm ௨ୗ ࡢὸ࠸ሙᡤ࡛ࡣ㸪῝࠸ሙᡤࡽぢ㏻ࡍ࡞ࡋ࡚Ỉ୰ࡢど⏺ࢆ☜ಖࡋࡓ㸬▼ࡸ᳜ ≀➼࡛ど⏺ࡀ㐽ࡽࢀࡿሙྜࡣ㸪ᑠᯞ➼࡛ྍど㒊ࢆ่⃭ࡋ࡚⛶㨶ࢆ㏨㑊ࡉࡏ どㄆࡋࡓ㸬௨ୖࡢ᪉ἲ࡛どㄆࡋࡓ⛶㨶ࡢಶయᩘࢆィᩘࡋࡓ㸬ྛㄪᰝ⪅ࡣᾋୖ ⛶㨶ࡢᐃⅬࡽ㝣ୖࡢグ㘓⪅ 1 ྡ☜ㄆಶయᩘࢆ▱ࡽࡏ㸪グ㘓⪅ࡣᶆⅬࢆཧ ↷ࡋ࡚⦰ᑻ 1/2,500 ࡢᆅᅗୖಶయᩘࢆグ㘓ࡋࡓ㸬ࡇࢀࡽࡢ⛶㨶ಶయᩘࢆྛㄪ ᰝ༊⏬ࡢᕥྑᓊู㞟ィࡋ࡚ 112 ಶࡢࢹ࣮ࢱࢭࢵࢺࡋ㸪௨ୗࡢゎᯒ౪ࡋࡓ㸬 ࡲࡓ㸪2008–2009 ᖺࡢ₯Ỉ┠どほᐹࡣ㸪┠┒ࡁሷࣅ⟶㸦⣙ 1 m㸧ࢆ⏝࠸ ࡚ᾋୖ⛶㨶ᐃⅬࡢỈ῝㸦௨ୗ㸪ᐃỈ῝㸧ࢆ 1 cm ༢࡛ ᐃࡋࡓ㸬ᐃỈ ῝ࡀ 1 m ௨ୖࡢሙྜࡣ㸪ㄪᰝ⪅ࡢ㌟㛗ࡸἢᓊỈ῝ࡢ ᐃ⤖ᯝࡽ 0.1 m ༢࡛ Ỉ῝ࢆ᥎ᐃࡋࡓ㸬」ᩘಶయࡢᐃⅬ࡛ࡣ㸪㞟ᅋࡢ୰ᚰ㒊ศࡢỈ῝ࢆ ᐃࡋ࡚ྛ ಶయࡢ್ࡋࡓ㸬ࡉࡽ㸪2007 ᖺ 3 ᭶ࡣ㸪ᾋୖ⛶㨶ࡢ⏕ᜥᩘࡀከ࠸ 3 ༊⏬㸦ᚋ ㏙㸧࡛ᐃⅬࡢ 60%Ỉ῝ࡢὶ㏿ࢆ ᐃࡋࡓ㸬 ⤫ィゎᯒ 2008–2009 ᖺࡢ࣐ࢦᾋୖ⛶㨶ࡢᐃỈ῝ࡘ࠸࡚㸪Jonckheere-Terpstra ᳨ᐃ ࡼࡾ⤒ኚࡢഴྥࢆㄪࡓ㸬ྛㄪᰝ᪥ࡢᐃỈ῝ᕪࡀ࡞࠸ࡍࡿᖐ↓௬ ㄝᑐࡋ㸪༢ㄪቑຍࡲࡓࡣ༢ㄪῶᑡࡢഴྥࢆᑐ❧௬ㄝࡋ᳨࡚ᐃࢆ⾜ࡗࡓ㸦⚄ ⏣㸪2012㸧㸬 ࣐ࢦᾋୖ⛶㨶ࡢ⏕ᜥሙᡤ⏝㛵㐃ࡍࡿ⎔ቃせᅉࢆ᥎ᐃࡍࡿࡓࡵ㸪 55 ྛ༊⏬ࡢᕥྑᓊูࡢ⛶㨶ಶయᩘࢆᛂ⟅ኚᩘ㸪ྛ༊⏬ࡢὶ㊰༢ࢱࣉ㸦G㸪GR㸪 R㸪RP㸪P㸪PG㸧⎔ቃኚ㔞㸦Ỉ㠃ᖜ㸪᭱Ỉ῝㸪ᕥྑᓊูࡢἢᓊỈ῝㸪ὶ㏿㸪 ᳜≀⿕ᗘ㸪࠾ࡼࡧ⏘༸ᗋࡽࡢ㊥㞳㸧ࢆㄝ᫂ኚᩘࡋ࡚㸪୍⯡⥺ᙧΰྜࣔࢹ ࣝ㸦GLMM㸸Bolker et al., 2009㸪2012㸧ࢆᵓ⠏ࡋࡓ㸬ᾋୖ⛶㨶ಶయᩘࡢศᩓࡣ ࡰࡍ࡚ࡢㄪᰝ᪥࡛ᖹᆒࢆୖᅇࡾ㸪࣏ࢯࣥศᕸࢆ௬ᐃࡋࡓሙྜࡣ㐣ศᩓ ࡞ࡗࡓࡓࡵ㸪ᛂ⟅ኚᩘࡢ☜⋡ศᕸ㈇ࡢ㡯ศᕸࢆ௬ᐃࡋࡓ㸦⢑㇂㸪2012㸧 㸬 ศᯒඛ❧ࡕ㸪㔞ⓗኚᩘ࡛࠶ࡿ 6 ࡘࡢ⎔ቃኚ㔞ࡘ࠸࡚⥲ᙜࡓࡾ࡛ࣆࢯࣥࡢ ✚⋡┦㛵ಀᩘࢆồࡵࡓࡇࢁ㸪᭱Ỉ῝ἢᓊỈ῝ࡢ㛫࡛㧗࠸┦㛵ࡀ᳨ฟࡉࢀ ࡓ㸦r = 0.62㸧㸬ࡑࡇ࡛㸪Ỉ㠃ᖜࢆ᭱Ỉ῝࡛㝖ࡋࡓ್ࢆᕝᖜỈ῝ẚ㸦B/H㸧ࡋ ࡚ㄝ᫂ኚᩘຍ࠼㸪᭱Ỉ῝Ỉ㠃ᖜࢆ㝖ཤࡋࡓ㸬ᕝᖜỈ῝ẚࡣὶ㊰༢ࡢᙧ ែᐦ᥋㛵㐃ࡍࡿࡇࡀ▱ࡽࢀ࡚࠸ࡿ㸦ⴛሙ㸪2003b㸧 㸬ࡲࡓ㸪࢝ࢸࢦࣜ ࢝ࣝኚᩘ࡛࠶ࡿὶ㊰༢ࢱࣉࡣࢲ࣑࣮ኚᩘࡋ࡚ࣔࢹࣝ⤌ࡳ㎸ࡲࢀࡓ㸬ྛ ὶ㊰༢ࢱࣉࡢಀᩘࡢ᥎ᐃ⢭ᗘࢆྥୖࡉࡏࡿࡓࡵ㸪᭱ࡶ༊⏬ᩘࡢከ࠸ῡ㸦P㸧 ࢆཧ↷࢝ࢸࢦ࣮ࣜࡋࡓ㸬ࡇࢀࡼࡾῡࡢಀᩘࡣࢮࣟ࡞ࡾ㸪ࡢὶ㊰༢ࢱ ࣉࡢಀᩘࡣῡᑐࡍࡿ㔜ࡳࡋ࡚⾲ࡉࢀࡿ㸬ゎᯒࡣㄪᰝᖺࡈㄪᰝ᪥ࡢࢹ ࣮ࢱࢆࣉ࣮ࣝࡋ࡚⾜࠸㸪ྛㄪᰝ᪥ࢆࣛࣥࢲ࣒ຠᯝࡋࡓ㸬ࡍ࡚ࡢㄝ᫂ኚᩘࡢ ⥺ᙧ⤖ྜࡼࡿࣔࢹࣝࢆ࣮࣋ࢫࡋ࡚㸪ㄝ᫂ኚᩘࡢ⥲ᙜࡓࡾࡢ⤌ࡳྜࢃࡏ࡛ࣔ ࢹࣝࢆᵓ⠏ࡋ㸪AIC㸦㉥ụሗ㔞ᇶ‽㸧ࡀ᭱ᑠࡢࣔࢹࣝࢆ࣋ࢫࢺࣔࢹࣝࡋ࡚ 㑅ᢥࡋࡓ㸦Barton, 2015㸧 㸬ᾋୖ⛶㨶ಶయᩘ⎔ቃせᅉࡢ㛵ಀࡣ㸪࣋ࢫࢺࣔࢹࣝ ຍ࠼㸪࣋ࢫࢺࣔࢹࣝࡢ AIC ᕪ㸦'AIC㸧ࡀ 2 ᮍ‶ࡢࣔࢹࣝࢆࡍ࡚⪃៖ࡋ ࡚᥎ᐃࡋࡓ㸦Burnham and Anderson, 2002㸧㸬௨ୖࡢゎᯒࡣ⤫ィࢯࣇࢺ R㸦R Development Core Team, 2005㸧㸪࠾ࡼࡧࡇࢀࢆ࣮࣋ࢫࡋࡓࣇ࣮ࣜ⤫ィࢯࣇࢺ EZR㸦⚄⏣㸪2012㸧࡛⾜࠸㸪᭷ពỈ‽ࢆ 0.05 ࡋࡓ㸬 ⤖ ᯝ ⏘༸ᗋᾋୖ⛶㨶ࡢศᕸ ⏘༸ᗋࡣ 2006 ᖺ⛅ 4 ༊⏬㸦16㸪23㸪33㸪36㸧㸪2007 ᖺ⛅ 3 ༊⏬㸦12㸪21㸪 56 36㸧 㸪2008 ᖺ⛅ 3 ༊⏬㸦27㸪36㸪51㸧࡛☜ㄆࡉࢀ㸪ྛ༊⏬ࡢὶ㊰༢ࢱࣉ ࡣῡ–ᖹ℩㸦༊⏬ 16㸪23㸪36㸪51㸧 㸪ᖹ℩–᪩℩㸦༊⏬ 12㸪27㸧 㸪ᖹ℩㸦༊⏬ 21㸧㸪 ῡ㸦༊⏬ 33㸧࡛࠶ࡗࡓ㸦Figs. 4-2–4-4㸧 㸬ྛ༊⏬ࡢ⏘༸ᗋࡽࡢ㊥㞳ࡣ 0–650 m 㸦3 ᖺ㛫ࡢᖹᆒ್ ± ᶆ‽೫ᕪ = 223 ± 165 m㸧ィ⟬ࡉࢀࡓ㸦Table 4-1㸧㸬ㄪᰝ ᪥ࡈࡢ࣐ࢦᾋୖ⛶㨶ࡢಶయᩘࡣ 2007 ᖺ 50–730 ಶయ㸪2008 ᖺ 5–70 ಶ య㸪2009 ᖺ 60–575 ಶయኚືࡋࡓࡀ㸪ㄪᰝᖺෆ࡛ࡣ 1 ᭶ᮎ–2 ᭶୰᪪ᴟ 㐩ࡍࡿඹ㏻ࡢ⤒ኚࢆ♧ࡋࡓ㸦Figs. 4-2–4-4㸧 㸬ᕥྑᓊࡢ༊⏬࠶ࡓࡾࡢ⛶㨶 ಶయᩘࡣ 2007 ᖺ 0–111 ಶయ㸪2008 ᖺ 0–12 ಶయ㸪2009 ᖺ 0–185 ಶయ࡛࠶ ࡗࡓ㸦Figs. 4-2–4-4㸧㸬ᾋୖ⛶㨶ࡢศᕸࣃࢱ࣮ࣥࡣᖺࡈ␗࡞ࡾ㸪2007–2008 ᖺࡣୗὶഃࡽ୰ኸ㒊㸦ࡑࢀࡒࢀ༊⏬ 9–33 ༊⏬ 16–32㸧㸪2009 ᖺࡣἙ㐨 ⊃✽㒊ࢆྵࡴୖὶഃ㸦༊⏬ 35–50㸧࡛ಶయᩘࡀከࡗࡓ㸬ᾋୖ⛶㨶ࡢᐃỈ῝ ࡣ 㸪 2008 ᖺ 0.07–0.6 m ࡢ ⠊ ᅖ ࠶ ࡾ ⤒ ⓗ ᭷ ព ቑ ຍ ࡋ ࡓ ࡀ 㸦Jonckheere-Terpstra ᳨ᐃ㹙ᑐ❧௬ㄝ㸸༢ㄪቑຍ㹛 㸪JT = 1,526㸪P = 0.003㸧 㸪2009 ᖺ ࡣ 0.03–4.0 m ᗈ ⠊ ᅖ ࠾ ࡼ ࡧ ⤒ ⓗ ᭷ ព ῶ ᑡ ࡋ ࡓ 㸦Jonckheere-Terpstra ᳨ᐃ㹙ᑐ❧௬ㄝ㸸༢ㄪῶᑡ㹛㸪JT = 387,813㸪P = 0.023㸹Table 4-2㸧㸬2007 ᖺ 3 ᭶༊⏬ 10㸦᪩℩–ῡ㸧㸪11㸦᪩℩㸧 㸪26㸦᪩℩–ῡ㸧࡛ ᐃࡋࡓ ᐃⅬࡢὶ㏿ࡣ 0.7–19.7㸦9.4 ± 9.6㸧cm/s㸪Ỉ῝ࡣ 12–18㸦14.3 ± 0.3㸧cm ࡛࠶ࡗ ࡓ㸬 ᾋୖ⛶㨶ಶయᩘ⎔ቃせᅉࡢ㛵ಀ ㄪᰝᖺࡈࡢ࣋ࢫࢺࣔࢹࣝ'AIC ࡀ 2 ᮍ‶ࡢࣔࢹࣝࢆ Table 4-3–4-5 ♧ࡋ ࡓ㸬࣐ࢦᾋୖ⛶㨶ࡢಶయᩘ᭷ព࡞┦㛵ࢆᣢࡘ⎔ቃせᅉ㸦ಀᩘࡢ᥎ᐃ್ࡢ 95% ಙ㢗༊㛫 0 ࡀྵࡲࢀ࡞࠸㸧ࡋ࡚㸪2007 ᖺࡣࡍ࡚ࡢࣔࢹࣝ࠾࠸࡚ᖹ℩ –᪩℩᪩℩–ῡࡢὶ㊰༢ࢱࣉ㸪ἢᓊỈ῝㸪ᕝᖜỈ῝ẚ㸪⏘༸ᗋࡽࡢ㊥㞳 ࡀ㑅ᢥࡉࢀࡓ㸦Table 4-3㸧㸬2008 ᖺࡣࡍ࡚ࡢࣔࢹ࡛ࣝ⏘༸ᗋࡽࡢ㊥㞳ࡀ ᭷ព࡞┦㛵ࢆ♧ࡋࡓ㸪 ὶ㊰༢ࢱࣉࡀ㑅ᢥࡉࢀࡓࣔࢹ࡛ࣝࡣᖹ℩–᪩℩ ᪩℩–ῡࡢὶ㊰༢ࢱࣉࡀ᭷ព࡞┦㛵ࢆ♧ࡋࡓ㸦Table 4-4㸧㸬2009 ᖺࡣࡍ ࡚ࡢࣔࢹࣝ࠾࠸࡚ᖹ℩ࡢὶ㊰༢ࢱࣉ㸪ἢᓊỈ῝㸪ὶ㏿㸪⏘༸ᗋࡽࡢ㊥ 57 㞳ࡀ㑅ᢥࡉࢀࡓ㸦Table 4-5㸧 㸬᭷ព࡞┦㛵ࡀぢࡽࢀࡓኚᩘࡢ࠺ࡕ㸪ᖹ℩–᪩℩㸪 ᪩℩–ῡࡢὶ㊰༢ࢱࣉᕝᖜỈ῝ẚࡢಀᩘࡢ➢ྕࡣṇ㸪 ᖹ℩ࡢὶ㊰༢ࢱ ࣉ㸪ἢᓊỈ῝㸪ὶ㏿㸪⏘༸ᗋࡽࡢ㊥㞳࡛ࡣ㈇࡛࠶ࡗࡓ㸬ྛㄪᰝᖺ࠾ࡅࡿὶ ㊰༢ࢱࣉࡢಀᩘࡢᖹᆒ್㸦2007 ᖺ㸸0.57 ± 0.37㸹2008 ᖺ㸸0.93 ± 0.56㸹2009 ᖺ㸸-0.08 ± 0.68㸧ࡣ᭷ព␗࡞ࡾ㸦୍ඖ㓄⨨ศᩓศᯒ㸪P < 0.001㸧㸪2009 ᖺ ࡣ㸪2007 ᖺ 2008 ᖺẚ᭷ពప࠸್ࢆ♧ࡋࡓ㸦Tukey ἲࡼࡿከ㔜ẚ㍑㸸 2007 ᖺ 2008 ᖺ㸪P = 0.28㸹2007 ᖺ 2009 ᖺ㸪P = 0.004㸹2008 ᖺ 2009 ᖺ㸪 P < 0.001㸧 㸬 ⪃ ᐹ ࣐ࢦᾋୖ⛶㨶ಶయᩘ㛵㐃ࡍࡿ⎔ቃせᅉ ᮏ◊✲ࡼࡾ㸪࣐ࢦ⏕ᜥᇦୗὶഃ࠾࠸࡚ᾋୖ⛶㨶ࡢಶయᩘᙉࡃ┦㛵ࡍ ࡿὶ㊰༢ࢫࢣ࣮ࣝࡢ⎔ቃせᅉࡣ㸪ὶ㊰༢ࢱࣉ㸪ἢᓊỈ῝㸪⏘༸ᗋࡽࡢ ㊥㞳࡛࠶ࡿࡇࡀ᫂ࡽ࡞ࡗࡓ㸬ὶ㊰༢ࢱࣉ࡛ࡣ㸪᪩℩࿘㎶ࡢ༊⏬㸪≉ ᖹ℩–᪩℩᪩℩–ῡࡀ 2007 ᖺ 2008 ᖺ㧗࠸┦㛵ࢆ♧ࡋࡓ㸦Tables 4-3–4-5㸧㸬 ಀᩘࡢ➢ྜࡀṇ࡛࠶ࡗࡓࡇࡽ㸪ࡇࢀࡽࡢ༊⏬࡛ࡣཧ↷࢝ࢸࢦ࣮࡛ࣜ࠶ࡿῡ ẚᾋୖ⛶㨶ࡀከࡗࡓ࠸࠼ࡿ㸬 ୧༊⏬ࡣ᪩℩ࢆྵࡴ᪩℩᥋ࡍࡿࡓࡵ㸪 ࣐ࢦࡸࢧࢡ࣐ࣛࢫࡢᾋୖ⛶㨶࡛ሗ࿌ࡉࢀ࡚࠸ࡿᚤ⏕ᜥሙᡤࢫࢣ࣮ࣝ 㸦ᐃⅬ㸧 ࠾ࡅࡿᑠࡉ࡞ὶ㏿ᇦࡢ⏝㸦ྡ㉺㸪1988㸹Nagata and Yanai, 2002㸧▩ ┪ࡍࡿࡼ࠺ぢ࠼ࡿ㸬ࡓࡔࡋ㸪ࢱࢭࣚ࢘ࢧࢣ࡛ࡣ㐣ᑠ࡞ὶ㏿ࡀ㑊ࡅࡽࢀ 㸦Girard et al., 2004㸧 㸪ὶ㏿ࡀ㐣ᑠ࡞ᐃⅬࡢಶయࡣ⛣ືࡋࡸࡍ࠸㸦Steingrímsson and Grant, 2003㸧 㸬ࡲࡓ㸪」ᩘࡢ✵㛫ࢫࢣ࣮࡛ࣝࢧࢣ⛉㨶㢮ࡢ⏕ᜥሙᡤࢆㄪࡓ ◊✲࡛ࡣ㸪ᐃⅬࡢὶ㏿ࡣᑠࡉ࠸ࡀ㸪ࡼࡾୖࡢࢫࢣ࣮࡛ࣝࡣࡁ࡞ὶ㏿ࢆᣢ ࡘሙᡤ㏆᥋ࡍࡿሗ࿌ࡉࢀ࡚࠸ࡿ㸦Bozek and Rahel, 1991㸹Inoue and Nakano, 1999㸧㸬ࡑࡢࡼ࠺࡞ᚤ⏕ᜥሙᡤࡣ㸪㏿࠸ὶࢀ㏆᥋ࡋࡓᕝᓊࡸ᪩℩ෆ㒊ࡢ㸪᳜≀ ᳜⏕ࡸ▼࡞ࡢỈὶࡢ㐽࠸≀㏆⨨ࡋ㸦Moore and Gregory, 1988㸹Bozek and Rahel, 1991㸧㸪ᐃࡋࡓ⛶㨶ᑐࡋ࡚㸪ὶୗ㣵⏕≀ࡢᤕ㣗ᶵࡢቑ㐟Ὃ 58 ࢚ࢿࣝࢠ࣮ࡢ⠇⣙ࢆࡶࡓࡽࡋ㸪᭱ࡢ࢚ࢿࣝࢠ࣮ຠ⋡ࢆ࠼ࡿ㸦Fausch, 1984㸧㸬 ᮏㄪᰝỈᇦ࡛ࡶ㸪᪩℩࿘㎶ࡢ༊⏬࡛ ᐃࡋࡓ⛶㨶ࡢᐃⅬࡢὶ㏿ࡣ᪤ሗ㸦ྡ㉺ 㸪1988㸹Nagata and Yanai, 2002㸹ⱝᯘ㸪2003㸧ࡢ⠊ᅖෆ࠶ࡗࡓ㸬௨ ୖࡢࡇࡽ㸪ὶ㊰༢ࢫࢣ࣮ࣝ࠾ࡅࡿ᪩℩࿘㎶ࡢ༊⏬ࡢ⏝ࡣ㸪ᚤ⏕ᜥሙ ᡤࢫࢣ࣮ࣝ࠾ࡅࡿᑠࡉ࡞ὶ㏿ᇦࡢ⏝▩┪ࡏࡎ㸪ࡴࡋࢁ㸪ዲ㐺࡞ᚤ⏕ᜥሙ ᡤ࡛࠶ࡿࠕ㏿࠸ὶࢀ㏆࠸⦆ὶ㒊ศ㸦Bozek and Rahel, 1991㸹Inoue and Nakano, 1999㸧ࠖࡀᐇ⌧ࡉࢀࡿࡓࡵᚲせ࡞๓ᥦ᮲௳㸦Kramer et al., 1997㸧ᤊ࠼ࡿࡇ ࡀ࡛ࡁࡿ㸬 ḟ㸪ἢᓊỈ῝ࡢಀᩘࡢ➢ྕࡣ㈇࡛࠶ࡗࡓࡇࡽ㸪ἢᓊỈ῝ࡀὸ࠸༊⏬ ࣐ࢦᾋୖ⛶㨶ࡀከࡗࡓ࠸࠼ࡿ㸦Tables 4-3–4-5㸧㸬ᑠࡉ࡞Ỉ῝ࡣ㸪ࢧࢣ ⛉㨶㢮ࡢᾋୖ⛶㨶ࡢᚤ⏕ᜥሙᡤࡢ≉ᚩࡋ࡚ࡼࡃ▱ࡽࢀ࡚࠾ࡾ㸦Moore and Gregory, 1988㸹ྡ㉺㸪1988㸹Nagata and Yanai, 2002㸧 㸪ᮏ◊✲࠾࠸࡚ࡶ㸪 ὶ㊰༢ࢫࢣ࣮࡛ࣝἢᓊỈ῝ࡢὸ࠸༊⏬ࡣ㸪࣐ࢦᾋୖ⛶㨶ᑠࡉ࡞Ỉ῝ࡢᚤ ⏕ᜥሙᡤࢆᥦ౪ࡋࡓ⪃࠼ࡽࢀࡿ㸬 ⏘༸ᗋࡽࡢ㊥㞳ࡣ㸪⪃៖ࡋࡓࡍ࡚ࡢࣔࢹ࡛࣐ࣝࢦᾋୖ⛶㨶ಶయᩘ᭷ ព࡞┦㛵ࢆ♧ࡋࡓࡇࡽ㸦Tables 4-3–4-5㸧㸪⛶㨶ಶయᩘ᭱ࡶᐦ᥋࡞㛵ಀࢆ ᣢࡘ⎔ቃせᅉ⪃࠼ࡽࢀࡓ㸬ಀᩘࡢ➢ྕࡀ㈇࡛࠶ࡗࡓࡇࡣ㸪⏘༸ᗋࡽࡢ㊥ 㞳ࡀ㏆࠸༊⏬࣐ࢦᾋୖ⛶㨶ࡀከࡗࡓࡇࢆ♧ࡋ࡚࠸ࡿ㸦Tables 4-3– 4-5㸧㸬ࢧࢣ⛉㨶㢮ࡢᾋୖ⛶㨶ࡀ⏘༸ᗋ㏆ከ࠸ࡇࡣ㸪࢝ࢵࢺࢫ࣮ࣟࢺࢺࣛ ࢘ࢺ㸦Bozek and Rahel, 1991㸧 㸪ࢱࢭࣚ࢘ࢧࢣ㸦Foldvik et al., 2010㸹Teichert et al., 2011㸧➼࡛ሗ࿌ࡀ࠶ࡿ㸬ᾋୖ⛶㨶ࡀ⏘༸ᗋ㏆ከ࠸⌮⏤ࡣ㸪ప࠸㐟Ὃ⬟ຊ 㸦Moore and Gregory, 1988㸹┿ᒣ㸪1992㸧ࡢࡓࡵࡶ⪃࠼ࡽࢀࡿࡀ㸪Ỉὶࡼࡿ ཷືⓗ࡞ὶୗࡣ㸪㐟Ὃ⬟ຊ↓㛵ಀྍ⬟࡛࠶ࡿࡇࡽ㸦Einum et al., 2011, 2012㸧㸪ࢱࢭࣚ࢘ࢧࢣ࡛ࡣ⛣ືࡼࡗ࡚ዲ㐺࡞ᚤ⏕ᜥሙᡤ฿㐩࡛ࡁࡿಖド ࡀ࡞࠸ࡓࡵ㸪ࡲࡓࡣ✀࠾ࡼࡧྠ✀㧗㱋㨶ࡼࡿ⿕㣗ࣜࢫࢡࡀࡁ࠸ࡓࡵ㸪 ⛶㨶ࡣ⏘༸ᗋ㏆ࡽ⛣ືࡋ࡞࠸᥎ ࡉࢀ࡚࠸ࡿ㸦Foldvik et al., 2010㸹Einum et al., 2011, 2012㸧 㸬ࡇࢀ㛵ࡋ࡚㸪࣐ࢦࡸ࣐࣓ࣖࡢ⏘༸ᗋࡀᖹ℩ࡸῡᑼᙧᡂ ࡉࢀࡿࡇ㸦ⓑ▼㸪1957㸹ᮌᮧ㸪1972㸹୰ᮧ㸪1999a㸧㸪Ἑᕝࡢ୰࣭ୗὶᇦ 59 ࡛ࡣᖹ℩ࡢୗὶഃ᪩℩ࡀ⨨ࡍࡿࡇ㸦ྍඣ㸪1978㸹Ỉ㔝࣭ᚚໃ㸪2000㸹ⴛ ሙ㸪2003a㸧ࢆ⪃៖ࡍࡿ㸪࣐ࢦ⏕ᜥᇦୗὶഃ࡛ࡣ㸪⏘༸ᗋࡢୗὶഃ᪩ ℩ࡀᏑᅾࡍࡿࡇࡀ♧၀ࡉࢀࡿ㸬ᮏㄪᰝỈᇦ࡛ࡶ㸪࣐ࢦࡢ⏘༸ᗋࡣᖹ℩࿘㎶ ࡛☜ㄆࡉࢀࡓࡇࡽ㸦Figs. 4-2–4-4㸧㸪ᾋୖ⛶㨶ࡣཷືⓗ࡞ὶୗࡼࡗ࡚᪩℩ ࢆ㏻㐣࡛ࡁࡿ⪃࠼ࡽࢀࡿࡀ㸪ከࡃࡢ⛶㨶ࡣ⏘༸ᗋ㏆࠸༊⏬ࡸ᪩℩࿘㎶ࡢ༊ ⏬࡛☜ㄆࡉࢀࡓ㸦Tables 4-3–4-5㸧㸬ࡇࢀࡣప࠸㐟Ὃ⬟ຊࡸཷືⓗ࡞ὶୗ௨እࡢ せᅉࢆ♧၀ࡍࡿࡶࡢ࡛࠶ࡿ㸬ᮏ◊✲࡛ࡣ㸪㨶✀ࡲࡓࡣྠ✀㧗㱋㨶ࡼࡿ࣐ ࢦᾋୖ⛶㨶ࡢᤕ㣗ࢆㄪᰝࡋ࡚࠸࡞࠸ࡀ㸪ࢧࢡ࣐ࣛࢫ㸦࣐࣓ࣖ㸧ࡢᾋୖ⛶㨶࡛ࡣ 」ᩘࡢሗ࿌ࡀ࠶ࡾ㸦┿ᒣ㸪1999㸹ୖ㔝㸪2009㸧 㸪ᮏㄪᰝỈᇦࡣ࣐ࢦ㧗㱋 㨶ࡀ⏕ᜥࡋ࡚࠸ࡓ㸦ᮌᮏ㸪2015㸧㸬ࡋࡓࡀࡗ࡚㸪࣐ࢦ⏕ᜥᇦୗὶഃࡢᾋୖ ⛶㨶ࡶ㸪㣚㣹ࡸ⿕㣗ࢆ㑊ࡅࡿࡓࡵ⏘༸ᗋ㏆␃ࡲࡗ࡚࠸ࡿྍ⬟ᛶࡀ⪃࠼ࡽ ࢀࡿ㸬 ⏕ᜥሙᡤ⏝ࡢᖺኚືせᅉ ࣐ࢦᾋୖ⛶㨶ࡢಶయᩘ┦㛵ࡍࡿ⎔ቃせᅉࡣ㸪⏘༸ᗋࡽࡢ㊥㞳௨እ࡛ࡣ ㄪᰝᖺࡈ␗࡞ࡗࡓ㸦Tables 4-3–4-5㸧 㸬ἢᓊỈ῝ࡣ 2008 ᖺ᭷ព࡞┦㛵ࢆ♧ ࡉ࡞ࡗࡓࡀ㸪᳨ウࡋࡓࡍ࡚ࡢࣔࢹ࡛ࣝಀᩘࡢ➢ྕࡀ㈇࡛࠶ࡗࡓࡇࡽ 㸦Tables 4-3–4-5㸧 㸪2008 ᖺࡢᾋୖ⛶㨶ࡶࡢᖺྠᵝἢᓊỈ῝ࡢὸ࠸༊⏬ࢆ ⏝ࡋࡓ⪃࠼ࡽࢀࡿ㸬ὶ㊰༢ࢱࣉ࡛ࡣ㸪2007–2008 ᖺࡣ᪩℩࿘㎶ࡢ༊ ⏬ࡀᾋୖ⛶㨶ಶయᩘ᭷ព࡞ṇࡢ┦㛵ࢆ♧ࡋࡓࡀ㸪2009 ᖺࡣᖹ℩ࡢὶ㊰༢ ࢱࣉ࡛᭷ព࡞㈇ࡢ┦㛵ࡀぢࡽࢀ㸪ὶ㊰༢ࢱࣉࡢಀᩘࡢᖹᆒ್ࡣࡢᖺࡼ ࡾ᭷ពపࡗࡓ㸦Tables 4-3–4-5㸧㸬ὶ㊰༢ࢱࣉࡢಀᩘࡀཧ↷࢝ࢸࢦ࣮ࣜ ࡛࠶ࡿῡᑐࡍࡿ㔜ࡳ࡛࠶ࡿࡇࢆ⪃៖ࡍࡿ㸪 ࡇࢀࡽࡢࡇࡣ 2009 ᖺࡢᾋୖ ⛶㨶ࡀࡢᖺẚῡከࡗࡓࡇࢆ♧ࡍࡶࡢ࡛࠶ࡿ㸬ᐇ㝿㸪2009 ᖺᾋ ୖ⛶㨶ࡀከࡃぢࡽࢀࡓୖὶഃࡢ 16 ༊⏬㸦35–50㸧ࡢ࠺ࡕ 11 ༊⏬㸦37–47㸧ࡣῡ ࡛࠶ࡗࡓ㸬 ࡓࡔࡋ㸪༊⏬ 37–47 ࡢ⛶㨶ಶయᩘࡣ 2–3 ᭶ࡅ࡚ᛴῶࡋ࡚࠸ࡓ㸦Fig. 4-4㸧㸬 ࡲࡓ㸪ࢧࢣ⛉㨶㢮ࡢᾋୖ⛶㨶ࡢᐃỈ῝ࡣᡂ㛗క࠸ቑࡍࡿࡇࡀ▱ࡽࢀ㸦ྡ 60 ㉺㸪1988㸹Nagata and Yanai, 2002㸹ⱝᯘ㸪2003㸧㸪ᮏ◊✲࡛ࡶ 2008 ᖺ ࡣྠᵝࡢഴྥࡀぢࡽࢀࡓࡀ㸪2009 ᖺࡣࡴࡋࢁึᮇࡁࡃ⤒ⓗపୗࡋ ࡓ㸦Tables 4-2㸧㸬ࡇࢀࡣ㸪ࡁ࡞ᐃỈ῝ࢆ♧ࡍ༊⏬ 37–47 ศᕸࡋ࡚࠸ࡓ⛶ 㨶ࡀ 2–3 ᭶ࡅ࡚ῶᑡࡋࡓࡓࡵ⪃࠼ࡽࢀࡿ㸦Fig. 4-4㸧 㸬ᮏ◊✲࡛ࡣ⛶㨶ࢆ ಶయ㆑ูࡋ࡚࠸࡞࠸ࡓࡵ㸪༊⏬ 37–47 ࡛ぢࡽࢀ࡞ࡃ࡞ࡗࡓಶయࡢ⾜᪉ࡣศࡽ ࡞࠸ࡀ㸪࢝ࢵࢺࢫ࣮ࣟࢺࢺࣛ࢘ࢺࡢᾋୖ⛶㨶ࡣ㸪⦕㎶㒊ࡀษࡾ❧ࡕỈ῝ࡀࡁ ࠸ὶ㊰࡛ࡣ㸪ዲ㐺࡞⏕ᜥሙᡤ㐩ࡍࡿࡲ࡛ὶୗࡍࡿ᥎ ࡉࢀ࡚࠸ࡿ㸦Moore and Gregory, 1988㸧㸬ᮏㄪᰝỈᇦ࡛ࡶ㸪༊⏬ 37 ࡽୖὶഃࡣ୧ᓊࡀษࡾ❧ࡕỈ῝ ࡀࡁ࡞Ἑ㐨⊃✽㒊ࡢᬒほࢆ♧ࡍࡇࡽ㸪2009 ᖺ 2 ᭶༊⏬ 37–47 ࡛☜ㄆࡉ ࢀࡓ⛶㨶ࡣ㸪3 ᭶ࡅ࡚ୗὶ᪉ྥ⛣ືࡋࡓྍ⬟ᛶࡀ⪃࠼ࡽࢀࡿ㸬 ྠᵝ㸪2009 ᖺ 2–3 ᭶ࡣ༊⏬ 48–50 ࡛ࡶ⛶㨶ಶయᩘࡀᛴῶࡋࡓ㸦Fig. 4-4㸧 㸬 ࡇࢀࡽࡢ༊⏬ࡢὶ㊰༢ࢱࣉࡣከࡃࡢ⛶㨶ࡀ⏝ࡍࡿண ࡉࢀࡓ᪩℩–ῡ ᖹ℩–᪩℩࡛࠶ࡗࡓࡀ㸪ࡇࡢሙᡤࡣἙ㐨⊃✽㒊࡛࠶ࡿࡓࡵᕝᓊࡀ⊃ࡃ㸪ᒾ┙ࡀ 㟢ฟࡋ㸪᳜≀⿕ᗘࡀపࡗࡓ㸬ᕝᓊࡢ▼ࡸ᳜≀᳜⏕ࡣࡁ࡞ὶ㏿ࢆపῶࡋ࡚⦆ ὶ㒊ศࢆ⏕ᡂࡍࡿࡓࡵ㸪ࢧࢣ⛉㨶㢮ࡢᾋୖ⛶㨶ࡢᚤ⏕ᜥሙᡤ᮲௳ࡋ࡚㔜せ࡛ ࠶ࡿࡇࡀ▱ࡽࢀ࡚࠸ࡿ㸦Moor and Gregory, 1988㸹ර⸨㸪1992㸹Nagata and Yanai, 2002㸧 㸬ࡉࡽ㸪ࡇࡢㄪᰝᖺࡣ༊⏬ 51 ࡼࡾୖὶ࡛࣐ࢦᾋୖ⛶㨶ࡀ☜ ㄆࡉࢀ࡞ࡗࡓࡇࡽ㸦Fig. 4-4㸧 㸪2009 ᖺ 2–3 ᭶༊⏬ 48–50 ࡛ぢࡽࢀࡓ⛶ 㨶ࡢᛴῶࡣ㸪ᚤ⏕ᜥሙᡤ᮲௳ࡢ㐺ྜࡼࡿ⛶㨶ࡢὶୗࡼࡗ࡚⏕ࡌࡓ᥎ᐃ ࡉࢀࡓ㸬ࡇࢀࡽࡢ᥎ᐃࡀṇࡋࡅࢀࡤ㸪2009 ᖺࡢ⛶㨶ぢࡽࢀࡓῡࢆ⏝ࡍࡿഴ ྥࡣ㸪⛣ື୰ࡢಶయࢆほ ࡋࡓࡓࡵ᳨ฟࡉࢀࡓ⪃࠼ࡽࢀࡿ㸬 ࣐ࢦ⏕ᜥᇦୗὶഃ࠾ࡅࡿᾋୖ⛶㨶ࡢಖ Ἑᕝࡢୖὶᇦ࡛ࡣ㸪࣐ࢦࡸࢧࢡ࣐ࣛࢫࡢᾋୖ⛶㨶ࡣ㸪ὶ㊰༢ࢫࢣ࣮ࣝࡢ ⏕ᜥሙᡤࡋ࡚ᖹ℩㸦ஂಖ㸪1980㸧ࡲࡓࡣῡ㸦ྡ㉺㸪1988㸧ࢆ⏝ࡍࡿ ሗ࿌ࡉࢀ࡚࠾ࡾ㸪ᮏ◊✲࡛ぢࡽࢀࡓ᪩℩࿘㎶ࡢὶ㊰༢ࢱࣉࢆ⏝ࡍࡿഴྥ ࡣ␗࡞ࡗ࡚࠸ࡿ㸬࢝ࢵࢺࢫ࣮ࣟࢺࢺࣛ࢘ࢺ࡛ࡣ㸪ᾋୖ⛶㨶ࡢዲ㐺࡞ᚤ⏕ᜥሙ ᡤࡢ᮲௳ࡣ㸪 ୖࢫࢣ࣮ࣝࡢᆅᙧ᮲௳➼ࡀ␗࡞ࡿỈᇦ㛫࡛ࡶඹ㏻ࡋ࡚࠾ࡾ㸪ྛࠎ 61 ࡢỈᇦෆ␗࡞ࡿᙧ࡛ᡂ❧ࡍࡿᣦࡉࢀ࡚࠸ࡿ㸦Bozek and Rahel, 1991㸧 㸬ᮏ ㄪᰝỈᇦࡢ࣐ࢦᾋୖ⛶㨶ࡢᚤ⏕ᜥሙᡤ᮲௳㸦ᐃⅬࡢỈ῝ὶ㏿㸧ࡶ㸪ὶୗ ⛣ືࡀ᥎ᐃࡉࢀࡓ 2009 ᖺࢆ㝖ࡃ㸪 ୖὶᇦ࠾ࡅࡿ࣐ࢦࡸࢧࢡ࣐ࣛࢫࡢᾋୖ ⛶㨶㸦ྡ㉺㸪1988㸹Nagata and Yanai, 2002㸧ྠᵝ࡛࠶ࡗࡓ㸦Table 4-2㸧 㸬 ᪤ሗ㸦Bozek and Rahel, 1991㸹Inoue and Nakano, 1999㸧ࡀ♧ࡍࡼ࠺㸪ࢧࢣ⛉㨶 㢮ࡢᾋୖ⛶㨶ࡗ࡚ዲ㐺࡞ᚤ⏕ᜥሙᡤࡀࠕ㏿࠸ὶࢀ㏆࠸⦆ὶ㒊ศ࡛ࠖ࠶ࡿ ࡍࡿ㸪ὶ㊰༢ࢫࢣ࣮ࣝ࠾ࡅࡿୖ࣭ୗὶ㛫ࡢ⏕ᜥሙᡤ⏝ࡢ㐪࠸ࡣ㸪ዲ 㐺࡞ᚤ⏕ᜥሙᡤࡢ㓄⨨ࡀ୧⪅࡛␗࡞ࡿࡓࡵ⏕ࡌࡿ⪃࠼ࡽࢀࡿ㸬ࡘࡲࡾ㸪ୖ ὶࡽୗὶࡅ࡚ࡣ㸪Ỉ㠃ᖜࡢቑຍకࡗ࡚ᓊࡽὶᚰ㒊ࡲ࡛ࡢ㊥㞳ࡀቑຍ ࡍࡿ㸪Ἑᗋ໙㓄ࡢపୗక࠸ῡࡢᘏ㛗ࡀయ༨ࡵࡿẚ⋡ࡀቑຍࡋ㸪᪩℩ ᖹ℩࡛ࡣῶᑡࡍࡿ㸦ⴛሙ㸪2003a㸧㸬ࡑࡢࡓࡵ㸪ᓊἢ࠸ከࡃぢࡽࢀࡿὸ ࡃὶ㏿ࡢᑠࡉ࡞ᚤ⏕ᜥሙᡤࡢೃ⿵ᆅࡢ࠺ࡕ㸪㏿࠸ὶࢀ㏆࠸ሙᡤࡣ㸪ୖὶഃ࡛ ࡣ㐢ᅾࡍࡿࡀୗὶഃ࡛ࡣ᪩℩࿘㎶ᒁᅾࡍࡿணࡉࢀࡿ㸬ࡋࡓࡀࡗ࡚㸪࣐ ࢦᾋୖ⛶㨶ࡢὶ㊰༢ࢱࣉ࠾ࡅࡿୖ࣭ୗὶ㛫ࡢ⏕ᜥሙᡤ⏝ࡢ㐪࠸ࡣ㸪ὶ ⛬ἢࡗ࡚ኚࡍࡿࡁ࡞✵㛫ࢫࢣ࣮ࣝࡢ⎔ቃ໙㓄⏤᮶ࡋ㸪᪩℩࿘㎶ࡢὶ㊰ ༢ࢱࣉࡢ⏝ࡣ㸪࣐ࢦ⏕ᜥᇦୗὶഃ≉᭷ࡢ⌧㇟࡛࠶ࡿ⪃࠼ࡽࢀࡿ㸬 ௨ୖࡢࡇࡽ㸪࣐ࢦ⏕ᜥᇦୗὶഃ࡛ᾋୖ⛶㨶ࡢಖࡸቑṪࢆᅗࡿࡓࡵ ࡣ㸪ࡲࡎ㸪℩࣭ῡᵓ㐀ࡢಖࡀᚲせ⪃࠼ࡽࢀࡿ㸬࣐ࢦ⏕ᜥᇦୗὶഃࢆྵࡴ ᓙ㇂ࡽᡪ≧ᆅࡅ࡚ࡣ㸪ୖὶഃࡢ◁㜵ࢲ࣒ࡼࡿᅵ◁౪⤥ࡢไ㝈㸦㧗ᶫ㸪 2000㸧㸪ㆤᓊࡸᗋᅛᕤ➼ࡼࡿἙ㐨ࡢᅛᐃ㸦Yoshimura et al., 2005㸧➼ࡼࡾ㸪 ℩࣭ῡᵓ㐀ࡀᾘኻࡍࡿ㸬℩࣭ῡᵓ㐀ࡣᾋୖ⛶㨶ࡢ⏕ᜥሙᡤࡔࡅ࡛࡞ࡃ㸪⦾Ṫࢆ ྵࡴࢧࢣ⛉㨶㢮ࡢ⏕ά⯡ࢆつᐃࡍࡿࡇࡽ㸦๓ᕝ㸪2000㸧㸪ࡍ࡚ࡢಖᑐ ⟇ࡢᇶᮏࡍࡁ࡛࠶ࡿ㸬ḟ㸪᪤ሗ㸦Foldvik et al., 2010㸹Einum et al., 2011, 2012㸧 ࡛᥎ ࡉࢀࡿࡼ࠺㸪ὶୗ⛣ືࡼࡾᾋୖ⛶㨶ࡢ㣚㣹ࡸ⿕㣗ࣜࢫࢡࡀ㧗ࡲࡿ࡞ ࡽࡤ㸪ὶୗࢆᅇ㑊ࡍࡿᚲせࡀ࠶ࡿ⪃࠼ࡽࢀࡿ㸬࢝ࢵࢺࢫ࣮ࣟࢺࢺࣛ࢘ࢺᾋୖ ⛶㨶ࡢὶୗࡣዲ㐺࡞ᚤ⏕ᜥሙᡤࡢ㊊ࡼࡾ⏕ࡌࡿࡉࢀ㸦Moore and Gregory㸪 1988㸧 㸪ᮏㄪᰝỈᇦ࡛ࡶྠᵝࡢὶୗࡀ᥎ᐃࡉࢀࡓ㸦Fig. 4-4㸧㸬ࡋࡓࡀࡗ࡚࣐ࢦ ⏕ᜥᇦୗὶഃ࡛ࡣ㸪᪩℩࿘㎶ࡢὶ㊰༢ࢱࣉෆࡢὸ࠸ሙᡤ㸪ࡁ࡞ὶ㏿ࢆ 62 పῶࡍࡿ▼ࡸ᳜≀᳜⏕ࢆᩚഛࡋࠕ㏿࠸ὶࢀ㏆࠸⦆ὶ㒊ศ㸦Bozek and Rahel, 1991㸹Inoue and Nakano, 1999㸧 ࠖࢆ⏕ᡂࡍࡿࡇࡀ㔜せ࠸࠼ࡿ㸬᭱ᚋ㸪ࢧࢣ ⛉㨶㢮ࡢᾋୖ⛶㨶ࡢὶୗ⛣ືࡣᐦᗘ౫Ꮡⓗ⏕ࡌࡿࡇࡽ㸦Nagata, 2002㸹 Einum et al., 2011, 2012㸧 㸪⏕ᜥሙᡤࡢ⎔ቃᐜຊಶయ⩌ᐦᗘࡢࣂࣛࣥࢫࢆ⪃៖ ࡍࡿᚲせࡀ࠶ࡿ⪃࠼ࡽࢀࡿ㸬≉㸪ேᕤ⏘༸ሙࡢ㐀ᡂ㸦୰ᮧ㸪1999㸧 㸪Ⓨ║༸ ᨺὶ㸦Ṋᒃ㸪2009㸧 㸪ぶ㨶ᨺὶ㸦ᚫཎ㸪2010㸧࠸ࡗࡓ㸪ᾋୖᚋࡢ⛶㨶ࡢ⏕ ⫱ࢆ⮬↛⎔ቃࡺࡔࡡࡿᡭἲ࡛ࡣ㸪㔝⏕ࡶࡋࡃࡣ㔝⏕㏆࠸≧ែࡢ⛶㨶ࡀ☜ᐇ ࡘ㔞ᚓࡽࢀࡿࡀ㸪ࡑࢀぢྜࡗࡓᾋୖ⛶㨶ࡢ⏕ᜥሙᡤࡀᚲせ࡞ࡿண ࡉࢀࡿ㸬 ᚋࡣ㸪 ⛶㨶ࡢῧຍ㔞ዲ㐺࡞ᚤ⏕ᜥሙᡤࡢᏑᅾ㔞ࡢ㛵ಀࡘ࠸࡚㸪 ᐃ㔞ⓗ࡞ホ౯ࢆ⾜࠺ᚲせࡀ࠶ࡿ㸬 63 Ûޗ1 N Fukuoka Prefecture Survey area Ogata River Seto Inland Sea Oita Prefecture 1 km Ono River Kumamoto Prefecture Ûޗ1 1 Kohbaru Stream Mt. Koshiki Hakiai Stream Mt. Sobo 3 5 Û(ޗ Û(ޗ 7 9 11 N 13 15 riffle man-made dam waterfall boundary of section Ogata River 17 47 45 43 19 21 23 Flo w 25 27 41 39 29 100 m 31 33 35 49 51 53 55 37 Fig. 4-1. Map and location of study sites in the Ogata River, northeastern Kyushu Island, Japan. 64 Fig. 4-2. See the next page for figure caption because of its length. 65 Fig. 4-2. Longitudinal distribution of newly emerged fry of red-spotted masu salmon in a 1,400 m stretch of the Ogata River based on 11 weekly snorkeling surveys from 10 January to 20 March in 2007. Fry were counted by divers moving upstream. Open circles represent the number of fry observed along both banks. Asterisks show the locations of spawning redds. Other notations are the same as those in Fig. 4-1. 66 n=1 n = 5 (17 Jan.) * n = 10 n = 100 * * n = 70 (15 Feb.) n = 26 (11 Mar.) Fig. 4-3. Longitudinal distribution of newly emerged fry of amago in a 1,400 m stretch of the Ogata River based on three snorkeling surveys each on 17 January, 15 February, and 11 March in 2008. Notations are the same as those in Fig. 4-2. 67 Fig. 4-4. Longitudinal distribution of newly emerged fry of amago in a 1,400 m stretch of the Ogata River based on five snorkeling surveys each on 14 January, 10 and15 February, 2 and17 March in 2009. Notations are the same as those in Figs. 4-2. 68 Table 4-1. Environmental characteristics of 56 sections (25 m long each) surveyed in the Ogata River. B/H: Width-depth ratio; DFR: distance from spawning redds. Environmental variables Depth (m) Right Left Velocity (cm/s) Right Left Vegitation cover (%) Right Left Wetted width (m) * Maximum depth (m) * B/H* DFR (m) 2007 2008 2009 mean ± sd 0.64 ± 0.77 0.79 ± 0.94 21.8 ± 29.7 10.8 ± 16.2 0.05 ± 0.12 0.22 ± 0.4 12.7 ± 6.01 1.65 ± 1.09 12.7 ± 10.9 213 ± 163 222 ± 157 235 ± 175 Range 0.05 – 3.42 0.05 – 3.84 0.06 – 134 0.06 – 89.1 0 – 0.68 0 – 1.0 2.63 – 32 0.29 – 4.55 1.12 – 59.5 0 – 600 0 – 600 0 – 650 *Wetted width and maximum depth were not used for generalized linear mixed model due to multicollinearity but used for calculating the width-depth ratios (B/H). 69 Table 4-2. Water depth at focal points of newly emerged fry of red-spotted masu salmon in the Ogata River from January to March in 2008 and 2009. Date 2008 17 Jan. 15 Feb. 11 Mar. 2009 14 Jan. 10 Feb. 15 Feb. 2 Mar. 17 Mar. mean ± sd (m) 0.21 ± 0.13 0.20 ± 0.11 0.29 ± 0.14 0.48 ± 0.77 0.50 ± 0.49 0.30 ± 0.29 0.38 ± 0.28 0.37 ± 0.30 70 Range (m) 0.08 – 0.42 0.07 – 0.60 0.07 – 0.55 0.05 – 4.00 0.04 – 4.00 0.03 – 4.00 0.08 – 2.00 0.10 – 2.00 71 3665.8 3 Intercept 0.72** (2.76) 0.72** (2.76) 0.73** (2.77) CU-G 0.28 (0.94) 0.16 (0.55) 0.27 (0.89) CU-GR CU-R 0.91*** 0.41 (4.76) (1.81) 0.82*** 0.26 (4.49) (1.28) 0.90*** 0.41 (4.57) (1.81) Significans level: *P < 0.05; **P < 0.01: ***P < 0.001 3664.3 2 Model AIC 1 3663.9 DFR: distance from spawning redds. CU-RP CU-PG Depth 1.14*** 0.31 -0.34*** (6.89) (1.43) (-3.43) 1.05*** 0.25 -0.37*** (6.75) (1.18) (-3.68) 1.13*** 0.30 -0.34*** (6.74) (1.40) (-3.43) 0.004 (-1.54) Velocity 0.004 (-1.51) -0.05 (-0.27) Cover B/H DFR 0.03*** -0.002*** (5.00) (-3.85) 0.03*** -0.002*** (5.23) (-3.95) 0.03*** -0.002*** (4.89) (-3.84) CU: channel unit; G: glide; GR: glide-riffle; R: riffle; RP: riffle-pool; PG: pool-glide; Cover: vegetation cover; B/H: width-depth ratio; are shown in an ascending order of AIC. Bold values indicate coefficient estimates where the 95% confidence intervals do not include zero. emerged fry of red-spotted masu salmon in the Ogata River from January to March in 2007. The best model and the models with 'AIC < 2 Table 4-3. Coefficient estimates and Wald statistics (in parentheses) from the generalized linear mixed model fit to abundance of newly 72 339.6 340.7 340.8 341.0 341.0 341.2 341.3 2 3 4 5 6 7 8 Intercept -0.36 (-0.85) -0.12 (-0.26) -0.28 (-0.65) 0.07 (0.14) -0.86 (-1.44) -0.50 (-0.76) -0.08 (-0.18) -0.30 (-0.67) 0.92 (1.10) 0.86 (1.03) CU-G 1.23* (2.07) 1.57* (2.46) CU-GR 1.09 (1.81) 1.48* (2.24) CU-R Significans level: *P < 0.05; **P < 0.01: ***P < 0.001 AIC 339.5 Model 1 Notations are the same as those in Table 4-3. 1.12* (2.34) 1.14* (2.39) CU-RP -0.08 (-0.10) -0.03 (-0.03) CU-PG -0.16 (-0.65) -0.22 (-0.87) Depth -0.011 (-1.47) -0.016* (-1.98) -0.020* (-2.25) -0.008 (-1.14) -0.009 (-1.26) Velocity -0.22 (-0.44) -0.47 (-0.91) Cover -0.02 (-1.34) B/H DFR -0.006*** (-3.92) -0.006*** (-4.13) -0.005*** (-3.67) -0.007*** (-4.22) -0.005** (-2.91) -0.005** (-3.12) -0.006*** (-3.81) -0.006*** (-3.91) are shown in an ascending order of AIC. Bold values indicate coefficient estimates where the 95% confidence intervals do not include zero. emerged fry of red-spotted masu salmon in the Ogata River from January to March in 2008. The best model and the models with 'AIC < 2 Table 4-4. Coefficient estimates and Wald statistics (in parentheses) from the generalized linear mixed model fit to abundance of newly 73 Intercept 1.98*** (7.16) 2.09*** (7.16) 2.01*** (7.18) 2.10*** (7.17) CU-G -1.28* (-2.14) -1.24* (-2.08) -1.29* (-2.17) -1.26* (-2.11) CU-GR 0.28 (1.09) 0.47 (1.61) 0.27 (1.07) 0.45 (1.53) CU-R 0.49 (1.91) 0.66* (2.30) 0.53* (2.02) 067* (2.32) Significans level: *P < 0.05; **P < 0.01: ***P < 0.001 1697.1 1696.5 3 4 1695.4 2 Model AIC 1 1695.1 Notations are the same as those in Table 4-3. CU-RP 0.13 (0.65) 0.20 (0.95) 0.13 (0.61) 0.19 (0.89) CU-PG -0.34 (-1.34) -0.24 (-0.89) -0.32 (-1.24) -0.23 (-0.87) Depth -0.54*** (-4.40) -0.61*** (-4.45) -0.55*** (-4.45) -0.62*** (-4.47) Velocity -0.008* (-2.29) -0.008* (-2.39) -0.008* (-2.39) -0.009* (-2.44) -0.20 (-0.78) -0.12 (-0.48) Cover -0.01 (-1.14) -0.01 (-1.29) B/H DFR -0.004*** (-6.95) -0.003*** (-6.36) -0.004*** (-6.86) -0.003*** (-6.34) are shown in an ascending order of AIC. Bold values indicate coefficient estimates where the 95% confidence intervals do not include zero. emerged fry of red-spotted masu salmon in the Ogata River from January to March in 2009. The best model and the models with 'AIC < 2 Table 4-5. Coefficient estimates and Wald statistics (in parentheses) from the generalized linear mixed model fit to abundance of newly ➨ 5 ❶ ࣐ࢦಶయ⩌ࡢ㑇ఏⓗᵓ㐀 ࣐ࢦࡣࢧࢣ⛉ᒓࡍࡿ᪥ᮏᅛ᭷ࡢள✀࡛࠶ࡾ㸪ᮏᕞኴᖹὒᓊ㸪ᅄᅜᇦ㸪 ᕞࡢ℩ᡞෆᾏὶධἙᕝ⮬↛ศᕸࡍࡿ㸦ᓥ㸪1957㸧 㸬ᮏள✀ࡢἙᕝᆺಶయ⩌ ࡣ㸪ࣃ࣮࣐࣮ࢡ㸪㯮Ⅼ㸪ᮒⅬࢆḞࡃ↓ᩬᆺࡢ࣓࣡ࡀᏑᅾࡍࡿࡇࡀ▱ࡽࢀ ࡚࠸ࡿ㸦Kimura and Nakamura, 1961㸧㸬ࢃࡀᅜ࡛ࡣ㸪1970 ᖺ௦௨㝆㸪㈨※ቑṪࡢ ࡓࡵࡢ㣴Ṫ㨶ࡢᨺὶࡀ┒ࢇ⾜ࢃࢀ࡚ࡁࡓ㸦❧ᕝ࣭ᮏⲮ㸪1976㸧㸬ࡑࡢࡓࡵ⌧ᅾ ࡛ࡣ㸪㑇ఏⓗ⣧⢋࡞࣐ࢦಶయ⩌ࡣࡸࢲ࣒ࡼࡗ࡚㝸㞳ࡉࢀࡓ※ὶ㒊ࡔࡅ ⏕ᜥࡍࡿࡇࡀሗ࿌ࡉࢀ࡚࠸ࡿ㸦Kawamura et al., 2007㸧㸬࣐ࢦࡢศᕸ༡す ➃࡛࠶ࡿᕞ࡛ࡣ㸪ࡰࡍ࡚ࡢ࣐ࢦࡀἙᕝᆺಶయ⩌ࡋ࡚Ἑᕝୖὶᇦ⏕ ᜥࡋ࡚࠸ࡿ㸦Kimura, 1989㸧㸬ࡑࢀࡽἙᕝᆺಶయ⩌ࡣἙᕝෆ࡛ࡁࡃ⛣ືࡋ࡞࠸ ࡇࡀ▱ࡽࢀ࡚࠾ࡾ㸦Nakano et al., 1990㸧 㸪ࡑࡢ㧗࠸ᐃఫᛶࡣಶయ⩌㛫ࡢ㑇ఏⓗ ὶࢆไ㝈ࡍࡿࡶ㸦Yamamoto et al., 2004㸧 㸪ᒁᡤ㐺ᛂࢆࡶࡓࡽࡍண ࡉࢀࡿ㸦Garcia de Leaniz et al., 2007㸹Kikko et al., 2008㸹Drinan et al., 2012㸧㸬㣴 Ṫ㨶ࡢᨺὶࡣ㸪㔝⏕ಶయ⩌ࡢ୰㐺ᛂⓗ࡛࡞࠸㑇ఏᏊࢆὶධࡉࡏ㸪␗⣔㓄ᙅ ໃ㸦Miller et al., 2004㸹Tymchuk et al., 2007㸧ࡸᒁᡤ㐺ᛂࡢᔂቯ㸦Garcia de Leaniz et al., 2007㸧ࢆࡶࡓࡽࡍྍ⬟ᛶࡀ࠶ࡿ㸬ࡋࡓࡀࡗ࡚㸪ᕞࡢ࣐ࢦ㛵ࡋ࡚ࡣ㸪 ண㜵ⓗ࡞ࣜࢫࢡ⟶⌮ࡢほⅬࡽᅾ᮶ಶయ⩌ࢆ᥈ࡋฟࡋ࡚ಖㆤࡍࡿᚲせࡀ࠶ࡿ㸬 ㏆ᖺࡢ◊✲࡛ࡣ㸪࣐ࢦࢆྵࡴἙᕝᛶࢧࢣ⛉㨶㢮ࡢᅾ᮶ಶయ⩌ࡣ㸪࠾ࡶᨺ ὶᒚṔᨺὶ㨶ࡢୖὶ᪉ྥࡢ⛣ືไ㝈㸦ࡸࢲ࣒㸧ࡢ⨨㸦୰ᮧ㸪2001㸧 㸪࠾ ࡼࡧᨺὶࡀ࡞ࡉࢀ࡚࠸࡞࠸㞄᥋ࡋࡓ」ᩘࡢಶయ⩌㛫࠾ࡅࡿ㑇ఏᏊᆺࡢ୍㈏ᛶ ࡸ㑇ఏᏊᆺ㢖ᗘࡢẚ㍑ࡽ᥎ᐃ࡛ࡁࡿࡇࡀሗ࿌ࡉࢀ࡚࠸ࡿ㸦Kawamura et al., 2007㸹Kubota et al., 2007㸹Sato et al., 2010㸧 㸬ศ┴ࡢከࡃࡢ℩ᡞෆᾏὶධἙᕝ ୖὶ㒊ࡣ࣐ࢦࡀ⏕ᜥࡋ࡚࠸ࡿࡀ㸪ᨺὶࡢᙳ㡪ࢆཷࡅ࡚࠸࡞࠸⪃࠼ࡽࢀࡿ ࣐ࢦಶయ⩌ࡣ㸪㔝ᕝỈ⣔ࡢ୍ᨭὶ㸪⚄ཎᕝࡢ᭱ୖὶ㒊ࡔࡅ⏕ᜥࡍࡿྍ⬟ ᛶࡀ♧ࡉࢀ࡚࠸ࡿ㸦Fig. 5-1㸹ᚨගಇ㹙ศ┴ᾏὒỈ⏘◊✲ࢭࣥࢱ࣮ෆỈ㠃◊ ✲ᡤ㸪ᮍⓎ⾲㸪ෆỈ㠃⁺ᴗ༠ྠ⤌ྜᑐࡍࡿᨺὶᒚṔࡢ⪺ࡁྲྀࡾㄪᰝ㸪1999㸧 㸬 ࡇࡢỈᇦࡣ⁺ᴗ࣭㐟⁺ࡘ࠸࡚㛤ᨺࡉࢀ࡚࠸ࡿࡀ㸪⚄ཎᕝࡢ㞄᥋ᨭὶ࡛࠶ࡿἼ 74 ᮌྜᕝࡣ㸪࣓࣡࠾ࡼࡧ࣐ࢦಖㆤࡢࡓࡵࡢ⚗⁺༊ࡀタࡅࡽࢀ࡚࠸ࡿ㸬࣡ ࣓ࡣᙜึ㸪Ἴᮌྜᕝࡢ࣓ࣥࣀࢶࣛ㇂⏕ᜥࡍࡿ᪂✀ࡢࢧࢣ⛉㨶㢮ࡋ࡚ሗ࿌ࡉ ࢀࡓ㸦Kimura and Nakamura, 1961㸧㸬ࡑࡢᚋ㸪ᮏᕞᅄᅜࡽ 6 ࡘࡢ࣓࣡ಶయ ⩌ࡀ㸪㏻ᖖࡢ࣐ࢦ࠶ࡿ࠸ࡣ࣐࣓ࣖࡶⓎぢࡉࢀ࡚࠸ࡿ㸦Kimura 1989㸹 Kano et al., 2010㸧㸬⌧ᅾ㸪࣓࣡ࡣ࣐ࢦࡢ㓄ᐇ㦂㸦ᒣෆ㸪1982㸹ᚨග㸪2003㸧 ᇶ࡙ࡁ㸪ྠᡤⓗ⏕ᜥࡍࡿ㏻ᖖࡢ࣐ࢦ࠶ࡿ࠸ࡣ࣐࣓ࣖࡢ↓ᩬᆺ✺↛ኚ␗య ⪃࠼ࡽࢀ࡚࠸ࡿ㸦Kimura, 1989㸹Kano et al., 2006㸧 㸬Ἴᮌྜᕝ࣓ࣥࣀࢶࣛ㇂ࡢ ࣓࣡ಶయ⩌ࡣ㸪㞄᥋ࡍࡿࡲࢇࡾࡻ࠺㇂ࡢ࣐ࢦࡶ 1966 ᖺศ┴ࡢኳ ↛グᛕ≀ᣦᐃࡉࢀ㸪1970 ᖺࡽࡣ⏕ᜥᆅࡀ⚗⁺࡞ࡗ࡚࠸ࡿ㸬ࡋࡋ㸪ࡲࢇ ࡾࡻ࠺㇂࡛ࡣ 1990 ᖺ௦㐟⁺⪅ࡀ㠀බᘧ࣐ࢦࢆᨺὶࡋࡓࡢሗࡀ࠶ࡿ 㸦➉⏣ᕷෆࡢᆅඖఫẸࡽࡢሗ㸧 㸬⚄ཎᕝ⏕ᜥࡍࡿ࣐ࢦࡘ࠸࡚㸪ࡇࢀࡲ ࡛」ᩘࡢ◊✲㸦ᮌᮏ㸪2013㸪2015㸧ࡀ⾜ࢃࢀ࡚ࡁࡓࡀ㸪࣐ࢦࡢ㑇ఏⓗᵓ 㐀ࡸ࣓࣡ࡢ㑇ఏⓗ㛵ಀࡘ࠸࡚ࡣሗࡀ࡞࠸㸬 ᮏ◊✲࡛ࡣ㸪㔝ᕝỈ⣔⚄ཎᕝࡢᅾ᮶࣐ࢦಶయ⩌ࢆ≉ᐃࡍࡿࡓࡵ㸪⚄ཎᕝ Ἴᮌྜᕝ⏕ᜥࡍࡿ 3 ࡘࡢ࣐ࢦಶయ⩌㸦࣓࣡ 1 ಶయ⩌ࢆྵࡴ㸧ࡢ㑇ఏⓗ ᵓ㐀ࢆ mtDNA ࡼࡾゎᯒࡋࡓ㸬ࡲࡓ㸪ᅾ᮶ಶయ⩌ࡢ᥎ᐃ⤖ᯝࢆ᳨ドࡍࡿࡓࡵ㸪 ⚄ཎᕝయ࡛᥇㞟ࡋࡓ⛶㨶ࡲࢇࡾࡻ࠺㇂ᨺὶࡉࢀࡓணࡉࢀࡿ㣴㨶ሙࡢ ⣔⤫ࡘ࠸࡚ࡶㄪᰝࡋࡓ㸬ࡇࢀࡽࡢ⤖ᯝᇶ࡙ࡁ㸪ᙜヱỈᇦࡢᅾ᮶࣐ࢦಶయ ⩌ࡢಖᡭἲࢆᥦࡋࡓ㸬 ᮦᩱ᪉ἲ ㄪᰝሙᡤ ᮏ◊✲࡛ࡣ㸪⥴᪉ᕝ⚄ཎᕝࡢ࣐ࢦ⏕ᜥᇦ㸦ᮌᮏ㸪2013㸧࡛࠶ࡿ⥴᪉ ᕝ༑ゅᕝࡢྜὶⅬࡽྜ┠ࡲ࡛ࡢ༊㛫ࢆㄪᰝᑐ㇟Ỉᇦࡋ㸪㨶㢮ࡢ⛣ື ไ㝈࡞ࡿ ࡸࢲ ࣒ ࡢ⨨ ᇶ࡙ ࠸࡚ 7 ࡘࡢ㐃⥆ࡋࡓㄪ ᰝ ༊ࢆタ ᐃࡋࡓ 㸦Section 1–7㸪Fig. 5-1㸧 㸬⚄ཎᕝ⥴᪉ᕝ࡛ࡣ㸪ࡇࢀࡲ࡛⚗⁺ᥐ⨨ࡀࡽࢀࡓࡇ ࡣ࡞࠸㸬⁺༠ࡼࡿබᘧ࡞ᨺὶࡣ㸪⚄ཎᕝᨭὶἼᮌྜᕝࡢྜὶⅬࡼࡾୗὶ 75 ഃ࡛ࡣẖᖺ⾜ࢃࢀ࡚࠸ࡿࡀ㸪⚄ཎᕝࡢ᭱ୖὶ㒊㸦Section 7㸪Fig. 5-1㸧࡛ࡣᐇ ࡉࢀ࡚࠸࡞࠸㸦㔝ᕝ⁺ᴗ༠ྠ⤌ྜࡽࡢሗ㸧 㸬ࡲࡓ㸪ᮏㄪᰝỈᇦࡼࡾୖὶ ࡣ㨶㢮ࡀ⏕ᜥࡋ࡚࠸࡞࠸㸦ᮌᮧ㸪1959㸹ᮌᮏ㸪ᮍⓎ⾲㸧 㸬 ࣓ࣥࣀࢶࣛ㇂ࡲࢇࡾࡻ࠺㇂ࡣࡶἼᮌྜᕝࡢᨭὶ࡛࠶ࡾ㸦Fig. 5-1㸧㸪Ἴ ᮌྜᕝࡢⓑỈᶫࡽ࣓ࣥࣀࢶࣛ㇂᭱ୖὶ㒊ࡢ◁㜵ࢲ࣒ࡲ࡛ࡢ༊㛫㸪࠾ࡼࡧ࣓ࣥ ࣀࢶࣛ㇂ࡲࢇࡾࡻ࠺㇂ࡢྜὶⅬࡽࣄ࣮ࣂࢳࡢࡲ࡛ࡢ༊㛫ࡣ 1970 ᖺࡽ ⚗⁺࡞ࡗ࡚࠸ࡿ㸬࣓ࣥࣀࢶࣛ㇂ࡣ࣓࣡ࡀ⏕ᜥࡋ࡚࠾ࡾ㸪ࡑࡢ⏕ᜥୖ㝈ࡣ 㙚ῡࡢ㸦ⴠᕪ 16 m㸧ࡽ 400 m ୖὶࡢῡࡲ࡛࡛࠶ࡗࡓࡀ㸪1998 ᖺ 5 ᭶⏕ ᜥᇦୖὶ➃ࡢ࣓࣡ᾋୖ⛶㨶 47 ಶయࡀ 520 m ୖὶ⛣᳜ᨺὶࡉࢀ⏕ᜥᇦࡀᣑ ࡉࢀࡓ㸦ᬒᖹ࣭ᚨග㸪2003㸧 㸬࣓ࣥࣀࢶࣛ㇂ࡢ㙚ῡࡢࡼࡾୗὶഃ࡛ࡣ㸪࣡ ࣓ࡣ࣐ࢦඹᏑࡋ࡚࠾ࡾ㸪࣓ࣥࣀࢶࣛ㇂௨እࡢἼᮌྜᕝయࡣ࣐ࢦࡀ⏕ ᜥࡋ࡚࠸ࡿ㸦㏆⸨࣭➉ୗ㸪2005㸧 㸬⪺ࡁྲྀࡾㄪᰝ࡛ࡣ㸪ࡲࢇࡾࡻ࠺㇂ࡢࣄ࣮ࣂࢳ ࡢࡢୖὶഃ 1990 ᖺ௦㣴Ṫ㨶ࡢ㠀බᘧ࡞ᨺὶࡀ࡞ࡉࢀࡓࡢሗࡀᚓࡽ ࢀ࡚࠸ࡿ㸦➉⏣ᕷෆࡢᆅඖఫẸࡽࡢሗ㸧㸬 ౪ヨ㨶ࡢ᥇㞟 ⚄ཎᕝ࡛ࡣ㸪2005 ᖺ 9 ᭶᭱ୖὶ㒊ࡢ Section 7㸦Fig. 5-1㸧࠾࠸࡚࣐ࢦ 63 ಶయ㸦KOH-7A ⾲グ㸪௨ୗྠᵝ㸧ࢆ㣵㔮ࡾࡼࡾ᥇ᤕࡋࡓ㸦Table 5-1㸧 㸬 Section 7 ࡢὶ⛬ࡣ⣙ 1 km ࡛࠶ࡾ㸪୍ྜ┠㸦ⴠᕪ 16 m㸧ࡼࡗ࡚ୗὶഃㄪᰝ ༊ࡽ㝸㞳ࡉࢀ࡚࠸ࡿ㸬ࡇࡢㄪᰝ༊ࡢ㞟ᅋࢧࢬࡣ㸪2005 ᖺ 9 ᭶Ⅼ࡛ 500 ಶయ௨ୗ᥎ᐃࡉࢀࡓ㸦ᮌᮏ㸪ᮍⓎ⾲㸧 㸬ࡲࡓ㸪㔝ᕝ⁺ᴗ༠ྠ⤌ྜ ࡼࡿ࣐ࢦ⛶㨶ࡢᨺὶࡀ⾜ࢃࢀࡿ๓ࡢ 2004 ᖺ 2–3 ᭶ࡣ㸪Section 1–7 㸦Fig. 5-1㸧 ࠾࠸࡚࣐ࢦᾋୖ⛶㨶 121 ಶయ㸦KOH-1F–7F㸧ࢆࡓࡶ⥙࡛᥇㞟ࡋࡓ㸬ᘵ⛶ 㨶ࡢ᥇㞟ࢆྍ⬟࡞㝈ࡾᅇ㑊ࡍࡿࡓࡵ㸪ㄪᰝ༊ 1 km ࡘࡁ 10 ಶయ㸪୍ࡘࡢ⩌ࢀ ࡽࡣ 1 ಶయࢆ㝈ᗘࡋ࡚᥇㞟ࢆ⾜ࡗࡓ㸬 Ἴᮌྜᕝࡢ࣓ࣥࣀࢶࣛ㇂࡛ࡣ㸪2005 ᖺ 3–6 ᭶㙚ῡࡢࡼࡾୖὶഃࡢᮏ᮶ࡢ ⏕ᜥᇦ࡛࣓࣡ 10 ಶయࢆ㸪2008 ᖺ 2 ᭶⛣᳜ᨺὶ༊㛫ࡽ࣓࣡ 13 ಶయࢆ㣵 㔮ࡾࡼࡾ᥇ᤕࡋࡓ㸦ྜィ 23 ಶయ㸸MEN㸧 㸬᥇ᤕ༊㛫ࡢὶ⛬ࡣ⣙ 1 km ࡛㸪㙚 76 ῡࡢࡼࡾୗὶഃỈᇦ㝸㞳ࡉࢀ࡚࠸ࡿ㸬㙚ῡࡢࡼࡾୖὶഃࡢ࣡ ࣓ࡢ㞟ᅋࢧࢬࡣ㸪2005 ᖺ 6 ᭶ 2008 ᖺ 5 ᭶ࡶ⣙ 500 ಶయ᥎ᐃࡉࢀ ࡓ㸦ᬒᖹ㸪2009㸧 㸬 Ἴᮌྜᕝࡢࡲࢇࡾࡻ࠺㇂࡛ࡣ㸪 2007 ᖺ 3 ᭶࣓ࣥࣀࢶࣛ㇂ࡢྜὶⅬࡽ 200 m ୖὶࡢሖሐࡽࣄ࣮ࣂࢳࡢࡲ࡛ࡢ༊㛫࡛࣐ࢦ 20 ಶయ㸦MAN㸧ࢆ㣵㔮ࡾ ࡼࡾ᥇ᤕࡋࡓ㸬᥇ᤕ༊㛫ࡢὶ⛬㛗ࡣ⣙ 1 km ࡛㸪ࡲࢇࡾࡻ࠺㇂࣓ࣥࣀࢶࣛ ㇂ࡢྜὶⅬࡢୖὶഃ⨨ࡍࡿ◁㜵ࢲ࣒ࡼࡗ࡚ୗὶഃỈᇦࡽ㝸㞳ࡉࢀ࡚࠸ ࡿ㸬ࡲࢇࡾࡻ࠺㇂ࡢ࣐ࢦࡢ㞟ᅋࢧࢬ㛵ࡍࡿሗࡣ࡞࠸㸬 2008 ᖺ 5 ᭶ࡣ㸪 1990 ᖺ௦ࡲࢇࡾࡻ࠺㇂ᨺὶࡉࢀࡓ㣴Ṫ㨶ࢆ⏕⏘ࡋࡓ ᥎ᐃࡉࢀࡿศ┴ෆࡢ 1 㣴㨶ሙࡀಖᣢࡍࡿ࣐ࢦ⣔⤫㸦HAT-A㸧࣐࣓ࣖ⣔⤫ 㸦HAT-M㸧ࡽ㸪 ࡑࢀࡒࢀ 10 ಶయࢆ᥇ྲྀࡋࡓ㸬ࡇࡢ㣴㨶ሙ࡛ࡣ㸪 HAT-A HAT-M ࡢ 2 ⣔⤫ࡔࡅࡀᩬ⣠ᙧ㉁ࡢ⨾ࡋࡉ㸦ࣃ࣮࣐࣮ࢡ㸪ᑠ㯮Ⅼ㸪ᮒᑠⅬࡢᩘ㸪ᙧែ㸪 㓄⨨➼㸧ᇶ࡙࠸࡚ 20 ᖺ௨ୖࢃࡓࡾ㑅ᢤ⫱✀ࡉࢀ࡚ࡁࡓ㸬࣐࣓ࣖࡣᕝỈ⣔ 㸦ศ┴ࡢ᭱༡➃⨨ࡋኴᖹὒすഃὶධ㸧※ὶ㒊࡛᥇ᤕࡉࢀࡓಶయ⏤᮶ ࡍࡿࡀ㸪࣐ࢦࡢ⏤᮶ࡣ࡛᫂࠶ࡿ㸬 ⛶㨶௨እࡢ᥇ᤕ㨶ࡣ㸪㔮⋓ᚋ 2-ࣇ࢙ࣀ࢟ࢩ࢚ࢱࣀ࣮࡛ࣝ㯞㓉ࡋ࡚ᑿཫ㛗ࢆ ᐃࡋ㸪⬡㫅ࡲࡓࡣᑿ㫅ࡢ୍㒊ࢆษ㝖ࡋ࡚ 99.5%࢚ࢳࣝࣝࢥ࣮ࣝಖᏑࡍࡿ ࡶ㸪㯞㓉ࡽᅇࡋࡓࡢࡕ㔮⋓ሙᡤᨺὶࡋࡓ㸬⛶㨶ࡣ᥇㞟ᚋ◊✲ᐊ ᣢࡕᖐࡾ㸪㯞㓉ࡋ࡚ᑿཫ㛗ࢆ ᐃࡋࡓࡢࡕ㸪㨶యࢆ 99.5%࢚ࢳࣝࣝࢥ࣮ ࣝ୰ಖᏑࡋࡓ㸬ࡍ࡚ࡢヨᩱࡣศᯒࡲ࡛-30°C ࡛ಖᏑࡋࡓ㸬 㑇ఏᏊศᯒ ౪ヨヨᩱࡽ DNeasy Tissue Kit㸦Qiagen, Hilden, Germany㸧ࢆ⏝࠸࡚ DNA ࢆᢳฟࡋࡓ㸬Kawamura et al.㸦2007㸧ᚑ࠸㸪ࢩࢺࢡ࣮࣒ࣟ b ࡽ 12S rRNA 㸦tRNA-Thr㸪tRNA-Pro㸪ㄪ⠇㡿ᇦ࠾ࡼࡧ tRNA-Phe ࢆྵࡴ㸧ࡲ࡛ࡢ⣙ 2.0 kb ࡢ DNA ᩿∦㸦D-loop 㡿ᇦ㸧 㸪࠾ࡼࡧ 16S rRNA ࡽ tRNA-Met㸦tRNA-Leu㸪NADH subunit 1 㡿ᇦ㸪tRNA-Ile ࠾ࡼࡧ tRNA-Gln ࢆྵࡴ㸧ࡲ࡛ࡢ⣙ 2.0 kb ࡢ DNA ᩿ ∦㸦ND1 㡿ᇦ㸧ࢆ PCR ࡼࡾቑᖜࡋࡓ㸬ࣉ࣐࣮ࣛࢭࢵࢺࡣ㸪D-loop 㡿ᇦ 77 ࡘ࠸࡚ CB3R-/ ƍ-CAY ATY MAR CCM GAA TGR TAT TT-ƍ 12SAR-H ƍ-ATA RTR GGG TAT CTA ATC CYA GTT-ƍ㸧 㸦Palumbi et al., 1991㸧 㸪ND1 㡿ᇦ ࡘ࠸࡚ࡣ ƍ-ACC CCG CCT GTT TAC CAA AAA CAT-ƍ ƍ-GGT ATG AGC CCG ATA GCT TA-ƍ㸦Hall and Nawrocki, 1995㸧ࢆ⏝࠸ࡓ㸬PCR ᛂᾮࡣ 1 ಸ⃰ᗘࡢ GeneAmp PCR Gold Buffer㸪0.1 mM ࡢ dNTPs㸪2 mM ࡢ MgCl2㸪ȝ0 ࡢྛࣉ ࣐࣮ࣛ㸪0.5 U ࡢ AmpliTaq Gold DNA polymerase㸦Applied Biosystems㸧㸪ȝO ࡢ⁛⳦␃Ỉ࠾ࡼࡧ 10–50 ng ࡢᢳฟ DNA ࢆΰྜࡋࡓ ȝO ࡢ⣔ࡋ㸪PCR ᛂ᮲௳ࡣ㸪GeneAmp PCR System 2720㸦Applied Biosystems, Foster City, CA, USA㸧 ࢆ⏝࠸࡚㸪95ºC ࡛ 10 ศ㛫ࡢึᮇ⇕ኚᛶᚋ㸪⇕ኚᛶ ɋ ⛊㸪ࢽ࣮ࣜࣥࢢ 55ºC/30 ⛊㸪ఙ㛗ᛂ 72ºC/2 ศ㛫ࡢࢧࢡࣝࢆ 35 ᅇ⧞ࡾ㏉ࡋࡓࡢࡕ㸪᭱⤊ఙ㛗 ᛂࢆ 72ºC/10 ศ㛫ࡋࡓ㸬ᚓࡽࢀࡓ PCR ⏘≀ࡢ㛗ࡉࢆ㟁ẼὋືࡼࡾ☜ㄆࡋ ࡓࡢࡕ㸪௨ୗࡢ 16 ✀㢮ࡢไ㝈㓝⣲ࢆ⏝࠸࡚ PCR-RFLP ศᯒࢆ⾜ࡗࡓ㸹Aci I㸪 Afa I㸪Alu I㸪Bst UI㸪Cfr 13I㸪Dde I㸪Hae III㸪Hha I㸪Hinc II㸪Hinf I㸪Hsp 92II㸪 Mbo I㸪Msp I㸪Scr FI㸪Taq I ࠾ࡼࡧ Tsp EI㸦New England Biolabs, Beverly, MA, USA; Promega, Madison, WI, USA; Takara Shuzo, Kyoto, Japan; Toyobo, Tokyo, Japan㸧㸬ྛ ヨᩱࡢ PCR ⏘≀ 5–ȝO ᑐࡋ࡚㸪ྛไ㝈㓝⣲ࢆㄝ᫂᭩ᚑ࠸స⏝ࡉࡏࡓ㸬ᾘ ᚋࡢ DNA ᩿∦ࡣ㸪3% NuSieve 3㸸1 ࣮࢞ࣟࢫ㸦Lonza, Basel, Switzerland㸧 ࢆ⏝࠸࡚ 50 V/1.5 㛫ࡢ㟁ẼὋືࡼࡾศ㞳ࡋ㸪⮯࢚ࢳࢪ࣒࡛࢘ᰁⰍࡋࡓࡢ ࡕ㸪⣸እ⥺↷᫂ୗ࡛ᙳࡋࡓ㸬ྛไ㝈㓝⣲ࡈᚓࡽࢀࡓ DNA ᩿∦㛗ࡢࣃࢱ ࣮ࣥࢆࡍ࡚⤫ྜࡋ࡚ mtDNA ࣁࣉࣟࢱࣉࢆᐃ⩏ࡋࡓ㸬ࡉࡽ㸪PCR-RFLP ࡛ ᚓࡽࢀࡓࡍ࡚ࡢࣁࣉࣟࢱࣉࡘ࠸࡚ࢲࣞࢡࢺࢩ࣮ࢡ࢚ࣥࢫࡼࡾሷᇶ㓄 ิࢆỴᐃࡋࡓ㸬ྛ PCR ⏘≀ࡣ illustra ExoProStar㸦GE Healthcare㸧࡛⢭〇ࡋ㸪 PCR-RFLP ྠࡌࣉ࣐࣮ࣛࢭࢵࢺࢆ⏝࠸࡚ BigDye Terminator v3.1 Cycle Sequencing Kit ࡼࡾ ABI PRISM 3130xl Genetic Analyzer㸦Applied Biosystems㸧 ࡛ᛂࢆ⾜ࡗࡓ㸬ሷᇶ㓄ิࡣ CLUSTAL W㸦Thompson et al. 1994㸧ࢆ⏝࠸࡚ ࣓ࣛࣥࣥࢺࡋࡓࡢࡕ㸪Kawamura et al.,㸦2007㸧ࡀྂᗙᕝࡢ࣐ࢦࡘ࠸࡚ሗ ࿌ࡋࡓሷᇶ㓄ิ㸦GenBank accession nos. AB236731 for D-loop and AB236732 for ND1㸧ẚ㍑ࡋࡓ㸬ᮏ◊✲࡛ᚓࡽࢀࡓࣁࣉࣟࢱࣉ㸦Hap-1㸹ᚋ㏙㸧ࡣࢪ࣮ࣥࣂ 78 ࣥࢡⓏ㘓ࡋࡓ㸦GenBank under accession nos. LC055411 for D-loop and LC055412 for ND1㸧 㸬 㑇ఏᏛⓗゎᯒ ࡍ࡚ࡢ mtDNA ࣁࣉࣟࢱࣉ㛫ࡢ⣔⤫㛵ಀࢆㄪࡿࡓࡵ㸪ࢥࣥࣆ࣮ࣗࢱ࣮࣭ ࣉࣟࢢ࣒ࣛ TCS version 1.21㸦Clement et al. 2000㸧ࢆ⏝࠸࡚ࣁࣉࣟࢱࣉ࣭ࢿࢵ ࢺ࣮࣡ࢡ㸦95㸣⤫ィⓗ᭱⠇⣙ࢿࢵࢺ࣮࣡ࢡ㸧ࢆᵓ⠏ࡋࡓ㸬ྛ㞟ᅋෆࡢ㑇ఏⓗከ ᵝᛶࡢᣦᶆࡋ࡚㸪ࣁࣉࣟࢱࣉᩘ㸪ࣁࣉࣟࢱࣉከᵝᗘ㸦h㸹᰿㸪1990㸧 㸪 ሷᇶከᵝᗘ㸦ʌ㸹᰿㸪 1990㸧ࢆ ARLEQUIN version 3.5 㸦Excoffier and Lischer 2010㸧 ࢆ⏝࠸࡚ᐃ㔞ࡋࡓ㸬㞟ᅋ㛫ࡢ㑇ఏⓗከᵝᛶࡣ㸪3 ࡘࡢ㝸㞳㞟ᅋ㸦KOH-7A㸪 MEN ࠾ࡼࡧ MAN㸧ࡔࡅࡘ࠸࡚ㄪ㸪ᾋୖ⛶㨶ࡢ㞟ᅋ㸦KOH-1F–7F㸧ࡘ࠸ ࡚ࡣ family effect ࡀணࡉࢀࡓࡓࡵ㸪 ศᯒࢆᐇࡋ࡞ࡗࡓ 㸦Nielsen et al. 2001㸧 㸬 㑇ఏⓗከᵝᛶࡢᣦᶆࡋ࡚㸪3 ࡘࡢ㝸㞳㞟ᅋ㛫ࡢ ĭST㸦ĭ-statistics; Excoffier et al., 1992㸧ࢆ ARLEQUIN ࡼࡾ⟬ฟࡋ㸪10,000 ᅇࡢ୪᭰࠼᳨ᐃࡼࡾ᭷ពᛶࡢ ุᐃࢆ⾜ࡗࡓ㸬ࡲࡓ㸪㞟ᅋ㛫ࡢࣁࣉࣟࢱࣉ㢖ᗘࡢᕪ␗ࢆ ARLEQUIN ࡢṇ☜ ☜⋡᳨ᐃ㸦Raymond and Rousset㸪1995㸧ࡼࡾㄪ㸪10,000 ᅇࡢ de-memorization step ࢆྵࡴ 100,000 ᅇࡢ࣐ࣝࢥࣇ㐃㙐ࢧࣥࣉࣜࣥࢢࡼࡿ᭷ពᛶุᐃࢆ⾜ࡗࡓ㸬 ⤖ ᯝ ࣁࣉࣟࢱࣉࡢ≉ᚩศᕸ ࡍ࡚ࡢ౪ヨ㨶㸦247 ಶయ㸧ࡘ࠸࡚㸪mtDNA ࡢ D-loop 㡿ᇦ ND1 㡿ᇦࡢ PCR-RFLP ࡽᚓࡽࢀࡓከᆺࣃࢱ࣮ࣥࢆ⤫ྜࡋࡓ⤖ᯝ㸪11 ✀㢮ࡢࣁࣉࣟࢱࣉ ࡀᚓࡽࢀࡓ㸦௨ᚋ㸪Hap-1–11 ⾲グ㸹Table 5-2㸧 㸬D-loop 㡿ᇦ࡛ࡣ DNA 㛗ࡢከ ᆺࡀぢࡽࢀ㸪Hap-1–4 Hap-6–11 ࡣ 1,944-bp ࡛࠶ࡗࡓࡀ㸪Hap-5 ࡛ࡣㄪ⠇㡿ᇦ ࡢ 839 ␒┠ሷᇶࡢ⨨ 82-bp ࡢᤄධࡀ࠶ࡾ㸪DNA 㛗ࡣ 2,026-bp ࡛࠶ࡗࡓ㸬 ND1 㡿ᇦ࡛ࡣ㸪ᮏ◊✲᳨࡛ฟࡉࢀࡓࣁࣉࣟࢱࣉ㛫 DNA 㛗ከᆺࡣࡳࡽࢀ࡞ ࡗࡓ㸬ᚓࡽࢀࡓ 11 ✀㢮ࡢࣁࣉࣟࢱࣉ㛫ࡢሷᇶ㓄ิࡢᕪ␗ࡣ 0.53 ± 0.29 % 79 㸦ᖹᆒ್ ± ᶆ‽೫ᕪ㸧࡛࠶ࡗࡓ㸬D-loop 㡿ᇦࡣ㌿ࡀ 8 ᡤ㸪㌿ࡀ 2 ᡤ࠶ࡾ㸪ND1 㡿ᇦࡣ㌿ࡀ 9 ᡤ㸪㌿ࡀ 4 ᡤㄆࡵࡽࢀࡓ㸦Table 5-2㸧 㸬 Hap-1 ࡣ 3 ࡘࡢ㝸㞳㞟ᅋ࡛ඹ᭷ࡉࢀ㸪࣓ࣥࣀࢶࣛ㇂ࡢ࣓࣡ಶయ⩌㸦MEN㸧 ࡛ᅛᐃ㸦100%㸧㸪⚄ཎᕝࡢ࣐ࢦಶయ⩌㸦KOH-7A㸧࡛ඃ༨ࡋࡓࡀ㸦90.5%㸧㸪 ࡲࢇࡾࡻ࠺㇂ࡢ࣐ࢦಶయ⩌㸦MAN㸧࡛ࡣᑡ࡞ࡗࡓ㸦10.0%㸹Fig. 5-1㸪Table 5-3㸧㸬࣐ࢦಶయ⩌ KOH-7A ࡛ࡣ㸪Hap-4 ࡀᑡᩘ㸦9.5%㸧ࡳࡽࢀࡓ㸬୍᪉㸪 ࣐ࢦಶయ⩌ MAN ࡛ࡣ㣴㨶ሙࡢ࣐ࢦ⣔⤫㸦HAT-A㸧ࡳࡽࢀࡿ Hap-8 ࡀඃ༨ ࡋ㸦70.0%㸧㸪Hap-2㸦20.0%㸧ࡀࡇࢀḟ࠸ࡔ㸬Hap-1 ࡣࡍ࡚ࡢ⛶㨶ಶయ⩌ 㸦KOH-1F–7F㸧ࡶඹ᭷ࡉࢀ㸪ࡍ࡚ࡢࣁࣉࣟࢱࣉ୰࡛᭱ࡶ㧗࠸㢖ᗘ㸦62.8%㸧 ࢆ♧ࡍࡶ㸪᭱ୖὶ㒊ࡢಶయ⩌ KOH-7F ࡛ࡣ༢୍࡛࠶ࡗࡓ㸬Hap-1 ௨እࡢ ࣁࣉࣟࢱࣉࡢ࠺ࡕ 8 ✀㢮㸦Hap-2–3㸪Hap-5–10㸧ࡣ㸪㔝ᕝ⁺ᴗ༠ྠ⤌ྜࡼ ࡾẖᖺබᘧ࡞ᨺὶࡀ࡞ࡉࢀ࡚࠸ࡿㄪᰝỈᇦࡢ୰ὶࡽୗὶഃ㸦Section 1–4㸧ࡢ ಶయ⩌㸦KOH-1F–4F㸧ࡽᚓࡽࢀࡓࡀ㸦Table 5-3㸧 㸪Hap-4 ࡣୖὶഃ㸦Section 5– 6㸧ࡢಶయ⩌㸦KOH-5F–6F㸧ࡔࡅࡽᚓࡽࢀࡓ㸬㣴㨶ሙࡢ⣔⤫ࡢ࠺ࡕ㸪࣐ࢦ 㸦HAT-A㸧࡛ࡣ Hap-8 Hap-11 ࡀࡑࢀࡒࢀ 40.0% 60.0%᳨ฟࡉࢀࡓࡢᑐࡋ㸪 ࣐࣓ࣖ㸦HAT-M㸧࡛ࡣ Hap-7 ࡔࡅࡀᚓࡽࢀࡓ㸬Hap-7 Hap-8 ࡣ㸪ࡑࢀࡒࢀ⛶ 㨶ಶయ⩌㸦KOH-3F–4F㸧࡛ࡶ☜ㄆࡉࢀࡓ㸦Table 5-3㸧 㸬ࣁࣉࣟࢱࣉࢿࢵࢺ࣮࣡ ࢡ࡛ࡣ㸪⚄ཎᕝࡢ࣐ࢦಶయ⩌㸦KOH-7A㸧Ἴᮌྜᕝࡢ࣓࣡ಶయ⩌㸦MEN㸧 ࡛㢖ᗘࡢ㧗࠸ Hap-1 ࡣ㸪2 ࡘࡢ࣐ࢦࡢ㝸㞳ಶయ⩌㸦KOH-7A MAN㸧࡛☜ㄆ ࡉࢀࡓ 3 ࡘࡢࣁࣉࣟࢱࣉ㸦Hap-2㸪Hap-4㸪Hap-8㸧 7–9 ሷᇶࡢ⨨࡛㝸࡚ࡽ ࢀࡓ㸦Fig. 5-2㸧㸬㣴㨶ሙࡢ࣐ࢦ㸦HAT-A㸧ࡽᚓࡽࢀࡓ Hap-8 Hap-11 ࡢᕪ ࡣ 1 ሷᇶ࡛࠶ࡗࡓ㸬 㞟ᅋෆ࠾ࡼࡧ㞟ᅋ㛫ࡢ㑇ఏⓗከᵝᛶ ࣁࣉࣟࢱࣉከᵝᗘሷᇶከᵝᗘࡣࡑࢀࡒࢀ 0–0.714㸪0–0.0035 ࡢ⠊ᅖ࠶ ࡗࡓ㸦Table 5-3㸧㸬ࣁࣉࣟࢱࣉከᵝᗘࡣ㸪බᘧ࡞ᨺὶࡀ࠶ࡿỈᇦ㸦Sction 2–4㸧 ࡢ࣐ࢦᾋୖ⛶㨶ಶయ⩌㸦KOH-F2–F4㸧㠀බᘧ࡞ᨺὶࡀࢃࢀࡿࡲࢇࡾࡻ࠺ ㇂ࡢ࣐ࢦಶయ⩌㸦MAN㸧࡛㧗ࡗࡓ㸬ሷᇶከᵝᗘࡣ㸪ࡁࡃ␗࡞ࡿࣁࣉࣟࢱ 80 ࣉࢆྵࡴ⛶㨶ࡢಶయ⩌ 㸦KOH-3F ࠾ࡅࡿ Hap-5 ࡢࣁࣉࣟࢱࣉ㸪KOH-6F ࠾ࡅࡿ Hap-4 ࡢࣁࣉࣟࢱࣉ㸧࡛㧗ࡗࡓ㸬3 ࡘࡢ㝸㞳ಶయ⩌㸦KOH-7A㸪 MEN ࠾ࡼࡧ MAN㸧యࡢ ĭST ࡣ㧗࠸್ࢆ♧ࡋࡓ㸦ĭST = 0.65, P < 0.001㸧㸬3 ಶ య⩌ࡢ⤌ࡳྜࢃࡏࡢ࠺ࡕ㸪ࡲࢇࡾࡻ࠺㇂ࡢ࣐ࢦಶయ⩌㸦MAN㸧ࡢಶయ⩌ ࡢ⤌ࡳྜࢃࡏ࡛ࡣ㸪ĭST ್ࡣ᭷ព 0 ␗࡞ࡗ࡚࠸ࡓࡀ㸦ĭST = 0.715–0.838㸪 P < 0.001㸹Table 5-4㸧 㸪⚄ཎᕝࡢ࣐ࢦಶయ⩌㸦KOH-7A㸧࣓ࣥࣀࢶࣛ㇂ࡢ ࣓࣡㸦MEN㸧ࡢ⤌ࡳྜࢃࡏ࡛ࡣ 95%ಙ㢗༊㛫 0 ࡀྵࡲࢀࡓ㸦ĭST = 0.039, P = 0.182㸧 㸬ࣁࣉࣟࢱࣉ㢖ᗘࡘ࠸࡚ࡶ KOH-7A MEN ࡢ㛫࡛ࡣ᭷ព࡞ᕪࡀࡳ ࡽࢀ࡞ࡗࡓࡀ㸦P = 0.186䠅䠈䛾⤌䜏ྜ䜟䛫䛷䛿᭷ព䛺ᕪ䛜☜ㄆ䛥䜜䛯䠄P < 0.001䠅䠊 ⪃ ᐹ ᅾ᮶ಶయ⩌ࡢ≉ᐃ ᮏ◊✲࡛ᚓࡽࢀࡓ mtDNA ࣁࣉࣟࢱࣉ Hap-1 ࡣ㸪ࡇࢀࡲ࡛ᨺὶࡀ⾜ࢃࢀ࡞ ࡗࡓணࡉࢀࡿἼᮌྜᕝ࣓ࣥࣀࢶࣛ㇂ࡢ࣓࣡ಶయ⩌㸦MEN㸧࡛ᅛᐃࡋ㸪 ⚄ཎᕝࡢ࣐ࢦಶయ⩌㸦KOH-7A㸧࡛ඃ༨ࡋ࡚࠸ࡓ㸬ࡉࡽ㸪Hap-1 ࡣࡍ࡚ࡢ ࣐ࢦᾋୖ⛶㨶ಶయ⩌㸦KOH-1F-7F㸧ࡶࡳࡽࢀ᭱ࡶ㧗࠸㢖ᗘࢆ♧ࡋࡓࡀ㸪㣴 㨶ሙࡢ࣐ࢦࡽࡣࡃ᳨ฟࡉࢀ࡞ࡗࡓ㸦Table 5-3㸧㸬ࡇࢀࡽࡢ⤖ᯝࡽ㸪 Hap-1 ࡣ⚄ཎᕝἼᮌྜᕝ⏕ᜥࡍࡿ࣐ࢦࡢᅾ᮶ࡢࣁࣉࣟࢱࣉ࡛࠶ࡿ⪃ ࠼ࡽࢀࡿ㸬ࡶࡋ௬㸪㐣ཤ㣴㨶ሙࡢ࣐ࢦࡀ࣓࣡ಶయ⩌㸦MEN㸧⏕ᜥᇦ ᨺὶࡉࢀ㸪࣓࣡㓄ࡋ࡚࠸ࡓ௬ᐃࡍࡿ㸪㣴㨶ሙࡢ࣐ࢦࡣከࡃࡢሙྜ ㏻ᖖࡢయⰍࢆᣢࡘ⪃࠼ࡽࢀࡿࡇࡽ㸪ᨺὶࡢ㊧ࡀ࣓࣡ಶయ⩌୰࠾ࡅ ࡿ࣐ࢦࡢฟ⌧ࡸ࣐ࢦࡢ㢖ᗘቑຍࡋ࡚ṧࡿ⪃࠼ࡽࢀࡿ㸬ࡋࡋᐇ㝿ࡣ㸪 ࣓࣡ࡣグ㍕ࡉࢀࡓ 1961 ᖺ㸦Kimura and Nakamura, 1961㸧௨᮶Ꮡ⥆ࡋ࡚࠾ࡾ㸪 ≉᭱㏆ࡢᑡ࡞ࡃࡶ 20 ᖺ㛫ࡣ㙚ῡࡢࡢୖὶഃ࡛༢⊂⏕ᜥᇦࢆᙧᡂࡋ࡚࠸ ࡿ㸦ᬒᖹ࣭ᚨග㸪2003㸹㏆⸨࣭➉ୗ㸪2005㸧㸬ࡇࢀࡽࡢࡇࡣ㸪࣓ࣥࣀࢶࣛ㇂ࡢ ࣓࣡ಶయ⩌㸦MEN㸧ࡀᅾ᮶ࡢࣁࣉࣟࢱࣉࢆಖᣢࡋ࡚࠸ࡿࡇ㸪ࡍ࡞ࢃࡕ㸪 81 ᅾ᮶ಶయ⩌࡛࠶ࡿࡇࢆᙉࡃ♧၀ࡍࡿ㸬ࡇࡢ᥎ㄽࡀṇࡋࡅࢀࡤ㸪ྠࡌ Hap-1 ࡀ ඃ༨ࡋ࡚࠸ࡓ⚄ཎᕝࡢ࣐ࢦಶయ⩌㸦KOH-7A㸧ࡶ㸪ᅾ᮶ಶయ⩌࡛࠶ࡿ⪃࠼ ࡿࡇࡀ࡛ࡁࡿ㸬 ୍᪉㸪ࡲࢇࡾࡻ࠺㇂ࡢ࣐ࢦ㸦MAN㸧࡛ࡣ㸪㣴㨶ሙࡢ࣐ࢦ㸦HAT-A㸧ࡳ ࡽࢀࡓ Hap-8 ࡀඃ༨ࡋ࡚࠸ࡓ㸦Fig. 5-1㸪Table 5-3㸧㸬Ỉᇦ࡛ࡢඛ⾜◊✲ 㸦Kawamura et al. 2007㸹Kubota et al. 2007㸧࡛ᣦࡉࢀ࡚࠸ࡿ࠾ࡾ㸪ศ┴ෆ ࡛ࡶ㸪ከࡃࡢ㣴㨶ሙ࠾࠸࡚」ᩘࡢἙᕝࡸ㣴㨶ሙ⏤᮶ࡍࡿ⣔⤫ࡀ㣴Ṫࡉࢀ࡚ ࠸ࡿ㸦ᮌᮏ㸪ᮍⓎ⾲㸧 㸬ࡋࡋ㸪ᮏ◊✲࡛ㄪᰝࡋࡓ㣴㨶ሙ࡛ࡣ㸪2 ࡘࡢ⣔⤫ 㸦HAT-A HAT-M㸧ࡔࡅࡀయⰍࡸᩬ⣠ᙧ㉁ࡢ⨾ࡋࡉᇶ࡙࠸࡚ 20 ᖺ௨ୖࢃ ࡓࡾ㑅ᢤ⫱✀ࡉࢀ࡚࠸ࡿ㸬ࡇࡢ࠺ࡕࡢ࣐ࢦ⣔⤫㸦HAT-A㸧ࡀ㸪1990 ᖺ௦ࡲ ࢇࡾࡻ࠺㇂ᨺὶࡉࢀࡓ⪃࠼ࡽࢀࡿࡇࡽ㸪 ᮏ◊✲࡛ 2008 ᖺ᥇ྲྀࡋࡓ ࣐ࢦ⣔⤫㸦HAT-A㸧ࡢಶయࡣ㸪ᨺὶࡉࢀࡓᙜࡢࣁࣉࣟࢱࣉࢆಖᣢࡋ࡚࠸ࡿ ᮇᚅࡉࢀࡿ㸬ࡲࡓ㸪HAT-A ࡽࡣ 2 ࡘࡢࣁࣉࣟࢱࣉ㸦Hap-8 Hap-11㸧ࡀ ᚓࡽࢀࡓࡀ㸪୧⪅ࡢ㐪࠸ࡣࢃࡎ 1 ሷᇶࡢ⨨࡛࠶ࡗࡓ㸬ࡇࢀࡽࡢ mtDNA ศ ᯒ⤖ᯝࡽ㸪ࡲࢇࡾࡻ࠺㇂ࡢ࣐ࢦಶయ⩌㸦MAN㸧ࡢ㒊ศࡣ㸪㠀බᘧᨺὶ ࡉࢀࡓ㣴㨶ሙࡢ࣐ࢦ⣔⤫⨨ࡁࢃࡗࡓࡶࡢ⪃࠼ࡽࢀࡿ㸬ࡋࡋ㸪ಶయ⩌ MAN ࡣࣄ࣮ࣂࢳࡢࡼࡾୗὶഃ࡛᥇ᤕࡉࢀࡓಶయ࡛ᙧᡂࡉࢀ࡚࠾ࡾ㸪ࡲࢇࡾ ࡻ࠺㇂⏕ᜥࡍࡿࡍ࡚ࡢ࣐ࢦಶయ⩌ࢆ௦⾲ࡍࡿࡶࡢ࡛ࡣ࡞࠸㸬㣴Ṫ㨶ࡢ㠀 බᘧ࡞ᨺὶࡣࣄ࣮ࣂࢳࡢࡼࡾୖὶഃ࡛⾜ࢃࢀࡓࡶࡢࡢ㸪࣐ࢦἙᕝᆺಶయ⩌ ࡢ㧗࠸ᐃఫᛶ㸦Nakano et al., 1990㸹㏆⸨࣭➉ୗ㸪2005㸧ࢆ⪃៖ࡍࡿ㸪࡛㝸 㞳ࡉࢀࡓࡇูࡢሙᡤᅾ᮶㨶ࡀ⏕ᜥࡋ࡚࠸ࡿྍ⬟ᛶࡣྰᐃ࡛ࡁ࡞࠸㸬 ಖ༢ Ἑᕝᛶࢧࢣ⛉㨶㢮ࡢᅾ᮶ಶయ⩌ࡣ㸪ࢲ࣒ࡸ⮬↛࡛㝸㞳ࡉࢀࡓ⊃࠸※ὶᇦ ⏕ᜥࡍࡿࡇࡀከࡃ㸪ࡋࡤࡋࡤప࠸㑇ఏⓗከᵝᛶ㧗࠸㏆ᙅໃࡢࣜࢫࢡࢆ♧ ࡍ㸦Yamamoto et al., 2004㸹Kubota et al., 2007㸧㸬ᮏ◊✲࠾࠸࡚ࡶ㸪ᨺὶࡀ⾜ࢃ ࢀ࡚࠸࡞࠸ணࡉࢀࡓ 2 ࡘࡢ㝸㞳㞟ᅋ㸦KOH-7A MEN㸧ࡣ㸪ᨺὶࡀ⾜ࢃࢀ ࡚࠸ࡿỈᇦࡢ㞟ᅋẚప࠸㑇ఏⓗከᵝᛶࢆ♧ࡋࡓ㸦Table 5-3㸧 㸬ࡇࢀࡣ㸪㑇ఏ 82 Ꮚὶືࡢไ㝈ࡸ㝸㞳ࡉࢀࡓ⏕ᜥᆅࡢᑠࡉ࡞⎔ቃᐜຊࡼࡗ࡚ࡶࡓࡽࡉࢀࡓ ⪃࠼ࡽࢀࡿ㸦Kawamura et al. 2007㸹Kubota et al. 2007㸹Sato et al. 2010㸧㸬ࡇࢀࡽ ࡢ㑇ఏⓗ࡞⬣ጾࢆ⦆ࡍࡿࡓࡵࡢ᪉ἲࡋ࡚㸪ಖ༢ෆ㒊ࡢศ㞟ᅋ㛫࠾ࡅ ࡿᅾ᮶㨶ࡢ⛣᳜ࡀᥦࡉࢀ࡚࠸ࡿ㸦Kawamura et al., 2007㸹Kanaiwa and Harada 2010㸹Sato et al., 2010㸧 㸬୍᪉㸪ྛศ㞟ᅋࡀᒁᡤ㐺ᛂࡋ࡚࠸ࡿሙྜࡣ㸪୰❧㑇 ఏ࣐࣮࣮࢝࠾ࡅࡿ㑇ఏⓗᵓ㐀ࡢᅾࡢࡳᇶ࡙࠸࡚ศ㞟ᅋ㛫࡛⛣᳜ࢆ⾜࠺㸪 ␗⣔㓄ᙅໃࡀ⏕ࡌࡿྍ⬟ᛶࡀᣦࡉࢀ࡚࠸ࡿ㸦Stockwell et al., 2003㸧 㸬ࡋࡓࡀ ࡗ࡚㸪ಖ༢ࡢỴᐃࡣ㑇ఏⓗ㸪ᙧែᏛⓗ㸪⏕ែᏛⓗ࡞≉ᚩᇶ࡙ࡁៅ㔜⾜ ࢃࢀࡿᚲせࡀ࠶ࡿ㸦Crandall et al., 2000㸹Garcia de Leaniz et al., 2007㸧㸬Ἴᮌྜᕝ ࣓ࣥࣀࢶࣛ㇂ࡢ࣓࣡ಶయ⩌㸦MEN㸧⚄ཎᕝࡢ࣐ࢦಶయ⩌㸦KOH-7A㸧ࡣ㸪 mtDNA ศᯒ࡛ࡣ⤫ィᏛⓗ༊ูࡍࡿࡇࡀ࡛ࡁ࡞ࡗࡓࡀ㸦Table 5-4㸧 㸪୧⪅ ࡣయⰍ࠾࠸࡚␗࡞ࡗ࡚࠾ࡾ㸪ࡇࡢయⰍࡢ㐪࠸ࡣ᰾ DNA ࠾ࡅࡿ㑇 ఏⓗᇶ┙ࡀᏑᅾࡍࡿࡇࡀᐇ㦂ࡽ᫂ࡽ࡛࠶ࡿ㸦ᒣෆ㸪1982㸹ᚨග㸪2003㸧㸬 ࡉࡽ㸪㏆⸨࣭➉ୗ㸦2005㸧ࡣ࣓ࣥࣀࢶࣛ㇂࡛ࡣ࣓࣡ࡢ⏘༸ᮇࡀྠᡤⓗ⏕ ᜥࡍࡿ࣐ࢦࡼࡾ㐜ࢀࡿࡇࢆሗ࿌ࡋ࡚࠸ࡿ㸦ࡓࡔࡋ Kano et al., 2006 ࡶཧ↷㸧㸬 ࢹ࣮ࢱࡀ♧ࡉࢀ࡚࠸࡞࠸ࡶࡢࡢ㸪ྠᵝࡢᣦࡣ࣓࣡ࢆኳ↛グᛕ≀ᣦᐃࡍࡿ 㝿ࡢ⏦ㄳ᭩㢮㸦⏣ᓥ㸪1965㸧ࡶグ㏙ࡉࢀ࡚࠸ࡿ㸬ࡋࡓࡀࡗ࡚㸪Ἴᮌྜᕝ࣓ࣥ ࣀࢶࣛ㇂ࡢ࣓࣡ಶయ⩌㸦MEN㸧⚄ཎᕝࡢ࣐ࢦಶయ⩌㸦KOH-7A㸧ࡣ㸪ࡑ ࢀࡒࢀ⊂❧ࡋࡓ༢ࡋ࡚ಖࡍࡁ⪃࠼ࡽࢀࡿ㸬 ಖࡢᥦᐇ㊶ Ἴᮌྜᕝࡲࢇࡾࡻ࠺㇂ࡢ࣐ࢦಶయ⩌㸦MAN㸧ࡣ㸪ࣄ࣮ࣂࢳࡢࡢୗὶഃࡢ ⚗⁺༊⏕ᜥࡋ࡚࠾ࡾ㸪 㐟⁺⪅ࡼࡿ㠀බᘧ࡞ᨺὶࡣ 1990 ᖺ௦ࡢୖὶഃ࡛ ⾜ࢃࢀࡓ㸬㣴Ṫ㨶ࡀᨺὶࡉࢀࡓ⌮⏤ࡣ㸪⚗⁺༊ࡢୖ㝈࡛࠶ࡿࣄ࣮ࣂࢳࡢࡼࡾ ୖὶഃ࡛ࡣ㐟⁺ࡀྍ⬟࡛࠶ࡿࡓࡵ⪃࠼ࡽࢀ㸦㏆⸨࣭➉ୗ㸪2005㸧 㸪ࡲࢇࡾࡻ࠺ ㇂ࡢ࣐ࢦಶయ⩌㸦MAN㸧ࡣ㸪ࡢୖὶഃࡽὶୗࡋ࡚ࡁࡓ㣴Ṫ㨶ࡢ㑇ఏᏊᾐ ㏱ࢆཷࡅࡓ⪃࠼ࡽࢀࡿ㸬ࡶࡋྠᵝࡢᨺὶࡀ࣓ࣥࣀࢶࣛ㇂ࡢ⚗⁺༊ࡢୖ㝈㸦᭱ ୖὶ㒊ࡢ◁㜵ሖሐ㸧ࡼࡾୖὶഃ࡛࡞ࡉࢀࡓሙྜࡣ㸪ᨺὶᆅⅬࡽὶୗࡋ࡚ࡁ 83 ࡓ࣐ࢦࡢ㓄ࡼࡗ࡚㸪࣓࣡ಶయ⩌ࡣᾘ⁛ࡍࡿྍ⬟ᛶࡶ⪃࠼ࡽࢀࡿ㸬ࡋ ࡓࡀࡗ࡚㸪Ἴᮌྜᕝ࣓ࣥࣀࢶࣛ㇂ࡢ࣓࣡ಶయ⩌㸦MEN㸧ࡲࢇࡾࡻ࠺㇂ࡢ ࣐ࢦಶయ⩌㸦MAN㸧ࢆಖࡍࡿࡓࡵࡣ㸪㔝ᕝ⁺ᴗ༠ྠ⤌ྜศ┴ࡢつ๎ ࡛タᐃࡉࢀ࡚࠸ࡿ⚗⁺༊ࡢୖ㝈ࢆ᧔ᗫࡋ㸪ࡉࡽᨺὶࢆ⚗Ṇࡍࡿࡇࡀᚲせ࡛ ࠶ࡿ㸬 ⚄ཎᕝࡢ࣐ࢦಶయ⩌㸦KOH-7A㸧ࡣ㸪ࡇࢀࡲ࡛᥎ㄽࡋ࡚ࡁࡓ࠾ࡾ㸪⚄ཎ ᕝୖὶᇦ࠾ࡅࡿᅾ᮶ಶయ⩌࡛࠶ࡿ⪃࠼ࡽࢀࡿ㸬ࡲࡓ㸪ᕞ࠾࠸࡚ᴟࡵ࡚ ᕼᑡ࡞ᅾ᮶࣐ࢦಶయ⩌ࡢ 1 ࡘ࡛࠶ࡿࡶ㸪࣓ࣥࣀࢶࣛ㇂ࡢ࣓࣡ಶయ⩌ 㸦MEN㸧᭱ࡶ⥭ᐦ࡞㛵ಀ࠶ࡿ࣐ࢦಶయ⩌ணࡉࢀࡿࡇࡽ㸪࣐ࢦ ಶయ⩌ KOH-7A ࡣಖࡍࡿᚲせࡀ࠶ࡿ⪃࠼ࡽࢀࡿ㸬ࡇ࠺ࡋࡓⴭࡋ࠸⊂⮬ᛶ ࡶࢃࡽࡎ㸪 ᮏ◊✲ࡀ⾜ࢃࢀࡿࡲ࡛ KOH-7A ࡣ⁺ᴗ࣭㐟⁺㛤ᨺࡉࢀ࡚ࡁࡓ㸬 ᮏ◊✲⤖ᯝᇶ࡙ࡁ㸪ศ┴ࡣ㔝ᕝ⁺ᴗ༠ྠ⤌ྜ㸪➉⏣ᕷ㸪ᆅඖ㐟⁺⪅ᅋ య➼ᑐࡋ࡚௨ୗࡢ 3 ࡘࡢᥦࢆ⾜ࡗࡓ㸸1㸧Ἴᮌྜᕝ࠾ࡅࡿ⚗⁺༊ୖ㝈ࡢ᧔ ᗫ㸪2㸧ྠἙᕝࡢ⚗⁺༊࠾ࡅࡿᨺὶࡢ⚗Ṇ㸪3㸧⚄ཎᕝࡢ୍ྜ┠ࡼࡾୖὶ ᇦ࠾ࡅࡿ⁺ᴗ࣭㐟⁺࠾ࡼࡧᨺὶࡢ⚗Ṇ㸬ࡇࢀࡽࡢᥦࡣཷࡅධࢀࡽࢀ㸪2009 ᖺࡽᐇ㊶ࡉࢀࡓ㸦ヲ⣽ࢆ➨ 6 ❶ ⥲ྜ⪃ᐹ࡛グ㏙ࡍࡿ㸧 㸬 84 Û( Û( Hap-1 KOH-1F N Sec. 1 Hap-2 Hap-3 KOH-2F Hap-4 Sea of Japan Hap-5 Hap-7 Pacific Ocean Û1 Sec. 2 Hap-6 Japan Hap-8 Ogata River KOH-3F Hap-9 Û(ޗ Ûޗ1 Û(ޗ Tosumi River Hap-10 Hap-11 Oita Prefecture Sec. 3 KOH-4F Seto Inland Sea man-made dam Ono River waterfalls Sec. 4 Ûޗ1 Kohbaru Stream Mt. Sobo KOH-5F Sec. 5 HAT-A KOH-7A MAN Hakiai Stream D KOH-6F Sec. 6 A HAT-M B KOH-7F Sec. 7 Hatchery strains C 1 km Manryo-dani MEN E Mennotsura-dani Fig. 5-1. Map of sampling sections with pie diagrams showing the distribution of mtDNA haplotype frequencies among amago and iwame populations in the Ogata River and its tributaries, the Kohbaru and the Hakiai streams, northeastern Kyushu, Japan. Sampling sections are circled with broken lines. Capital letters (A–E) indicate falls (A: Ichigome Falls, B: Yoroibuchi Falls, C: Hiibachi Falls), a bridge (D: Shirouzu Bridge), and an erosion control dam (E). No fishing has been allowed since 1970 from the Shirouzu Bridge (D) to the uppermost dam (E) in Mennotsura-dani and from the junction with Mennotsura-dani to Hiibachi Falls (C) in Manryo-dani. For details of samples, see Table 5-1. 85 6 8 9 2 3 11 10 1 4 7 87 7 5 mt1 11 Fig. 5-2. The most parsimonious network of mtDNA haplotypes detected from two amago populations, one iwame population, seven populations of amago fry from the Kohbaru and Hakiai streams, and hatchery populations of amago and masu salmon. Numeral in each circle corresponds to the haplotype number in Fig. 5-1 and Tables 5-2 and 5-3, and mt1 represents the haplotype previously observed in amago by Kawamura et al. (2007). Circles surrounded by heavy lines represent the haplotypes detected from the three isolated populations. Small circles indicate missing haplotypes. 86 87 MAN HAT-A HAT-M Abbreviation KOH-1F KOH-2F KOH-3F KOH-4F KOH-5F KOH-6F KOH-7F KOH-7A MEN Date Feb. 2004 Feb. 2004 Feb. 2004 Feb. 2004 Feb. 2004 Feb.-Mar. 2004 Feb.-Mar. 2004 Sep. 2005 Mar.-Jun. 2005 Feb. 2008 Mar. 2007 May 2008 May 2008 n 6 15 33 37 10 10 10 63 10 13 20 10 10 FL (mm) 30-48 29-42 27-40 25-42 26-35 28-33 23-31 70-227 100-225 136-324 93-235 154-196 160-185 Species and status of samples Newly-emerged amago fry* Newly-emerged amago fry* Newly-emerged amago fry* Newly-emerged amago fry* Newly-emerged amago fry Newly-emerged amago fry Newly-emerged amago fry, unstocked** Amago, unstocked** Iwame, fishing prohibited after 1970 Iwame, fishing prohibited after 1970 Amago, fishing prohibited after 1970*** Amago Masu salmon *** Some amago produced by a hatchery were stocked in the 1990s (local residents, personal communication). ** No amago have been stocked into Section 7 (OFC staff, personal communication). * Amago have been annually stocked into Section 1–4 by the Onogawa Fisheries Cooperative (OFC). Hatchery Manryou-dani Locality Kohbaru Stream Section 1 Section 2 Section 3 Section 4 Section 5 Section 6 Section 7 Section 7 Hakiai Stream Mennotsura-dani Table 5-1. Specimens of amago and iwame (Oncorhynchus masou ishikawae) and masu salmon (O. masou masou) used in this study. 88 Control region * 2 6 3 9 2 8 9 3 G G T . . . . . . . . . . . C . . . . A . . . . . . . T . . . . . 12S rRNA * 1 2 6 3 1 0 5 0 G G G . . . . A . . . A . . . C . . . . . . . . . . . . . . . . . 8 1 A G G G G G G G G G G 3 8 1 A . . . . G . G . . . 6 G . . . . . . . . A . ND1 region (1,963bp) 16S rRNA tRNA Leu ** An insertion (82bp) was observed in D-loop control region (between 838-839bp) * Transversion Haplotype Position D-loop region (1,944bp) Cb tRNA Pro 1 1 9 0 0 2 4 6 2 1 9 8 6 Hap-1 G T G A Hap-2 A . A . Hap-3 . . . . Hap-4 A . A . Hap-5** . . . . Hap-6 A . . . Hap-7 . . . G Hap-8 A . . . Hap-9 A C A . Hap-10 . . . . Hap-11 A . . . Hap-1. ND1 * * 1 5 0 4 5 C A . . . . . . A C . . . . . . . . . . . . * 1 4 1 G T . T . T . T T . T 1 4 7 T . . . C . . . . . . 4 7 7 A . . . . . G . . . . 6 4 5 G A . A . . . . A . . 6 7 2 G A A A A A A A A A A 6 8 4 A . . . G . . . . . . * 7 8 3 T G . G G G . G G . G 9 0 3 A G . G . G . G G . G iwame population, seven amago fry populations, and hatchery populations of amago and masu salmon. Dots indicate identity with Table 5-2. Variable nucleotide positions of D-loop and ND1 regions of mtDNA haplotypes detected from two amago populations, one 89 Number of samples 6 15 33 37 10 10 10 63 23 20 10 10 * Newly emerged fry KOH-1F* KOH-2F* KOH-3F* KOH-4F* KOH-5F* KOH-6F* KOH-7F* KOH-7A MEN MAN HAT-A HAT-M Population Numberof individuals with haplotypes 1 2 3 4 5 6 7 8 9 10 11 1 5 7 1 3 4 21 4 1 2 3 2 24 7 3 2 1 9 1 4 6 10 57 6 23 2 4 14 4 6 10 hatchery strains of amago (HAT-A) and masu salmon (HAT-M). Number of Haplotype diversity haplotypes h +SD 2 0.333 ± 0.215 4 0.714 ± 0.081 6 0.581 ± 0.094 5 0.548 ± 0.083 2 0.200 ± 0.154 2 0.533 ± 0.095 1 0.000 ± 0.000 2 0.175 ± 0.060 1 0.000 ± 0.000 3 0.484 ± 0.113 2 0.533 ± 0.095 1 0.000 ± 0.000 Nucleotide diversity S +SD 0.0011 ± 0.0007 0.0011 ± 0.0007 0.0035 ± 0.0018 0.0009 ± 0.0005 0.0005 ± 0.0003 0.0012 ± 0.0007 0.0000 ± 0.0000 0.0004 ± 0.0003 0.0000 ± 0.0000 0.0006 ± 0.0004 0.0001 ± 0.0001 0.0000 ± 0.0000 amago populations (KOH-7A, MAN), one iwame population (MEN), seven amago fry populations (KOH-1F–7F), and Table 5-3. Haplotype distribution, number of haplotypes, haplotype diversity (h), and nucleotide diversity (p) of two Table 5-4. Estimated SDLUZLVH ĭ67 EHORZ diagonal) and P-value of the pairwise exact test of population differentiation (above diagonal) among three isolated populations (i.e., two amago populations [KOH-7A, MAN] and one iwame population [MEN]). Population KOH-7A MEN MAN KOH-7A MEN 0.186 0.039 0.715* 0.838* ĭST overall value = 0.650 (P < 0.0001). * P < 0.0001 90 MAN < 0.001 < 0.001 ➨ 6 ❶ ⥲ྜ⪃ᐹ ᮏ◊✲࡛ࡣ㸪ᕞᮾ㒊࠾ࡅࡿ࣐ࢦࡢಖࢆᅗࡿࡓࡵ㸪ศ┴㔝ᕝỈ ⣔⥴᪉ᕝࡑࡢᨭὶ㸪⚄ཎᕝἼᮌྜᕝࢆᑐ㇟Ỉᇦࡋ࡚㸪⏕ᜥᇦయࢃࡓ ࡿ࣐ࢦࡢὶ⛬ศᕸࡢ≉ᚩࢆᢕᥱࡍࡿࡓࡵࡢ㨶㢮⩌㞟ゎᯒ㸦➨ 2 ❶㸧㸪⮬↛⏕ ⏘ࡢᐇែࢆᢕᥱࡍࡿࡓࡵࡢᾋୖ⛶㨶ศᕸㄪᰝ㸦➨ 3 ❶㸧㸪⮬↛⏕⏘ࡢಁ㐍ᚲ せ࡞▱ぢࢆᢕᥱࡍࡿࡓࡵࡢᾋୖ⛶㨶ࡢ⏕ᜥሙᡤ⏝ㄪᰝ㸦➨ 4 ❶㸧 㸪ࡑࡋ࡚ᅾ᮶ ࣐ࢦಶయ⩌ࡢ⏕ᜥỈᇦࢆ≉ᐃࡍࡿࡓࡵࡢ㑇ఏᵓ㐀ศᯒ㸦➨ 5 ❶㸧ࢆᐇࡋࡓ㸬 ᮏ❶࡛ࡣ㸪ྛ❶ࡢෆᐜࢆᕞᮾ㒊࠾ࡅࡿ࣐ࢦࡢಖ࠸࠺┠ⓗἢࡗ ࡚ᨵࡵ࡚ホ౯㸪ゎ㔘ࡍࡿࡶ㸪ྛ❶࡛ಶูᚓࡽࢀࡓ⤖ᯝࡢ㛵㐃ᛶࢆ᳨ウ ࡍࡿࡇ࡛㸪ᮏㄪᰝỈᇦ࠾ࡅࡿ࣐ࢦಖ⟇ࡢయീࢆᑟฟࡍࡿࡇࢆ┠ⓗ ࡍࡿ㸬ࡉࡽ㸪ᚋࡢㄢ㢟ࡸᡂᯝࡢά⏝ࡘ࠸࡚ࡶ⪃ᐹࡍࡿ㸬 ὶ⛬ἢࡗࡓ㨶㢮⩌㞟ࡢኚ ➨ 2 ❶࡛ࡣ㸪㔝ᕝỈ⣔⥴᪉ᕝࡑࡢᨭὶ㸪⚄ཎᕝἼᮌྜᕝ࠾࠸࡚⏕ᜥ 㨶㢮ࡢ₯Ỉ┠どほᐹࢆᐇࡋ㸪4 ┠ 7 ⛉ 14 ✀㸦ள✀ࢆྵࡴ㸧ࢆ☜ㄆࡋࡓ㸬ࡲࡓ㸪 㨶㢮⩌㞟ࡣୖὶഃࡢ࣐ࢦࢱ࢝ࣁࣖࡀඃ༨ࡍࡿ༢⣧࡞⤌ᡂࡽ㸪ୗὶഃࡢ࢘ ࢢࡸ࣒࢝࣡ࢶ➼ࡢࢥ⛉㨶㢮ࡀඃ༨ࡍࡿࡼࡾ」㞧࡞⤌ᡂ⛣⾜ࡋ㸪ࡇࡢኚ ࡣὶ⛬ἢࡗ࡚༢ㄪ࡞ኚࢆ♧ࡍࡁ࡞✵㛫ࢫࢣ࣮ࣝࡢ⎔ቃ໙㓄㸦ᶆ㧗㸪Ἑ ᗋ໙㓄㸪Ỉ㠃ᖜ㸪ὶ㔞㸪Ỉ 㸧┦㛵ࡍࡿࡇࢆ᫂ࡽࡋࡓ㸬୍᪉㸪ࢥ⛉ 㨶㢮ࡢಶయ⩌ᐦᗘࡣ༢ㄪ࡞ኚࢆ♧ࡉࡎ㸪≉ࢱ࢝ࣁࣖࡢಶయ⩌ᐦᗘࡣ㸪࣐ ࢦ࣭ࢱ࢝ࣁࣖࢆ୰ᚰࡍࡿୖὶഃࡢ⩌㞟ࡽࢥ⛉㨶㢮ࢆ୰ᚰࡍࡿୗὶഃࡢ ⩌㞟ࡢ⛣⾜ᖏ㸦transition zone㸹Rahel and Hubert, 1991㸧࠾࠸࡚ⴭࡋࡃప ࡗࡓ㸬⛣⾜ᖏࡢୖὶഃࡣሖሐ⩌ࡀᏑᅾࡋ࡚࠸ࡓࡇࡽ㸪⛣⾜ᖏ࡛ࡣேⅭᨵ ኚࡢᝏᙳ㡪ࡀ⏕ࡌ࡚࠸ࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡿ㸦ᮌᮏ㸪2015㸧 㸬 㨶㢮⩌㞟ࡢ⛣⾜ᖏ࡛⎔ቃࡢேⅭᨵኚࡢᝏᙳ㡪ࡀ⏕ࡌࡿྍ⬟ᛶࢆ⪃៖ࡍࡿࡇ ࡣ㸪ᕞᮾ㒊࠾࠸࡚࣐ࢦࡢಖࢆᅗࡿୖ࡛㠀ᖖ㔜せ⪃࠼ࡽࢀࡿ㸬࡞ ࡐ࡞ࡽ࣐ࢦࡣ෭Ỉᛶ㨶㢮࡛࠶ࡾ㸦Kimura, 1989㸹Ỉ㔝࣭ᚚໃ㸪2000㸧 㸪ἙᕝỈ 91 ࡣ୍⯡ୗὶ᪉ྥୖ᪼ࡍࡿࡓࡵ㸦Vannote et al., 1980㸧㸪ᕞࢆྵࡴす᪥ᮏ࡛ ࣐ࢦಶయ⩌ᐦᗘࡀୗὶ᪉ྥపୗࡍࡿࡇࡣ㸪௬ேⅭᨵኚࡢᝏᙳ㡪ࡼࡿ ⌧㇟࡛࠶ࡗࡓࡋ࡚ࡶ㸪⎔ቃ໙㓄ἢࡗࡓ⮬↛⌧㇟༊ูࡍࡿࡇࡀ㞴ࡋ࠸ࡓ ࡵ࡛࠶ࡿ㸬≉ᮏㄪᰝỈᇦ࡛ࡣ㸪࣐ࢦࡢ⏕ᜥᇦࡢ 7 ௨ୖࡀ⛣⾜ᖏࢆྵࡴୗ ὶഃỈᇦᏑᅾࡋࡓࡇࡽ㸪⛣⾜ᖏேⅭᨵኚࡢ㛵ಀࡣ㔜せ࡛࠶ࡿ㸬࣐ࢦ ಶయ⩌ᐦᗘࡢୗὶ᪉ྥࡢపୗせᅉࢆ⮬↛⌧㇟ேⅭᨵኚࡢᙳ㡪ศ㞳ࡍࡿࡓ ࡵࡣ㸪ேⅭᨵኚࢆཷࡅ࡚࠸࡞࠸࣐ࢦࡢ⏕ᜥἙᕝ࠾࠸࡚㸪ಶయ⩌ᐦᗘࡢὶ ⛬ኚᵝᘧࢆ᫂ࡽࡍࡿᚲせࡀ࠶ࡿ㸬ࡋࡋ㸪※ὶ㒊ࡽᡪ≧ᆅࡅ࡚ࡣ Ἑᗋ໙㓄ࡸἙᕝᙧែᆺ➼ࡀᛴ⃭ኚࡍࡿࡇ㸦ⴛሙ㸪2013㸧 㸪Ἑᕝࡢ◁㜵ᕤ ࡀ※ὶ㒊ࡽᡪ≧ᆅࡲ࡛ࡢὶ࡛⾜ࢃࢀࡿࡇ㸦㧗ᶫ㸪2000㸧 㸪ࡑࡋ࡚ᅜෆࡢἙ ᕝࡣࡍ࡛ኰࡋ࠸ᩘࡢ◁㜵ࢲ࣒ࡀᘓタࡉࢀ࡚࠸ࡿࡇ㸦᳃⏣࣭ᒣᮏ㸪2004㸹 㐲⸨㸪2006㸹ᮌᮏ㸪2013㸹2015㸧ࢆ⪃៖ࡍࡿ㸪す᪥ᮏࡢከࡃࡢἙᕝ ࠾࠸࡚㨶㢮⩌㞟ࡢ⛣⾜ᖏࡣ◁㜵ࢲ࣒⩌ࡢ┤ୗ⨨ࡍࡿࡶࡢ⪃࠼ࡽࢀࡿ㸬 ࡋࡓࡀࡗ࡚㸪࣐ࢦࡢಶయ⩌ᐦᗘࡢὶ⛬ኚࢆㄪࡿࡇࡼࡗ࡚⛣⾜ᖏ࠾ ࡅࡿேⅭᨵኚࡢᙳ㡪ࢆ᥎ᐃࡍࡿࡇࡣ㸪⌧Ⅼ࡛ࡣᅔ㞴ᛮࢃࢀࡿ㸬 ୍᪉㸪ከࡃࡢࢥ⛉㨶㢮ࢆྵࡴ Ỉ㨶ࡣᗈ ᛶ࡛࠶ࡾ㸪ࡑࡢ⏕ᜥ㔞ࡣỈ ௨ እࡢせᅉࡶᙉࡃつᐃࡉࢀࡿࡇࡽ㸦Ỉ㔝࣭ᚚໃ㸪2000㸧 㸪ࢥ⛉㨶㢮ࡢಶయ ⩌ᐦᗘࡢὶ⛬ኚࢆㄪ࡚⛣⾜ᖏ࠾ࡅࡿேⅭᨵኚࡢᙳ㡪ࢆ᥎ᐃࡍࡿࡇࡣ᭷ ⏝⪃࠼ࡽࢀࡿ㸬ᮏ◊✲࡛ࡣ㸪ࢱ࢝ࣁࣖ㸪࢘ࢢ㸪࣒࢝࣡ࢶࡢಶయ⩌ᐦᗘࡢὶ ⛬ኚࡣ㸪⛣⾜ᖏෆ㒊࠾࠸࡚㸪Ỉ ࡸᆅᙧ࠸ࡗࡓࡁ࡞✵㛫ࢫࢣ࣮ࣝࡢ⎔ ቃ໙㓄୍⮴ࡋ࡞ࡗࡓ㸬⛣⾜ᖏෆ㒊ࡢ㨶㢮⏕ᜥ㔞ࡣ㸪ࡼࡾᑠࡉ࡞✵㛫ࢫࢣ࣮ ࣝࡢせᅉつᐃࡉࢀࡿࡇࡽ㸦Jackson et al., 2001㸹Inoue and Nunokawa, 2002㸹 Torgersen et al., 2006㸧㸪ࡇࢀࡽࡢ㨶✀ࡢಶయ⩌ᐦᗘỈ῝㸪ὶ㏿㸪Ἑᗋᮦᩱ➼ࡢ ⎔ቃኚ㔞ࡢ㛵ಀࢆㄪࡿࡇࡀ᥎ዡࡉࢀࡿ㸬ࡉࡽ㸪ᑠࡉ࡞✵㛫ࢫࢣ࣮࡛ࣝࡣ㸪 㨶㢮ࡢ⏕ᜥ㔞ࡣ≀⌮⎔ቃࡼࡾࡶ✀㛫➇த➼ࡢ⏕≀㛫┦స⏝ᙉࡃᙳ㡪ࢆཷࡅ ࡿࡇࡀ▱ࡽࢀ࡚࠸ࡿ㸦Jackson et al., 2001㸧㸬≉ࢱ࢝ࣁࣖࡣ㸪࣐ࢦ㸪࣒࢝࣡ ࢶ㸪࢘ࢢ➼ඹᏑࡋࡓሙྜࡣ✀㛫㛵ಀ࡛᭱ୗ࡞ࡾ㸪࡞ሙᡤ࡛⏕ά ࡍࡿ㸦ᯈ㸪1976㸪1979㸹Ỉ㔝࣭ᚚໃ㸪2000㸧 㸬ேⅭᨵኚࡢᙳ㡪ࡣ㸪✀㛫㛵ಀࢆ 92 ㏻ࡌ࡚ࡶࡗࡶ㡰ࡢప࠸ࢱ࢝ࣁࣖࡢ⏕ᜥ㔞㢧ⴭᫎࡉࢀࡿྍ⬟ᛶࡶ⪃࠼ ࡽࢀࡿࡇࡽ㸪ࢱ࢝ࣁࣖࡢ⏕ᜥ㔞ࡣ㸪⛣⾜ᖏ࠾ࡅࡿேⅭᨵኚࡢᙳ㡪ࢆ᥎ᐃ ࡍࡿୖ࡛≉᭷⏝࡞ᣦᶆ࡞ࡿྍ⬟ᛶࡀ⪃࠼ࡽࢀࡿ㸬 ࡲࡓ㸪⥴᪉ᕝࡢୗὶഃࡣࢩ࣐ࢻࢪࣙ࢘ᒓ㨶㢮ࡀ⏕ᜥࡋ࡚࠸ࡓ㸬ᮏ◊✲࡛ࡣ ₯Ỉ┠どほᐹࢆ⾜ࡗࡓࡓࡵᒓࡲ࡛ࡢศ㢮ࡋࡓࡀ㸪ඛ⾜◊✲࡛ࡣ㸪⥴᪉ᕝࡢࢩ ࣐ࢻࢪࣙ࢘㢮ࡣ㸪㞄᥋ࡍࡿศᕝỈ⣔ࡢࢩ࣐ࢻࢪࣙ࢘㢮ࡶࢩ࣐ࢻࢪࣙ࢘ 㸦4 ಸయ㸧Cobitis biwae ྠᐃࡉࢀ࡚࠸ࡿ㸦㧗℈㸪2006㸹ᕝ㸪2009㸧㸬 ➹⪅ࡽࡶ㸪2013 ᖺ 11 ᭶ St. 1–2㸦Fig. 2-1㸧࠾࠸࡚ࢩ࣐ࢻࢪࣙ࢘ᒓ㨶㢮ࢆࢱ ࣔ⥙࡛᥇㞟ࡋ㸪⬚㫅ᇶ㒊㦵㉁┙ࡢᙧ≧➼㸦୰ᆓ㸪2013㸧ᇶ࡙ࡁࢩ࣐ࢻࢪࣙ࢘ 㸦4 ಸయ㸧࡛࠶ࡿࡇࢆ☜ㄆࡋࡓ㸦ᮌᮏ㸪ᮍⓎ⾲㸧 㸬ࢩ࣐ࢻࢪࣙ࢘㸦4 ಸయ㸧 ࡣ㸪୰ᅜ࣭ᅄᅜᆅ᪉ࡢ℩ᡞෆഃす㒊ᆅᇦᗈࡃศᕸࡍࡿࡀ㸪ᕞ࡛ࡣศᕝỈ ⣔㔝ᕝỈ⣔ࡔࡅࡽሗ࿌ࡉࢀ࡚࠾ࡾ㸦ᕝ㸪2009㸧 㸪ࢠࢠ Pseudobagrus nudiceps㸦Mizoiri et al., 1997㸹Watanabe, 1998㸹Watanabe and Nishida, 2003㸧ࡸࢻ ࣥࢥ㸦Sakai et al., 1998㸧ࡢᆅ⌮ⓗศᕸ࠾ࡼࡧ㑇ఏⓗ㢮⦕ᛶࡶ㸪ࡇࢀࡽᆅ ᇦᕞᮾ㒊Ἑᕝࡢ῝࠸⏕≀ᆅ⌮Ꮫⓗ㛵ࢃࡾࢆ♧ࡍ㸦ᕝ㸪2009㸧 㸬ࡇࡢ ࡇࡣ㸪➨ 5 ❶࡛᫂ࡽ࡞ࡗࡓ⚄ཎᕝࡢᅾ᮶࣐ࢦಶయ⩌ࡢ⣔⤫㛵ಀࢆ⪃ᐹ ࡍࡿୖ࡛㸪ᚋ㸪୰ᅜ࣭ᅄᅜᆅ᪉ࡢ℩ᡞෆഃす㒊ᆅᇦࡢ࣐ࢦಶయ⩌ࡢ㛵ಀ ࢆ⪃៖ࡍࡿᚲせࡀ࠶ࡿࡇࢆ♧ࡋ࡚࠸ࡿ㸬 ࣐ࢦᾋୖ⛶㨶ࡢὶ⛬ศᕸ ➨ 3 ❶࡛ࡣ㸪ㄪᰝỈᇦ㸦16.3 km㸧ࡢ₯Ỉ┠どほᐹࡼࡾ࣐ࢦᾋୖ⛶㨶ࡢ ὶ⛬ศᕸࢆㄪ㸪ಶయ⩌ᐦᗘࡣୖὶഃ࡛㧗ࡃୗὶഃ࡛ప࠸ࡇࢆ᫂ࡽࡋࡓ㸬 ࡲࡓ㸪ᾋୖ⛶㨶ࡢศᕸࡢ೫ࡾࡣ⣙ 1 ࣨ᭶㛫ኚࡏࡎ㸪ᗂ㨶ᮇ௨㝆ࡢ࣐ࢦࡢಶ య⩌ᐦᗘ᭷ព࡞┦㛵ࢆ♧ࡋࡓ㸬ࡇࢀࡽࡢ⤖ᯝ㸪ᗂ㨶ᮇ௨㝆ࡢ࣐ࢦ࡛ሗ࿌ ࡉࢀ࡚࠸ࡿ㧗࠸ᐃఫᛶࢆ⪃៖ࡍࡿ㸪 ࣐ࢦᾋୖ⛶㨶ࡣᑡ࡞ࡃࡶ⣙ 1 ᭶㛫㸪 ⏘༸ᗋ㏆␃ࡲࡿࡶࡢ⪃࠼ࡽࢀࡓ㸦ᮌᮏ㸪2013㸧 㸬 ୗὶഃ࡛ᾋୖ⛶㨶ࡢಶయ⩌ᐦᗘࡀప࠸⌮⏤ࡋ࡚㸪➨ 3 ❶࡛ࡣ㧗࠸㔮⋓ᅽ ࡼࡿぶ㨶ᩘࡢ㊊㸪ୗὶഃ࡛ᒾ┙Ἑᗋࡸࢺࣟࡢᆅᙧࡀከ࠸ࡇࡼࡿ⏘༸㐺 93 ᆅࡢ㊊ࢆᣦࡋࡓ㸬୍᪉㸪➨ 2 ❶ࡢ㨶㢮⩌㞟ࡢኚᵝᘧࢆ⪃៖ࡍࡿ㸪ᾋୖ ⛶㨶ࡢಶయ⩌ᐦᗘࡀప࠸ሙᡤࡣ㸪㨶㢮⩌㞟ࡢ⛣⾜ᖏࢆྵࡴୗὶഃỈᇦ୍⮴ࡍ ࡿ㸬ࡇࡢࡇࡣ㸪➨ 2 ❶࡛⪃ᐹࡋࡓேⅭᨵኚࡢᙳ㡪ࡀ㸪ୗὶഃỈᇦ࠾ࡅࡿ ࣐ࢦᾋୖ⛶㨶ࡢಶయ⩌ᐦᗘపୗࡶ㛵ࡍࡿྍ⬟ᛶࢆ♧ࡋ࡚࠸ࡿ㸬◁㜵ࢲ࣒ࡣ ୖὶࡽὶୗࡋࡓᅵ◁ࢆᤕᤊࡋ࡚ୗὶࡢᅵ◁౪⤥ࢆไ㝈ࡍࡿࡇࡽ 㸦㧗ᶫ㸪 2000㸧㸪ୗὶഃ࡛ࡣἙᕝᆅᙧࡢኚࡸἙᗋࡢపୗ㸦బᕝ࣭୰ᮧ㸪2010㸧㸪Ἑᗋࡢ 㟢┙㸦୰ᮧ㸪2011㸧ࡀ⏕ࡌ㸪Ỉ῝㸪ὶ㏿㸪Ἑᗋᮦᩱ➼ࡢኚࢆకࡗ࡚࣐ࢦ ࡢ⏘༸ሙᡤࢆຎࡉࡏ࡚࠸ࡿྍ⬟ᛶࡶ⪃࠼ࡽࢀࡿ㸬ᅜෆ࡛ࡣ㸪ከࡃࡢ◁㜵ࢲ࣒ ࡀ 1970 ᖺ௦௨㝆ᘓタࡉࢀ㸦᳃⏣࣭ᒣᮏ㸪2004㸧㸪◁㜵ࢲ࣒ࡼࡿἙᗋࡢ㟢┙ ࡣタ⨨ᚋࡢ㛫⤒㐣కࡗ࡚㐍⾜ࡍࡿࡇࡀ▱ࡽࢀ࡚࠸ࡿ㸦୰ᮧ㸪2011㸧㸬ᮏ ㄪᰝỈᇦࡢୗὶഃ㸦࠼ࡤ Sec. 2㸧࡛ࡣ㸪2 ᅇࡢㄪᰝࢆ㏻ࡌ࡚ࢇᾋୖ⛶㨶 ࡀ☜ㄆࡉࢀ࡞ࡗࡓࡀ㸪1970 ᖺ௦ࡣ࣐ࢦࡢ⏘༸⾜ືࡀ㢖⦾ほᐹࡉࢀ࡚࠸ ࡓ㸦㊊❧ᚭಙẶ㹙㣴㨶ሙ Ỉⱌ㹛 㸪⚾ಙ㸧㸬ࡇࡢࡇࡣ㸪◁㜵ࢲ࣒ࡼࡿ⏘༸⎔ ቃࡢຎࡀ㸪タ⨨ᚋ 30㹼40 ᖺࢆ⤒࡚㢧ᅾࡋࡓࡇࢆ♧ࡍࡢࡶࡋࢀ࡞࠸㸬 ࡲࡓ㸪➨ 5 ❶࡛⾜ࡗࡓ࣐ࢦᾋୖ⛶㨶ࡢ mtDNA ࣁࣉࣟࢱࣉศᯒࡼࡾ㸪 ẖᖺ㣴Ṫ㨶ࡀᨺὶࡉࢀ࡚࠸ࡿୗὶഃỈᇦࡣ㸪㠀ᅾ᮶ಶయࡀ⏕ᜥࡋ࡚࠸ࡿྍ⬟ ᛶࡀ♧ࡉࢀࡓ㸦Kimoto et al., ༳ๅ୰㸧 㸬ࡇࢀࡲ࡛㸪ࢧࢣ⛉㨶㢮ࡢᵝࠎ࡞ᙧ㉁ 㛵ࡋ࡚ᒁᡤ㐺ᛂࡢᏑᅾࡀሗ࿌ࡉࢀ㸪ᨺὶࡉࢀࡓ⎔ቃᑐࡍࡿ㣴Ṫ㨶ࡢ㐺ᛂᗘࡣ ᅾ᮶ಶయࡼࡾప࠸ࡇࡀணࡉࢀ࡚࠸ࡿ㸦Taylor, 1991㸹Garcia de Leaniz et al., 2007㸸Kikko et al., 2008㸹 Fraser et al., 2011㸹Primmer, 2011㸹Drinan et al., 2012㸧 㸬 ࣐ࢦࡢ⏕⏘㛵ࡋ࡚ᒁᡤ㐺ᛂࡢᏑᅾࢆ♧ࡍሗ࿌ࡣ࡞࠸ࡀ㸪⚄ཎᕝࡢ࣐ࢦ ⏕ᜥୖ㝈㸦ᮌᮧ㸪1959㸹ᮌᮏ㸪ᮍⓎ⾲㸧࡛࠶ࡿྜ┠ࡢ㸦ⴠᕪ 11 m㸧ࡢ ና࡛ࡣ㸪࣐ࢦ㸪࣐࣓ࣖࡢἙᕝᆺಶయ⩌ࡗ࡚Ⰻዲ࡞⏘༸ሙᡤࡉࢀࡿᖹ ℩㸦ⓑ▼㸪1957㸹ᮌᮧ㸪1972㸧ࡀࡰᏑᅾࡋ࡞࠸ࡶࢃࡽࡎ㸪2004–2010 ᖺ 4 ᭶ࡣᖖᾋୖ⛶㨶ࡀ☜ㄆࡉࢀࡓ㸦ᮌᮏ㸪ᮍⓎ⾲㸧㸬ናࡢ➃ࡸᗏ㒊࡛ ࡣ㸪ᨺ༸࣭ᨺ⢭ࡣᮍ☜ㄆ࡞ࡀࡽ࣐ࢦࡢ⏘༸⾜ືࡀほᐹࡉࢀࡿࡇࡽ㸦ᮌᮏ 㸪ᮍⓎ⾲㸧 㸪᭱ୖὶ㒊ࡢ࣐ࢦࡣୗὶഃࡣ␗࡞ࡿ⎔ቃ࡛ࡶ⏘༸ࡍࡿࡶࡢ ⪃࠼ࡽࢀࡿ㸬ࡇࢀࡀᒁᡤ㐺ᛂ┦ᙜࡍࡿ࠺ࡣ㸪⛣᳜ᐇ㦂㸦common garden 94 experiments㸧➼ࡼࡾ᫂ࡽࡍࡿᚲせࡀ࠶ࡿࡶࡢࡢ㸦Garcia de Leaniz et al., 2007㸹Fraser et al., 2011㸧 㸪࣐ࢦࡢ⏕⏘㛵ࡍࡿᙧ㉁࠾࠸࡚ࡶ㸪ᒁᡤ㐺ᛂ ࡀᏑᅾࡍࡿྍ⬟ᛶࢆ♧ࡍࡶࡢ࡛࠶ࡿ㸬ࡋࡓࡀࡗ࡚㸪ୗὶഃỈᇦ࠾ࡅࡿ࣐ࢦ ࡢ⮬↛⏕⏘ࡢపୗࡣ㸪㠀㐺ᛂⓗ࡞㣴Ṫ㨶ࡢᨺὶࡼࡗ࡚ࡶࡓࡽࡉࢀࡓྍ⬟ᛶ ࡶ⪃៖ࡍࡿࡁᛮࢃࢀࡿ㸬 ࣐ࢦᾋୖ⛶㨶ࡢ⏕ᜥሙᡤ⏝ ➨ 4 ❶࡛ࡣ㸪࣐ࢦ⏕ᜥᇦࡢୗὶഃ࡛㸪ὶ㊰༢ࢫࢣ࣮ࣝ࠾ࡅࡿᾋୖ⛶㨶 ࡢ⏕ᜥሙᡤ⏝ࢆㄪࡿࡓࡵ㸪⥴᪉ᕝࡢ 1.4 km ༊㛫࡛」ᩘᖺࢃࡓࡿ₯Ỉ┠ど ほᐹࢆ⾜ࡗࡓ㸬ᾋୖ⛶㨶ᩘࡣ㸪᪩℩࿘㎶ࡢὶ㊰༢ࢱࣉ᭷ព࡞ṇࡢ┦㛵㸪 ἢᓊỈ῝࠾ࡼࡧ⏘༸ᗋࡽࡢ㊥㞳᭷ព࡞㈇ࡢ┦㛵ࢆ♧ࡋࡓ㸬᪩℩࿘㎶ࡢὶ㊰ ༢ࢱࣉࢆ⏝ࡍࡿഴྥࡣୗὶഃỈᇦ≉᭷࡛࠶ࡾ㸪ࡇࢀࡣ㸪㧗࠸࢚ࢿࣝࢠ ࣮ຠ⋡ࢆࡶࡓࡽࡍዲ㐺࡞ᚤ⏕ᜥሙᡤ㸦㏿࠸ὶࢀ㏆࠸⦆ὶ㒊ศ㸧ࡀ㸪ୖὶഃ࡛ ࡣ㐢ᅾࡍࡿࡀ㸪ୗὶഃ࡛ࡣ᪩℩㏆ᒁᅾࡍࡿࡓࡵ⪃࠼ࡽࢀࡓ㸬ࡓࡔࡋ㸪ὶ ㊰༢ࢫࢣ࣮࡛ࣝࡇࢀࡽࡢ᮲௳ࡀ‶ࡓࡉࢀ࡚࠸࡚ࡶ㸪ᚤ⏕ᜥሙᡤࢫࢣ࣮ࣝࡢ⎔ ቃ᮲௳ࡀ༑ศ࡞ሙᡤ࡛ࡣ㸪 ᾋୖ⛶㨶ࡀୗὶ᪉ྥ⛣ືࡍࡿྍ⬟ᛶࡀ♧ࡉࢀࡓ㸬 ௨ୖࡢࡇࡽ㸪࣐ࢦ⏕ᜥᇦୗὶഃ࠾࠸࡚ᾋୖ⛶㨶ࡢಖࡸቑṪࢆᅗࡿࡓ ࡵࡣ㸪᪩℩࿘㎶ࡢὶ㊰༢ࢱࣉࡢෆ㒊ዲ㐺࡞ᚤ⏕ᜥሙᡤࢆᩚഛࡍࡿࡇ㸪 ⎔ቃᐜຊ⛶㨶ࡢቑṪ㔞ࡢࣂࣛࣥࢫࢆ⪃៖ࡍࡿࡇࡀ㔜せ⪃࠼ࡽࢀࡿ㸬 ᮏ◊✲࡛ࡣ㸪2009 ᖺࡢ 1–3 ᭶Ἑ㐨⊃✽㒊㸦༊⏬ 35–50㸧࡛☜ㄆࡉࢀࡓᾋୖ ⛶㨶ࡀୗὶ᪉ྥ⛣ືࡋࡓ᥎ᐃࡉࢀࡓ㸬ᮏ◊✲ࡣ࣐ࢦࡢ⮬↛⏘༸⏤᮶ࡍ ࡿᾋୖ⛶㨶ࢆᑐ㇟ࡋ㸪ཷ⢭༸ࡸⓎ║༸ᑐࡍࡿᶆ㆑ࢆ⾜ࢃ࡞ࡗࡓࡓࡵ㸪ཝ ᐦࡣ㸪☜ㄆࡉࢀࡓᾋୖ⛶㨶ࡀࡢ⏘༸ᗋ⏤᮶ࡍࡿࡢࡣ࡛᫂࠶ࡿ㸬ࡋ ࡋ㸪ࢧࢣ⛉㨶㢮ࡢᾋୖ⛶㨶ࡀ⏘༸ᗋ㏆␃ࡲࡿୗὶ᪉ྥ⛣ືࡍࡿ᥎ᐃ ࡉࢀ࡚࠸ࡿࡇ㸦Moore and Gregory, 1988㸹Nagata, 2002㸹Steingrímsson and Grant, 2003㸹Einum et al., 2011㸧 㸪2009 ᖺࡢㄪᰝᮇ㛫୰ࡣ༊⏬ 51 䜘䜚ୖὶ䛷䜰䝬䝂ᾋ ୖ⛶㨶䛜☜ㄆ䛥䜜䛺䛛䛳䛯䛣䛸ࢆ⪃៖ࡍࡿ㸪Ἑ㐨⊃✽㒊ࡢᾋୖ⛶㨶ࡢ㒊ศࡣ㸪 2008 ᖺ⛅༊⏬ 51 ᙧᡂࡉࢀࡓ⏘༸ᗋ⏤᮶ࡍࡿྍ⬟ᛶࡀ㧗࠸⪃࠼ࡽࢀࡿ㸬 95 ༊⏬ 51 ࡢ⏘༸ᗋࡣ㸪2008 ᖺ༊⏬ 50 ࡢୖὶ➃ฟ⌧ࡋࡓ᪂ࡋ࠸᪩℩ࡢୖὶഃ ࠶ࡿᖹ℩㒊ศᙧᡂࡉࢀ࡚࠸ࡓ㸬ࡇࡢሙᡤࡣ㸪2008 ᖺ 3 ᭶ࡣ᭱Ỉ῝ 1.7 m ࡢῡ࡛࠶ࡗࡓࡀ㸪2008 ᖺࡢ⛅ࡲ࡛ࡢ㛫㸪♟ࡀ㔞ሁ✚ࡍࡿࡇ࡛᪩℩ኚ ࡋࡓ㸬ὶ㊰యࢆᇙࡵࡓ⢏ᚄ 5 cm ๓ᚋࡢ♟ࡼࡾ㸪᪩℩ࡢୖὶ➃ࡢỈ῝ࡣ⣙ 0.2 m ὸࡃ࡞ࡾ㸪Ἑᗋࡣㄪᰝ⪅ࡢ㊊ࡀ 20–30 cm ỿࡴᰂࡽࡃ㸪࣐ࢦࡢ ⏘༸ሙࡋ࡚ඃࢀ࡚࠸ࡓ᥎ ࡉࢀࡿ㸦ᮌᮏ㸪ᮍⓎ⾲㸧㸬ࡇࡢࡼ࠺㸪✺↛ ⌧ࢀࡓ᪩℩ࡢୖὶഃ࡛ᐇ㝿࣐ࢦࡀ⏘༸ࡋࡓࡇࢆ⪃៖ࡍࡿ㸪ᮏㄪᰝỈᇦ ࡛ࡣ㸪ேᕤ⏘༸ሙ㐀ᡂ➼㸦୰ᮧ㸪1999b㸧ࡼࡗ࡚⏘༸ዲ㐺࡞⎔ቃࢆᩚഛࡍࡿ ࡇ࡛㸪⮬↛⏘༸ࢆㄏⓎ࡛ࡁࡿ⪃࠼ࡽࢀࡿ㸬ࡓࡔࡋ㸪୰ᮧ㸦1999b㸧ࡼࡿே ᕤ⏘༸ሙ㐀ᡂᡭἲࡣ㸪࣡ࢼࡀ⏘༸ࡍࡿᑠᨭὶࢆᑐ㇟ࡋ࡚㛤Ⓨࡉࢀࡓᢏ⾡࡛ ࠶ࡿ㸬࣐ࢦࡸ࣐࣓ࣖࡣ㸪࣡ࢼẚ࡚Ἑᕝࡢᮏὶ࡛⏘༸ࡍࡿഴྥࡀᙉ࠸ࡇ ࡽ㸦୰ᮧ㸪1999a㸧㸪ᚋࡣ㸪ᮏㄪᰝỈᇦࡢࡼ࠺࡞Ἑᕝࡢᮏὶ࠾࠸࡚ࡶ㸪 ேᕤ⏘༸ሙࡢ㐀ᡂᢏ⾡ࢆ㛤Ⓨࡍࡿᚲせࡀ࠶ࡿ⪃࠼ࡽࢀࡿ㸬 ➨ 4 ❶࡛ࡣ㸪ࡇࢀࡽࡢᾋୖ⛶㨶ࡀὶୗࡋࡓ┤᥋ⓗཎᅉࡋ࡚㸪ᾋୖࡋࡓሙᡤ ᪩࠸ὶࢀࢆ⦆ࡍࡿ▼ࡸ᳜⏕➼ࡢᵓ㐀ࡀ࡞ࡗࡓࡇࢆᣲࡆࡓ㸬ࡇࡢ᥎ᐃࡀ ṇࡋ࠸௬ᐃࡍࡿ㸪࣐ࢦᾋୖ⛶㨶ࡀὶୗ⛣ືࡍࡿ࠺ࡣ㸪ᾋୖ⛶㨶ࡢ ࡢၥ㢟࡛ࡣ࡞ࡃ㸪ぶ㨶ࡢ⏘༸ሙᡤ㑅ᢥつᐃࡉࢀࡿࡇ࡞ࡿ㸬ࢱࢭࣚ࢘ ࢧࢣ࡛ண ࡉࢀࡿࡼ࠺㸪ࢧࢣ⛉㨶㢮ᾋୖ⛶㨶ࡢὶୗ⛣ືࡀ㐺ᛂᗘࢆୗࡆࡿ⾜ ື㸦Foldvik et al., 2010㸹Einum et al., 2011, 2012㸧࡛࠶ࡿ࡞ࡽ㸪ぶ㨶ࡢ⏘༸ሙᡤ 㑅ᢥࡣ⛶㨶ࡢὶୗ⛣ືࢆᅇ㑊ࡍࡿᶵᵓࡀᏑᅾࡍࡿࡶࡋࢀ࡞࠸㸬ࡋࡋ㸪ࢧ ࢣ⛉㨶㢮ࡢ⏘༸ሙᡤ㑅ᢥ㛵ࡋ࡚㸪ᾋୖ⛶㨶ࡢ⏕ᜥሙᡤࡀᇶ‽࡞ࡿ࠸࠺ሗ ࿌ࡣ࡞࠸㸦࠼ࡤ㸪ⓑ▼㸪1957㸹ᮌᮧ㸪1972㸹Geist and Dauble, 1998㸹୰ᮧ㸪 1999a㸹Mull and Wilzbach, 2007㸹Cram et al., 2012㸧 㸬௬㸪࣐ࢦࡢ⏘༸ሙᡤ㑅 ᢥ࡛ࡣ㸪ᮏ᮶ⓗᾋୖ⛶㨶ࡢ⏕ᜥሙᡤࡀ⪃៖ࡉࢀ࡞࠸ࡍࡿ㸪ᾋୖ⛶㨶ࡢὶ ୗ⛣ືࡣ㸪⏘༸ᗋୗὶഃࡢ⎔ቃၥ㢟ࡀ࠶ࡿࡓࡵ⏕ࡌࡿ࠸࠼ࡿ㸬ᮏㄪᰝỈ ᇦࡣ㸪ேⅭᨵኚࡢᙳ㡪ࡀ᥎ᐃࡉࢀࡓ㨶㢮⩌㞟ࡢ⛣⾜ᖏෆ㒊⨨ࡍࡿࡇࡽ 㸦➨ 2 ❶㸧㸪 ⏘༸ᗋୗὶഃࡢ⎔ቃࡀຎࡋ࡚࠸ࡓྍ⬟ᛶࡶ⪃࠼ࡽࢀࡿ㸬 ࡲࡓ㸪 2008 ᖺ༊⏬ 50 ࡢୖὶ➃ฟ⌧ࡋࡓ᪂ࡋ࠸᪩℩ࡣ㸪2009 ᖺ⛅ࡣ♟ࡢὶኻࡼࡾ 96 ᾘ⁛ࡋ࡚࠸ࡓ㸦ᮌᮏ㸪 ᮍⓎ⾲㸧㸬ࡇࡢࡼ࠺࡞୍ⓗ࡞᪩℩ࡸேᕤ⏘༸ሙ࡛ࡣ㸪 㛗ᮇ㛫ࢃࡓࡗ࡚Ꮡᅾࡋ࡚࠸ࡿ⮬↛ࡢ᪩℩ࡣ࿘㎶ࡢ⎔ቃ㸦᳜≀᳜⏕ࡸ▼ࡢሁ ✚➼㸧ࡀ␗࡞ࡿࡢࡶࡋࢀ࡞࠸㸬ࡇࡢሙྜࡣ㸪➨ 4 ❶࡛᥎ᐃࡉࢀࡓ࠾ࡾ㸪 ⏘༸ᗋୗὶഃࡢ᪩℩㏆᪩࠸ὶࢀࢆ⦆ࡍࡿᒾࡸ᳜⏕➼ࡢᵓ㐀ࢆ㐀ᡂࡍࡿࡇ ࡀ᭷⏝࡛࠶ࡿ㸬᪉㸪ᾋୖ⛶㨶ࡢ⏕ᜥሙᡤࡀ⏘༸ሙᡤࡢ㑅ᢥᇶ‽࡞ࡿࡍ ࢀࡤ㸪ࡑࡢሙᡤ⏘༸ࡋࡓぶ㨶ࡢ⾜ືၥ㢟ࡀ࠶ࡿࡇ࡞ࡿ㸬➨ 5 ❶࡛♧ࡋ ࡓࡼ࠺㸪ᮏㄪᰝỈᇦ࡛ࡣ㣴Ṫ㨶ࡀᨺὶࡉࢀ࡚࠸ࡿࡇࡽ㸦Kimoto et al., ༳ ๅ୰㸧㸪 㠀㐺ᛂⓗ࡞ᨺὶಶయࡀ⏘༸ࡋ࡚࠸ࡓྍ⬟ᛶࡶ⪃࠼ࡽࢀࡿ㸬 ࡇࡢሙྜࡶ㸪 ୖグྠᵝ⏕ᜥሙᡤࡢ㐀ᡂࡣ᭷ຠ࡛࠶ࡿࡀ㸪㛗ᮇⓗࡣ㸪ࡇࡢሙ㐺ᛂࡋࡓ ᅾ᮶ಶయࡼࡿ⮬↛⏕⏘ࢆಁ㐍ࡍࡁ⪃࠼ࡽࢀࡿ㸬ᚋࡣ㸪࣐ࢦぶ㨶ࡢ ⏘༸ሙᡤ㑅ᢥᾋୖ⛶㨶ࡢ⏕ᜥሙᡤࡢ㛵ಀࢆㄪࡿᚲせࡀ࠶ࡿࡔࢁ࠺㸬 ࣐ࢦಶయ⩌ࡢ㑇ఏⓗᵓ㐀 ➨ 5 ❶࡛ࡣ㸪⚄ཎᕝᨭὶἼᮌྜᕝ⏕ᜥࡍࡿ 3 ࡘࡢ࣐ࢦ㝸㞳ಶయ⩌㸦⚄ ཎᕝ※ὶ㒊ࡢ࣐ࢦ㸪Ἴᮌྜᕝࡲࢇࡾࡻ࠺㇂ࡢ࣐ࢦ㸪Ἴᮌྜᕝ࣓ࣥࣀࢶࣛ㇂ ࡢ࣓࣡㸧ࡢ㑇ఏⓗᵓ㐀ࢆ mtDNA ศᯒࡼࡾㄪࡓ㸬ᚓࡽࢀࡓ 11 ✀㢮ࡢࣁࣉ ࣟࢱࣉࡢ࠺ࡕ Hap-1 ࡣࡍ࡚ࡢ㝸㞳ಶయ⩌ぢࡽࢀ㸪࣓࡛࣡ᅛᐃ㸦100%㸧㸪 ⚄ཎᕝ※ὶ㒊ࡢ࣐ࢦ࡛ඃ༨ࡋࡓ㸦90.5%㸧㸪⚄ཎᕝయ࡛᥇㞟ࡋࡓ࣐ࢦ ࡢᾋୖ⛶㨶࡛ࡶඃ༨ࡋ࡚࠸ࡓ㸦62.8%㸧ࠋ୍᪉㸪ࡲࢇࡾࡻ࠺㇂ࡢ࣐ࢦ࡛ࡣ Hap-1 ࡣᑡ࡞ࡃ㸦10.0%㸧㸪㐣ཤᨺὶࡉࢀࡓ᥎ᐃࡉࢀࡿ㣴㨶ሙࡢ࣐ࢦࡢࣁࣉࣟࢱ ࣉࡀඃ༨ࡋ࡚࠸ࡓ㸬௨ୖࡢ⤖ᯝࡽ㸪⚄ཎᕝ※ὶ㒊ࡢ࣐ࢦ࣓ࣥࣀࢶࣛ㇂ ࡢ࣓࣡ࡣᅾ᮶ಶయ⩌࡛࠶ࡾ㸪ࡲࢇࡾࡻ࠺㇂ࡢ࣐ࢦࡣ㣴Ṫ㨶ࡰ⨨ࡉࢀ ࡚࠸ࡿ᥎ ࡉࢀࡓࠋࡇࢀࡽᇶ࡙࠸࡚ 3 㝸㞳ಶయ⩌ࡑࢀࡒࢀࢆ༢ࡍࡿಖ ࢆᥦࡋ㸪⁺ᴗ㛵㐃つ๎ࢆᨵṇࡋࡓࠋ ⚄ཎᕝ※ὶ㒊ࡢ⁺ᴗ㛵㐃つ๎ࡢᨵṇ࡛ࡣ㸪୍ྜ┠ࡽୖὶᇦ࠾࠸࡚⁺ ᴗ࣭㐟⁺࠾ࡼࡧᨺὶࢆ⚗Ṇࡋࡓࡀ㸪୍ྜ┠ࡽୖὶࡢ࣐ࢦಶయ⩌ࡣ㸪 㠀ᅾ᮶ணࡉࢀࡿࣁࣉࣟࢱࣉ㸦Hap-4㸧ࢆᣢࡘಶయࡀΰᅾ㸦9.5%㸧ࡋ࡚࠸ ࡓ㸬୍ྜ┠ࡢୖὶࡣ୕ྜ┠㸦ⴠᕪ 10 m㸧ྜ┠㸦ⴠᕪ 11 m㸧ࡀ࠶ 97 ࡾ㸪Hap-4 ࡣ୍ྜ┠୕ྜ┠ࡢ㛫ࡔࡅ᳨࡛ฟࡉࢀࡓ㸬ࡋࡓࡀࡗ࡚㸪ཝᐦ࡞ ᅾ᮶ಶయ⩌⏕ᜥỈᇦࡣ୕ྜ┠ࡼࡾୖὶ࠸࠼ࡿࡀ㸪୕ྜ┠ࡣⓏᒣ㐨ࡽ㞳 ࢀ࡚࠸ࡿࡓࡵᐦ⁺ࡢ┘どࡀᅔ㞴ணࡉࢀࡓ㸬 ୍᪉㸪୍ྜ┠ࡣᯘ㐨ࡽ㏆ࡃ㸪 ཱྀ㏆ࡣ♽ẕᒣࡢⓏᒣᐈ⏝ࡢ㥔㌴ሙࡀᩚഛࡉࢀ࡚࠸ࡿࡓࡵ㸪┘どࢆ⾜࠸ࡸ ࡍ࠸㸬ࡇࢀࡽࡢ⌮⏤ࡼࡾ㸪୍ྜ┠ࡽୖὶᇦࢆಖᑐ㇟Ỉᇦࡋࡓ㸬 Hap-4 ࡣᾋୖ⛶㨶ࡽࡶ᳨ฟࡉࢀࡓࡀ㸪ࡢࣁࣉࣟࢱࣉ␗࡞ࡾ㸪୍ྜ┠ ࡽ⣙ 1.5 km ୗὶࡲ࡛ࡢ⊃࠸⠊ᅖぢࡽࢀ㸪୍ྜ┠ࡢ┤ୗ࡛᭱ࡶ㧗࠸㢖ᗘ 㸦60%㸧ࢆ♧ࡋࡓ㸬ࡇ࠺ࡋࡓ㝈ࡽࢀࡓศᕸ୍ྜ┠ࡢࡑࡤ 1990 ᖺ௦ࡲ࡛㣴 㨶ሙࡀ✌ാࡋ࡚࠸ࡓࡇ㸦㊊❧ᚭಙẶ㹙㣴㨶ሙ Ỉⱌ㹛 㸪⚾ಙ㸧ࢆ⪃៖ࡍࡿ㸪 Hap-4 ࡣࡇࡢ㣴㨶ሙࡢ㣫⫱㨶⏤᮶ࡍࡿྍ⬟ᛶࡀ㧗࠸⪃࠼ࡽࢀࡿ㸬ᮏ◊✲࡛ ࡣ㸪㠀ᅾ᮶ணࡉࢀࡿ Hap-4 ࢆྵࡵࡓಖࡀ㸪㔝ᕝ⁺ᴗ༠ྠ⤌ྜ㸪➉⏣ ᕷ㸪ᆅඖ㐟⁺⪅ᅋయ➼ᢎࡉࢀ㸪⚗⁺༊ࡀタ⨨ࡉࢀࡓ㸦ᮌᮏ㸪2008㸧 㸬ᚋࡣ㸪 ⚗⁺༊ෆ࠾ࡅࡿ Hap-4 ࡢືែࢆࣔࢽࢱࣜࣥࢢࡍࡿᚲせࡀ࠶ࡿ㸬 ⚄ཎᕝࡢ※ὶ㒊ࡣ㸪ࡇࢀࡲ࡛⚗⁺ࡢᑐ㇟ᣦᐃࡉࢀࡓࡇࡀ࡞ࡃ㸪ࡋࡶୖ グࡢ࠾ࡾୗὶ➃㌴㐨ࡸ㥔㌴ሙࡀᩜタࡉࢀ࡚࠸ࡿࡇࢆ⪃៖ࡍࡿ㸪ᨺὶࡢ ᙳ㡪ࢆࢇཷࡅ࡚࠸࡞ࡗࡓࡇࡣወ㊧ⓗ࡞ᖾ㐠࠸࠼ࡿ㸬ᚋࡣ㸪ศ ┴ෆࡢἙᕝຍ࠼࡚㸪⏕≀ᆅ⌮Ꮫⓗ㛵ࢃࡾࡀ᥎ᐃࡉࢀࡿ୰ᅜ࣭ᅄᅜᆅ᪉ࡢ℩ ᡞෆഃす㒊ᆅᇦ⏕ᜥࡍࡿ࣐ࢦࡘ࠸࡚ࡶ㑇ఏᏛⓗ࡞ゎᯒࢆ⾜࠸㸪⣔⤫ⓗ ⨨ࡅࢆ᫂ࡽࡍࡿᚲせࡀ࠶ࡿࡔࢁ࠺㸬୍᪉㸪ࡇࢀࡲ࡛ḷᒣ┴ྂᗙᕝ 㸦Kawamura et al., 2007㸧㧗▱┴ዉ༙ᕝ㸦ᒸ㒊࣭ᑠᯇ㸪2012㸧ࡽᅾ᮶࣐ ࢦಶయ⩌ࡀሗ࿌ࡉࢀ࡚࠾ࡾ㸪ᮏ◊✲ࡣ๓⪅ྠࡌ mtDNA ࡢ㡿ᇦࢆゎᯒࡋࡓࡓ ࡵ⤖ᯝࡢẚ㍑ࡀྍ⬟࡛࠶ࡗࡓࡀ㸪ᚋ⪅ࡣ㡿ᇦࡀ␗࡞ࡗࡓࡓࡵẚ㍑࡛ࡁ࡞ࡗ ࡓ㸬ᮏ◊✲ Kawamura et al.㸦2007㸧࡛ゎᯒࡋࡓ㡿ᇦ㸦D-loop ND1㸧ࡣ㸪ሷ ᇶ㓄ิࡢỴᐃࡀᅔ㞴࡛࠶ࡾ㸪࠶ࡲࡾ࠸ᡭࡢⰋ࠸ゎᯒ㒊ࡣ࠸࠼࡞࠸㸦⡿ ຍ⏣ ᚭẶ㹙ᅜ❧◊✲㛤ⓎἲேỈ⏘⥲ྜ◊✲ࢭࣥࢱ࣮ቑ㣴Ṫ◊✲ᡤ㹛 㸪⚾ಙ㸧㸬 ᚋ㸪ᅜෆࡢᅾ᮶࣐ࢦಶయ⩌ࡢࢹ࣮ࢱ࣮࣋ࢫࢆᵓ⠏ࡍࡿሙྜࡣ㸪ᒸ㒊࣭ᑠᯇ 㸦2012㸧ࡀ⏝࠸ࡓ㡿ᇦ㸦ND5 Cb㸧ࡶ⪃៖ࡍࡁ⪃࠼ࡽࢀࡿ㸬 Ἴᮌྜᕝࡲࢇࡾࡻ࠺㇂ࡢ࣐ࢦಶయ⩌ࡀࡰ㣴Ṫ㨶⨨ࡉࢀࡓྍ⬟ᛶࡀ♧ 98 ࡉࢀࡓࡇࡽ㸪⚄ཎᕝ※ὶ㒊ࡢ࣐ࢦಶయ⩌ࡣ㸪༢㔝ᕝỈ⣔ࡢ୍ᨭὶࡢ ᅾ᮶࣐ࢦಶయ⩌࡛࠶ࡿࡔࡅ࡛࡞ࡃ㸪࣓࣡᭱ࡶ㏆࠸㛵ಀ࠶ࡿ࣐ࢦಶయ ⩌ࡋ࡚ᕼᑡᛶࡀ㧗࠸⪃࠼ࡽࢀࡿ㸬ᮏ◊✲࡛ࡣ㸪ᅾ᮶ಶయ⩌ࢆ᥎ᐃࡍࡿࡓࡵ mtDNA ࡢሗࡔࡅࢆ⏝࠸ࡓࡀ㸪ᚋ㸪࣐ࢦ࣓࣡ࡢ㑇ఏⓗ㛵ಀࢆ᫂ࡽ ࡍࡿࡓࡵࡣ㸪 ࡉࡽሗ㔞ࡢከ࠸࣐ࢡࣟࢧࢸࣛࢺࡸ AFLP ➼㸪 ᰾ DNA ࢆ⏝࠸ࡓゎᯒࢆᐇࡍࡿᚲせࡀ࠶ࡿ㸦Kawamura et al., 2007㸹Kubota et al., 2007㸹 Kikko et al., 2008㸹Sato et al., 2010㸧 㸬୍᪉㸪Kano et al.㸦2010㸧ࡣᅜෆࡢ࣓࣡ ಶయ⩌ࡢ⌧≧ࢆㄪ㸪ࡇࢀࡲ࡛ሗ࿌ࡉࢀࡓ 7 ಶయ⩌ࡢ࠺ࡕ㸪4 ಶయ⩌ࡣ㔞 ࡢ㣴Ṫ㨶ࡢᨺὶࡸ⮬↛⅏ᐖ➼࡛ࡰ⤯⁛ࡋࡓࡇ㸪Ᏻᐃⓗ࡞ಶయ⩌ࡣᮏㄪᰝỈ ᇦࡢ࣓ࣥࣀࢶࣛ㇂୕㔜┴ဨᘚᕝࡢ୕ᅜ㇂ࡔࡅᏑᅾࡍࡿࡇࢆሗ࿌ࡋ࡚࠸ࡿ㸬 ࡓࡔࡋ㸪ဨᘚᕝࡢ࣓࣡ࡣ༢⊂⏕ᜥᇦࢆᣢࡓࡎ࣐ࢦඹᏑࡋ࡚࠸ࡿࡇ 㸦Kano et al., 2006㸧 㸪2007 ᖺኳ↛グᛕ≀ᣦᐃࡉࢀࡓᚋࡶ㐟⁺㛤ᨺࡉࢀ࡚ ࠸ࡿࡇ㸦Kano et al., 2010㸧ࡽ㸪㣴Ṫࡉࢀࡓ࣐ࢦࡀ࣓࣡⏕ᜥᇦᨺὶࡉ ࢀ㸪ᅾ᮶ࡢ࣐ࢦࡸ࣓࣡㞧ࡍࡿ㸪ࡲࡓࡣ㞧ࡋࡓྍ⬟ᛶࢆ㝖࡛ࡁ࡞࠸㸬 㣴Ṫ㨶ࡢ㞧ࡀ㐍ࡵࡤ㸪ဨᘚᕝࡢ࣓࣐࣡ࢦࡢ㑇ఏⓗ㛵ಀ㸪ࡉࡽࡣ 㔝ᕝࡢ࣓࣡ࡢ㑇ఏⓗ㛵ಀࢆㄪࡿࡇࡣᴟࡵ࡚ᅔ㞴࡞ࡿࡇࡽ㸪ဨᘚ ᕝࡢ࣓࣡ಶయ⩌ࡢ᪩ᛴ࡞㑇ఏⓗᵓ㐀ࡢㄪᰝබⓗ࡞⚗⁺ᥐ⨨ࡀᮃࡲࢀࡿ㸬 ⤖ ㄽ ௨ୖࢆࡲࡵࡿ㸪➨ 2 ❶㸦ὶ⛬ἢࡗࡓ㨶㢮⩌㞟ࡢኚ㸧➨ 5 ❶㸦࣐ ࢦಶయ⩌ࡢ㑇ఏⓗᵓ㐀㸧ࡣ㸪ࡑࢀࡒࢀಶูࡢ⤖ᯝࢆ♧ࡋ࡞ࡀࡽ㸪➨ 3 ❶㸦࣐ ࢦᾋୖ⛶㨶ࡢὶ⛬ศᕸ㸧➨ 4 ❶㸦࣐ࢦᾋୖ⛶㨶ࡢ⏕ᜥሙᡤ⏝㸧࡛ぢࡽࢀ ࡓୗὶഃỈᇦ࠾ࡅࡿᾋୖ⛶㨶ࡢศᕸࡢไ㝈せᅉࡋ࡚㸪ࡑࢀࡒࢀேⅭᨵኚࡢ ᙳ㡪㸦➨ 2 ❶㸧㣴Ṫ㨶ࡢᨺὶࡢᙳ㡪㸦➨ 5 ❶㸧ࢆ♧၀ࡍࡿᵓ㐀࡞ࡗ࡚࠸ࡿ㸬 ࡋࡋ㸪⌧Ⅼ࡛ࡣ㸪ࡇࢀࡽࡀ┿⮬↛⏕⏘ࡢไ㝈せᅉ࡛࠶ࡿྰࢆ᫂ࡽ ࡍࡿࡇࡣᅔ㞴࡛࠶ࡿ㸬ࡑࢀࡣ㸪ẚ㍑ᑐ↷ࡍࡿࡁ◁㜵ࢲ࣒ࡸᨺὶࡢᙳ㡪 ࡀ࡞࠸ἙᕝࡀᏑᅾࡋ࡞࠸ࡓࡵ࡛࠶ࡾ㸪ࡲࡓ㸪ࡑࢀࡽࡢᙳ㡪ࡀ⏕ࡌࡿ๓ࡢ⚄ཎᕝ ࡸ⥴᪉ᕝࡢ≧ែࡀᐃ㔞ⓗ࡞ሗࡋ࡚ṧࡗ࡚࠸࡞࠸ࡓࡵ࡛ࡶ࠶ࡿ㸬ࡓࡔࡋ㸪ᐃ 99 ᛶⓗ࡞ሗ࡛ࡣ㸪ᮏㄪᰝỈᇦࡢ᭱ୗὶ㒊࡛࣐ࢦࡢ⮬↛⏘༸ࡀ㢖⦾ぢࡽࢀ࡚ ࠸ࡓࡇࡽ㸪୧せᅉࡀఱࡽࡢᝏᙳ㡪ࢆ࠼࡚࠸ࡿྍ⬟ᛶࡣྰᐃ࡛ࡁ࡞࠸㸬 ௨ୖࡽ㸪ᮏㄪᰝỈᇦࡢ࣐ࢦಶయ⩌ࡢ⌧≧ࡋ࡚௨ୗࡢ⤖ㄽࡀᚓࡽࢀࡓ㸬 1㸧⛣⾜ᖏࢆྵࡴୗὶഃỈᇦࡣ㸪ேⅭᨵኚࡢᝏᙳ㡪ࢆཷࡅ࡚࠸ࡿྍ⬟ᛶࡀ࠶ࡿ㸬 2㸧࣐ࢦ⏕ᜥᇦࡢ 7 ௨ୖࡣ㸪⛣⾜ᖏࢆྵࡴୗὶഃỈᇦྵࡲࢀࡿ㸬 3㸧࣐ࢦࡢ⮬↛⏕⏘ࡣ⛣⾜ᖏࡼࡾୖὶ࡛Ⰻዲ㸪ࡑࡢୗὶ࡛ࡣపㄪ࡛࠶ࡿ㸬 4㸧ᅾ᮶࣐ࢦಶయ⩌ࡣ⚄ཎᕝࡢ୍ྜ┠ࡼࡾୖὶἼᮌྜᕝࡢ࣓ࣥࣀࢶࣛ㇂ 㸦࣓࣡㸧⏕ᜥࡋ࡚࠸ࡿ㸬 5㸧ࡲࢇࡾࡻ࠺㇂࡛ࡣ㸪ࣄ࣮ࣂࢳࡢࡢୗὶഃࡣࡰ㣴Ṫ㨶ࡰ⨨ࡁࢃࡗ࡚ ࠸ࡿࡀ㸪ࡢୖὶഃࡣᅾ᮶࣐ࢦಶయ⩌ࡀ⏕ᜥࡋ࡚࠸ࡿྍ⬟ᛶࡀ࠶ࡿ㸬 6㸧⛣⾜ᖏࢆྵࡴୗὶഃỈᇦࡣ㠀ᅾ᮶ࡢ࣐ࢦࡀ⏕ᜥࡋ࡚࠸ࡿྍ⬟ᛶࡀ࠶ࡿ㸬 ࡇࢀࡽᇶ࡙ࡁ㸪ಖ᪉ἲ㸦ࢰ࣮ࢽࣥࢢ⟶⌮㸧ࡋ࡚௨ୗࡀᑟฟࡉࢀࡓ㸬 1㸧⚄ཎᕝࡢ୍ྜ┠ࡽୖὶᇦ࠾࠸࡚㸪⁺ᴗ㐟⁺࠾ࡼࡧᨺὶࢆ⚗Ṇ㸬 2㸧Ἴᮌྜᕝࡢ⌧⾜⚗⁺༊ࡢୖ㝈ࢆ᧔ᗫࡋ㸪ⓑỈᶫࡽୖὶᇦ࠾࠸࡚㸪⁺ᴗ 㐟⁺࠾ࡼࡧᨺὶࢆ⚗Ṇ㸬 3㸧୧⚗⁺༊ࡼࡾୗὶഃ࡛ࡣ⁺ᴗ㐟⁺࠾ࡼࡧᨺὶࢆྍ⬟ࡍࡿ㸬 4㸧ࡓࡔࡋ㸪⛣⾜ᖏࢆྵࡴୗὶഃỈᇦ࡛ࡣ㸪ேⅭᨵኚࡸᨺὶࡢᙳ㡪ࡼࡾ࣐ࢦ ࡢ⮬↛⏕⏘ࡀ㜼ᐖࡉࢀ࡚࠸ࡿྍ⬟ᛶࡀ࠶ࡿࡓࡵ㸪ᾋୖ⛶㨶ࡢ⏕ᜥሙᡤᩚഛࡸ ᨺὶࡢぢ┤ࡋࡼࡾ㸪⮬↛⏕⏘ࡢಁ㐍ࢆヨࡳࡿᚲせࡀ࠶ࡿ㸬 ➹⪅ࡽࡣ㸪ࡇࢀࡽࡢࢰ࣮ࢽࣥࢢࡢᑟධࡢྍྰࢆ㸪ᆅඖఫẸ㸪㔝ᕝ⁺ᴗ༠ ྠ⤌ྜ㸪㐟⁺⪅㸪ᆅඖ⾜ᨻ㸦➉⏣ᕷ㸧➼࡛⤌⧊ࡋࡓጤဨ࠾࠸࡚༠㆟ࡋࡓ㸬 ༠㆟ෆᐜࡢヲ⣽ࡣᮌᮏ㸦2008㸧ㆡࡿࡀ㸪⤖ㄽࡋ࡚ࡇࢀࡽࡢࡣཷࡅධࢀࡽ ࢀࡓ㸬ࡇࡢ࠺ࡕ㸪1㸧ࡣ㔝ᕝ⁺ᴗ༠ྠ⤌ྜࡢつ๎㸦⁺ᴗᶒ⾜つ๎㐟⁺つ๎㸧 ࡢ᪂タࢆ㸪2㸧ࡣ㔝ᕝ⁺ᴗ༠ྠ⤌ྜࡢつ๎ศ┴ࡢ⁺ᴗㄪᩚつ๎ࡢᨵṇ࣭᪂ タࢆᚲせࡋࡓ㸬㛵ಀ⪅ࡢດຊࡼࡾ㸪㔝ᕝ⁺ᴗ༠ྠ⤌ྜࡢつ๎ࡣ 2008 ᖺ 7 ᭶ 23 ᪥㸪ศ┴ࡢ⁺ᴗㄪᩚつ๎ࡣ 2009 ᖺ 10 ᭶ 16 ᪥࡛ኚ᭦ࡉࢀࡓ㸬 ⌧ᅾ㸪ࡇࡢࡼ࠺ࡋ࡚ᨵṇ࣭᪂タࡉࢀࡓ⁺ᴗ㛵㐃つ๎ᇶ࡙ࡁ㸪㔝ᕝ࠾ ࠸࡚࣓࣡ࢆྵࡴ࣐ࢦࡢಖࡀᐇࡉࢀ࡚࠸ࡿ㸦Fig. 6-1㸧㸬 100 Fig. 6-1. A billboard notifing the fishing and stocking-prohibited area above Ichigome Falls of the Kohbaru Stream. This area was established in July 2008 on the basis of this study. 101 せ ⣙ ➨ 1 ❶ ㅖゝ ࣐ࢦ Oncorhynchus masou ishikawae ࡣࢧࢣ⛉ᒓࡍࡿ᪥ᮏᅛ᭷ࡢள✀࡛࠶ ࡾ㸪⚄ዉᕝ┴௨すࡢᮏᕞኴᖹὒᓊ㸪ᅄᅜᇦ㸪ᕞࡢ℩ᡞෆᾏὶධἙᕝ⮬↛ ศᕸࡍࡿ㸬ศᕸ༡㝈ᇦ࡛࠶ࡿᕞࡣἙᕝᆺಶయ⩌ࡔࡅࡀ⏕ᜥࡋ࡚࠸ࡿ㸬࣐ ࢦࢆྵࡴᅾ᮶ࢧࢣ⛉㨶㢮࡛ࡣ㸪✀ⱑ⏕⏘ᢏ⾡ࡀ☜❧ࡉࢀࡓ 1970 ᖺ௦௨㝆㸪࠾ࡶ 㣴Ṫ㨶ࡢᨺὶࡼࡗ࡚㈨※ࡢ⥔ᣢ࣭ቑṪࡀ࡞ࡉࢀ㸪ຠᯝࢆୖࡆ࡚ࡁࡓ㸬ࡋ ࡋ㏆ᖺ㸪ᨺὶࡉࢀࡓ㣴Ṫ㨶ࡀ㔝⏕ಶయ⩌ࡢ୰㐺ᛂⓗ࡛࡞࠸㑇ఏᏊࢆὶධࡉࡏ㸪 ␗⣔㓄ᙅໃࡸᒁᡤ㐺ᛂࡢᔂቯࢆࡶࡓࡽࡍྍ⬟ᛶࡀ᫂ࡽࡉࢀ࡚࠸ࡿ㸬 ࡲࡓ㸪 㣴Ṫ㨶ࡢᨺὶࡀ㛗ᮇࢃࡓࡾ⥅⥆ࡉࢀࡓ⤖ᯝ㸪࣐ࢦࢆྵࡴᅜෆࡢἙᕝᛶࢧࢣ ⛉㨶㢮ࡢᅾ᮶ಶయ⩌ࡣ⤯⁛ࡢ༴ᶵ࠶ࡾ㸪⮬↛ࡸࢲ࣒࡛㝸㞳ࡉࢀࡓἙᕝ※ὶ 㒊ࡔࡅṧᏑࡋ࡚࠸ࡿ㸬ࡇࡢࡼ࠺㸪ᨺὶࡣ࣐ࢦࢆྵࡴ㔝⏕ࡢἙᕝᛶࢧࢣ⛉ 㨶㢮ࡢᏑ⥆ᝏᙳ㡪ࢆ࠼ࡿྍ⬟ᛶࡀ㧗࠸ࡇࡽ㸪㛗ᮇⓗ࡞ಶయ⩌ࡢಖࡢ ࡓࡵࡣᐇࡍࡁ࡛࡞࠸ࡀ㸪⁺ᴗ࣭㐟⁺ࡼࡿ▷ᮇⓗ࡞⏝ࡢࡓࡵࡣᚲせ ࡛࠶ࡿ㸬㏆ᖺ㸪 Ἑᕝᛶࢧࢣ⛉㨶㢮ࡢಖ⏝ࡢ୧❧ࢆᅗࡿࡓࡵࡢ᪉ἲࡋ࡚㸪 ᅾ᮶ಶయ⩌ࡢ⏕ᜥỈᇦ⮬↛⏕⏘ࡢᐇែᇶ࡙࠸࡚Ἑᕝࢆ༊ศࡋ㸪Ỉᇦࡈ ᨺὶ⁺ᴗ࣭㐟⁺⟶⌮ᡭἲࢆ࠸ศࡅࡿࢰ࣮ࢽࣥࢢࡀὀ┠ࡉࢀ࡚࠸ࡿ㸬ᮏ◊ ✲࡛ࡣ㸪ᅾ᮶࣐ࢦಶయ⩌ࡀ⏕ᜥࡍࡿྍ⬟ᛶࡢ㧗࠸ศ┴㔝ᕝỈ⣔⚄ཎᕝ ᨭὶἼᮌྜᕝࡢ୍㒊༊㛫࠾ࡼࡧ⚄ཎᕝࡀὶධࡍࡿ⥴᪉ᕝࡢୖὶഃ༊㛫ࢆㄪᰝᑐ ㇟Ỉᇦࡋ࡚㸪ࢰ࣮ࢽࣥࢢ⟶⌮ࢆᑟධࡍࡿࡓࡵᚲせ࡞▱ぢࢆᚓࡿࡓࡵ㸪௨ ୗࡢ 4 ࡘࡢㄪᰝࢆᐇࡋࡓ㸬 ➨ 2 ❶ ὶ⛬ἢࡗࡓ㨶㢮⩌㞟ࡢኚ 㔝ᕝỈ⣔⥴᪉ᕝᨭὶ⚄ཎᕝ࠾ࡼࡧἼᮌྜᕝ⣙ 1 km ࠾ࡁタࡅࡓ 15 ࡢ ㄪᰝⅬ࠾࠸࡚㸪2003 ᖺ 10 ᭶㸪2004 ᖺ 2 ᭶ 8 ᭶⏕ᜥ㨶㢮ࡢ₯Ỉ┠どほᐹ ࢆ⾜࠸㸪㨶㢮⩌㞟⤌ᡂࡢὶ⛬ἢࡗࡓኚᵝᘧࢆㄪࡓ㸬ࡍ࡚ࡢㄪᰝ࡛ 4 ┠ 7 ⛉ 14 ✀㸦ள✀ࢆྵࡴ㸧ࡀ☜ㄆࡉࢀ㸪⥲☜ㄆಶయᩘࡣ 18,015 ಶయ࡛࠶ࡗࡓ㸬 102 㨶㢮⩌㞟ࡣ㸪ୖὶഃㄪᰝⅬࡢ࣐ࢦࢱ࢝ࣁࣖࡀඃ༨ࡍࡿ༢⣧࡞⤌ᡂࡽ㸪ୗ ὶഃㄪᰝⅬࡢ࢘ࢢࡸ࣒࢝࣡ࢶ➼ࡢࢥ⛉㨶㢮ࡀඃ༨ࡍࡿࡼࡾ」㞧࡞⤌ᡂ ᫂☜⛣⾜ࡋࡓ㸬ከኚ㔞ゎᯒࡢ⤖ᯝ㸪15 ࡢㄪᰝⅬࡣ࣐ࢦࢱ࢝ࣁࣖࡀඃ༨ࡍ ࡿୖὶഃㄪᰝⅬ㸦ᶆ㧗 415–820 m㸧 㸪࢘ࢢ㸪࣒࢝࣡ࢶ㸪ࢱ࢝ࣁࣖࡀඃ༨ࡍࡿ⛣ ⾜ᖏࡢㄪᰝⅬ㸦260–397 m㸧࠾ࡼࡧࢥ⛉㨶㢮ࣚࢩࣀ࣎ࣜ㢮ࡀඃ༨ࡍࡿୗὶ ഃㄪᰝⅬ㸦232–255 m㸧ࡢ 3 ࡘࡢࢢ࣮ࣝࣉศ㢮ࡉࢀࡓ㸬ࡉࡽ㸪ࡇࢀࡽࡢ⩌ 㞟⤌ᡂࡢኚᵝᘧࡣ㸪ㄪᰝỈᇦࡢὶ⛬㸦16.3 km㸧ἢࡗ࡚༢ㄪ࡞ኚࢆ♧ࡍ 6 ࡘࡢ⎔ቃኚ㔞㸦Ἑᕝᙧែᆺ㸪ᶆ㧗㸪Ἑᗋ໙㓄㸪ᖹᆒỈ㠃ᖜ㸪ὶ㔞࠾ࡼࡧ᪥ᖹᆒ ᭱పỈ 㸧᭷ព┦㛵ࡋ࡚࠸ࡓࡇࡽ㸪ὶ⛬ἢࡗࡓ⎔ቃ໙㓄ࡁࡃつ ᐃࡉࢀࡿࡶࡢ⪃࠼ࡽࢀࡓ㸬୍᪉㸪ࢥ⛉㨶㢮㸪≉ࢱ࢝ࣁࣖࡢಶయ⩌ᐦᗘࡣ ὶ⛬ἢࡗࡓ༢ㄪ࡞ኚࢆ♧ࡉࡎ㸪◁㜵ሖሐୗὶഃࡢ⛣⾜ᖏ࡛ῶᑡࡋ࡚࠸ࡓ㸬 ࡇ࠺ࡋࡓ㨶✀ูࡢಶయ⩌ᐦᗘࡢኚࡣ㸪ேⅭᨵኚࡼࡿ⏕ᜥ⎔ቃࡢᝏࢆྵࡴ㸪 ࡼࡾᑠࡉ࡞✵㛫ࢫࢣ࣮ࣝࡢ⎔ቃせᅉつᐃࡉࢀࡿࡶࡢ⪃࠼ࡽࢀࡿ㸬 ➨ 3 ❶ ࣐ࢦᾋୖ⛶㨶ࡢὶ⛬ศᕸ 㔝ᕝỈ⣔⥴᪉ᕝᨭὶ⚄ཎᕝタࡅࡓ 14 ࡢㄪᰝ༊㛫㸦⥲ᘏ㛗 16.3 km㸧 ࠾࠸࡚㸪2005 ᖺ 1–4 ᭶ 2 ᅇ㸦11–32 ᪥㛫㝸㸧ࡢ₯Ỉ┠どほᐹࢆ⾜࠸㸪࣐ࢦ ᾋୖ⛶㨶ࡢὶ⛬ศᕸࢆㄪࡓ㸬ᾋୖ⛶㨶ࡢಶయ⩌ᐦᗘࡣୖὶഃࡢ 4 ㄪᰝ༊㛫࡛ 㧗ࡗࡓࡀୗὶഃ 6 ㄪᰝ༊㛫࡛ࡣⴭࡋࡃపࡃ㸪ὶ⛬ศᕸࡢ೫ࡾࡣ 2 ᅇࡢㄪᰝࢆ ㏻ࡌ࡚ኚࡋ࡞ࡗࡓ㸬ࡲࡓ㸪2003 ᖺ 10 ᭶㸪2004 ᖺ 2 ᭶࠾ࡼࡧ 8 ᭶ㄪࡓ ᗂ㨶ᮇ௨㝆ࡢಶయ⩌ᐦᗘࡶୖὶഃ࡛㧗ࡃ㸪Ꮨ⠇ࡈ᭷ព࡞┦㛵ࢆ♧ࡋࡓ㸬ࡉ ࡽ㸪2005 ᖺ 1–4 ᭶࠾ࡅࡿᾋୖ⛶㨶ࡢಶయ⩌ᐦᗘ㸪2004 ᖺ 8 ᭶࠾ࡅࡿᗂ 㨶ᮇ௨㝆ࡢಶయ⩌ᐦᗘࡣ᭷ព࡞┦㛵ࢆ♧ࡋࡓ㸬ࡇࢀࡽࡢ⤖ᯝ㸪᪤ሗࡀᣦࡍ ࡿ࣐ࢦࡢἙᕝᆺಶయ⩌࠾ࡅࡿ⏕άሙᡤ⏘༸ሙᡤࡢ㏆᥋ࢆ⪃៖ࡍࡿ㸪 ᕞࡢἙᕝୖὶᇦ࡛ࡣ㸪࣐ࢦᾋୖ⛶㨶ࡣᑡ࡞ࡃࡶ⣙ 1 ᭶㛫㸪⏘༸ᗋ㏆ ␃ࡲࡿࡶࡢ⪃࠼ࡽࢀࡿ㸬 ➨ 4 ❶ ࣐ࢦᾋୖ⛶㨶ࡢ⏕ᜥሙᡤ⏝ 103 ࣐ࢦ⏕ᜥᇦࡢୗὶഃ࡛࠶ࡿ㔝ᕝỈ⣔⥴᪉ᕝࡢ 1.4 km ༊㛫࠾࠸࡚㸪ὶ㊰ ༢ࢫࢣ࣮ࣝ࠾ࡅࡿᾋୖ⛶㨶ࡢ⏕ᜥሙᡤ⏝ᵝᘧࢆㄪࡓ㸬2007–2009 ᖺࡢ 1–3 ᭶ࡑࢀࡒࢀ 11㸪3㸪5 ᅇࡢ₯Ỉ┠どほᐹࢆ⾜࠸㸪㐃⥆ࡋࡓ 56 ༊⏬㸦ྛ 25 m㸧 ࡢ୧ᓊἢࡗ࡚ᾋୖ⛶㨶ᩘࢆィᩘࡍࡿࡶ㸪ྛ༊⏬ࡢ⎔ቃኚ㔞㸦ὶ㊰༢ ࢱࣉ㸪⏘༸ᗋࡽࡢ㊥㞳㸪ᕝᖜỈ῝ẚ࠾ࡼࡧᕥྑᓊࡢỈ῝㸪ὶ㏿㸪᳜⏕⿕ᗘ㸧 ࢆ ᐃࡋࡓ㸬ᾋୖ⛶㨶ᩘࡣ㸪᪩℩࿘㎶ࡢὶ㊰༢ࢱࣉ᭷ព࡞ṇࡢ┦㛵㸪ἢ ᓊỈ῝࠾ࡼࡧ⏘༸ᗋࡽࡢ㊥㞳᭷ព࡞㈇ࡢ┦㛵ࢆ♧ࡋࡓ㸬ࡇࡢ࠺ࡕ㸪᪩℩࿘ ㎶ࡢὶ㊰༢ࢱࣉࢆ⏝ࡍࡿഴྥࡣୗὶഃỈᇦ≉᭷࡛࠶ࡗࡓ㸬ࡇࢀࡣ㸪ᾋ ୖ⛶㨶㧗࠸࢚ࢿࣝࢠ࣮ຠ⋡ࢆࡶࡓࡽࡍዲ㐺࡞ᚤ⏕ᜥሙᡤ㸦㏿࠸ὶࢀ㏆࠸⦆ ὶ㒊ศ㸧ࡀ㸪ୖὶഃ࡛ࡣ㐢ᅾࡍࡿࡀ㸪ୗὶഃ࡛ࡣ᪩℩㏆ᒁᅾࡍࡿࡓࡵ⪃ ࠼ࡽࢀࡿ㸬୍᪉㸪ὶ㊰༢ࢫࢣ࣮࡛ࣝࡇࢀࡽࡢ᮲௳ࡀ‶ࡓࡉࢀ࡚࠸࡚ࡶ㸪ᚤ⏕ ᜥሙᡤࢫࢣ࣮ࣝࡢ⎔ቃ᮲௳ࡀ༑ศ࡞ሙᡤ࡛ࡣ㸪ᾋୖ⛶㨶ࡀୗὶ᪉ྥ⛣ືࡍ ࡿྍ⬟ᛶࡀ♧ࡉࢀࡓ㸬௨ୖࡢࡇࡽ㸪࣐ࢦ⏕ᜥᇦୗὶഃ࠾࠸࡚ᾋୖ⛶㨶 ࡢಖࡸቑṪࢆᅗࡿࡓࡵࡣ㸪᪩℩࿘㎶ࡢὶ㊰༢ࢱࣉࡢෆ㒊ዲ㐺࡞ᚤ⏕ ᜥሙᡤࢆᩚഛࡍࡿࡇ㸪⎔ቃᐜຊ⛶㨶ࡢቑṪ㔞ࡢࣂࣛࣥࢫࢆ⪃៖ࡍࡿࡇ ࡀ㔜せ⪃࠼ࡽࢀࡿ㸬 ➨ 5 ❶ ࣐ࢦಶయ⩌ࡢ㑇ఏⓗᵓ㐀 㔝ᕝỈ⣔ᨭὶ⚄ཎᕝࡢ᭱ୖὶ㒊⏕ᜥࡍࡿ࣐ࢦࡢ 3 㝸㞳ಶయ⩌㸦↓ᩬᆺ ࡢ࣓࣡ࢆྵࡴ㸧ࢆಖࡍࡿࡓࡵ㸪mtDNA ࢆ⏝࠸࡚㑇ఏⓗᵓ㐀ࢆㄪࡓࠋ⚄ཎ ᕝᨭὶἼᮌྜᕝࡢ࣓ࣥࣀࢶࣛ㇂ࡣ࣓࣡ࡀ⏕ᜥࡋ㸪㞄᥋ࡍࡿࡲࢇࡾࡻ࠺㇂ࡢ ࣐ࢦࡶ⏕ᜥᇦࡢ㒊ศࡀ⚗⁺࡛࠶ࡿࡀ㸪 ᚋ⪅࡛ࡣ 1990 ᖺ௦⚗⁺༊ࡢ ୖὶഃ࡛㠀බᘧ࡞㣴Ṫ㨶ࡢᨺὶࡀ⾜ࢃࢀࡓࡢሗࡀ࠶ࡿ㸬୍᪉㸪⚄ཎᕝ※ὶ 㒊ࡣ㛤ᨺࡉࢀ࡚࠸ࡿࡀ㸪ࡇࢀࡲ࡛㔝ᕝ⁺ᴗ༠ྠ⤌ྜࡼࡿබᘧ࡞ᨺὶグ㘓ࡣ ࡞࠸ࠋᚓࡽࢀࡓ 11 ✀㢮ࡢࣁࣉࣟࢱࣉࡢ࠺ࡕ Hap-1 ࡣ㸪ࡍ࡚ࡢ㝸㞳ಶయ⩌ ぢࡽࢀ㸪࣓࡛࣡ᅛᐃ㸦100%㸧 㸪⚄ཎᕝ※ὶ㒊ࡢ࣐ࢦ࡛ඃ༨㸦90.5%㸧ࡋࡓ 㸪⚄ཎᕝయ࡛᥇㞟ࡋࡓ࣐ࢦᾋୖ⛶㨶࡛ࡶඃ༨ࡋ࡚࠸ࡓ㸦62.8%㸧ࠋ୍᪉㸪 ࡲࢇࡾࡻ࠺㇂ࡢ࣐ࢦ࡛ࡣ Hap-1 ࡣᑡ࡞ࡃ㸦10.0%㸧 㸪㐣ཤᨺὶࡉࢀࡓ᥎ᐃ 104 ࡉࢀࡿ㣴㨶ሙࡢ࣐ࢦࡢࣁࣉࣟࢱࣉࡀඃ༨㸦70%㸧ࡋ࡚࠸ࡓ㸬ࡇࢀࡽࡢ㑇ఏ ⓗᵓ㐀࣓࣡ಶయ⩌ࡢ⌧ᅾࡲ࡛ࡢᏑ⥆ࢆ⪃៖ࡍࡿ㸪࣓ࣥࣀࢶࣛ㇂ࡢ࣓࣡ ⚄ཎᕝ※ὶ㒊ࡢ࣐ࢦࡣᅾ᮶ಶయ⩌࡛࠶ࡿࡀ㸪ࡲࢇࡾࡻ࠺㇂ࡢ࣐ࢦࡣ⚗⁺ ༊ࡢୖὶഃ࡛ᨺὶࡉࢀࡓ㣴Ṫ㨶ࡰ⨨ࡉࢀࡓ᥎ ࡉࢀࡓࠋࡇࢀࡽᇶ࡙ ࠸࡚ 3 㝸㞳ಶయ⩌ྛࠎࢆ༢ࡍࡿಖࢆసᡂࡋ㸪⁺ᴗ㛵㐃つ๎ࢆᨵṇࡋࡓࠋ ➨ 6 ❶ ⥲ྜ⪃ᐹ ➨ 2 ❶㸦ὶ⛬ἢࡗࡓ㨶㢮⩌㞟ࡢኚ㸧➨ 5 ❶㸦࣐ࢦಶయ⩌ࡢ㑇ఏⓗᵓ 㐀㸧ࡣ㸪ࡑࢀࡒࢀಶูࡢ⤖ᯝࢆ♧ࡋ࡞ࡀࡽ㸪➨ 3 ❶㸦࣐ࢦᾋୖ⛶㨶ࡢὶ⛬ศ ᕸ㸧➨ 4 ❶㸦࣐ࢦᾋୖ⛶㨶ࡢ⏕ᜥሙᡤ⏝㸧࡛ぢࡽࢀࡓୗὶഃỈᇦ࠾ࡅ ࡿᾋୖ⛶㨶ࡢศᕸࡢไ㝈せᅉࡋ࡚㸪ࡑࢀࡒࢀேⅭᨵኚࡢᙳ㡪㸦➨ 2 ❶㸧㣴 Ṫ㨶ࡢᨺὶࡢᙳ㡪㸦➨ 5 ❶㸧ࢆ♧၀ࡋ࡚࠸ࡿ㸬ྛ❶ࡢ⤖ᯝࡽ㸪ᮏㄪᰝỈᇦࡢ ࣐ࢦಶయ⩌ࡢ⌧≧ࡋ࡚௨ୗࡢ⤖ㄽࡀᚓࡽࢀࡓ㸹1㸧⛣⾜ᖏࢆྵࡴୗὶഃỈᇦ ࡣ㸪ேⅭᨵኚࡢᝏᙳ㡪ࢆཷࡅ࡚࠸ࡿྍ⬟ᛶࡀ࠶ࡿ㸬2㸧࣐ࢦ⏕ᜥᇦࡢ 7 ௨ୖ ࡣ⛣⾜ᖏࢆྵࡴୗὶഃỈᇦྵࡲࢀࡿ㸬3㸧࣐ࢦࡢ⮬↛⏕⏘ࡣ⛣⾜ᖏࡼࡾୖ ὶ࡛Ⰻዲ㸪ࡑࡢୗὶ࡛ࡣపㄪ࡛࠶ࡿ㸬4㸧ᅾ᮶࣐ࢦಶయ⩌ࡣ⚄ཎᕝࡢ୍ྜ┠ ࡼࡾୖὶἼᮌྜᕝࡢ࣓ࣥࣀࢶࣛ㇂㸦࣓࣡㸧⏕ᜥࡋ࡚࠸ࡿ㸬5㸧ࡲࢇࡾࡻ࠺ ㇂࡛ࡣ㸪ࣄ࣮ࣂࢳࡢࡢୗὶഃࡣࡰ㣴Ṫ㨶ࡰ⨨ࡁࢃࡗ࡚࠸ࡿࡀ㸪ࡢ ୖὶഃࡣᅾ᮶ಶయ⩌ࡀ⏕ᜥࡋ࡚࠸ࡿྍ⬟ᛶࡀ࠶ࡿ㸬6㸧⛣⾜ᖏࢆྵࡴୗὶഃỈ ᇦࡣ㸪㠀ᅾ᮶ಶయࡀ⏕ᜥࡋ࡚࠸ࡿྍ⬟ᛶࡀ࠶ࡿ㸬ࡇࢀࡽࡢᐇែᇶ࡙ࡁ㸪ಖ 㸦ࢰ࣮ࢽࣥࢢ㸧᪉ἲࡋ࡚௨ୗࡀᑟฟࡉࢀࡓ㸹1㸧⚄ཎᕝࡢ୍ྜ┠ࡽୖὶ ᇦ࠾࠸࡚⁺ᴗ㸪㐟⁺㸪ᨺὶࢆ⚗Ṇ㸬2㸧Ἴᮌྜᕝࡢ⌧⾜⚗⁺༊ࡢୖ㝈ࢆ᧔ᗫ ࡋ㸪ⓑỈᶫࡽୖὶᇦ࠾࠸࡚⁺ᴗ㸪㐟⁺㸪ᨺὶࢆ⚗Ṇ㸬3㸧୧⚗⁺༊ࡼࡾୗ ὶഃ࡛ࡣ⁺ᴗ㐟⁺࠾ࡼࡧᨺὶࢆྍ⬟ࡍࡿ㸬4㸧⛣⾜ᖏ࡛ࡣ㸪ᾋୖ⛶㨶ࡢ⏕ᜥ ሙᡤᩚഛࡸᨺὶࡢぢ┤ࡋࡼࡾ㸪⮬↛⏕⏘ࡢಁ㐍ࢆヨࡳࡿᚲせࡀ࠶ࡿ㸬 ࡇࢀࡽࡢࢰ࣮ࢽࣥࢢࡣ㸪ᆅඖఫẸ㸪㔝ᕝ⁺ᴗ༠ྠ⤌ྜ㸪㐟⁺⪅㸪ᆅඖ⾜ ᨻ㸦➉⏣ᕷ㸧➼࡛⤌⧊ࡋࡓጤဨཷࡅධࢀࡽࢀࡓ㸬2009 ᖺࡲ࡛⁺ᴗ㛵㐃つ ๎ࡀᨵṇࡉࢀ㸪⌧ᅾ㸪ᮏ◊✲⤖ᯝᇶ࡙ࡃ࣐ࢦࡢಖࡀᐇࡉࢀ࡚࠸ࡿ㸬 105 ㅰ ㎡ ᗈᓥᏛᏛ㝔⏕≀ᅪ⛉Ꮫ◊✲⛉ࡢ㛗⃝ஓᩍᤵࡣ㸪ᮏㄽᩥከࡃࡢ㐺ษ࡞ ࡞ࡈຓゝࢆୗࡉࡾᚰࡽឤㅰࡢពࢆ⾲ࡍࡿ㸬ࡲࡓ㸪ᮏㄽᩥ᭷┈࡞ࡈຓゝࢆୗ ࡉࡗࡓྠ◊✲⛉ࡢἙྜᖾ୍㑻ᩍᤵ㸪 ᆏ㝧୍ᩍᤵ㸪 ᾏ㔝ᚭஓᩍᤵឤㅰࡍࡿ㸬 ᮏ◊✲ࡢከࡃࡢ㔝እㄪᰝࡣ㸪ᙜࡢศ┴㎰ᯘỈ⏘◊✲ࢭࣥࢱ࣮Ỉ⏘ヨ㦂ሙ ෆỈ㠃◊✲ᡤ࠾ࡽࢀࡓᬒᖹ┿᫂Ặ㸪␁ᆅஂẶ㸪⚟⏣♸୍Ặࡢࡈ༠ຊࡀ࡞ࡅ ࢀࡤ㐙⾜ࡍࡿࡇࡣ࡛ࡁ࡞ࡗࡓ㸬ࡲࡓ㸪ᙜࡢෆỈ㠃◊✲ᡤࡢᒾᮏ㑳⏕Ặ ⊷Ώ ᐇẶ㸪ᙜࡢศ┴Ỉ⏘⯆ㄢࡢ୕ᾆៅ୍Ặࡣ㸪㔝እㄪᰝࢆᡭఏࡗ࡚ୗࡉ ࡗࡓ㸬ඖศᏛྡᩍᤵࡢᨾྜྷ⏣ṇ㞝༤ኈࡣ㸪⤊ጞ᠓ษ࡞ࡈᣦᑟບࡲࡋ ࢆ࠸ࡓࡔ࠸ࡓ㸬ᙜᕞᏛࡢ㏆⸨༟ဢ༤ኈࡣ㸪◊✲᪉㔪㛵ࡍࡿⓗ☜࡞ࡈᣦ ᑟࡶ㔝እㄪᰝࢆᡭఏࡗ࡚ୗࡉࡗࡓ㸬ᕞᏛỈ⏘ᐇ㦂ᡤࡢ㨣ᚨ㞝༤ኈ ࡣࢹ࣮ࢱゎᯒ᪉ἲࢆࡈᣦᑟ࠸ࡓࡔ࠸ࡓ㸬୕㔜ᏛࡢἙᮧຌ୍༤ኈ㸪ᰣᮌ┴Ỉ ⏘ヨ㦂ሙࡢஂಖ⏣ோᚿ༤ኈ㸪㈡┴Ỉ⏘ヨ㦂ሙࡢட⏥Ṋᚿ༤ኈࡣ㸪㑇ఏᏊゎᯒ ᡭἲ㛵ࡋ࡚ࡈᩍᤵୗࡉࡗࡓ㸬⊂❧⾜ᨻἲேỈ⏘⥲ྜ◊✲ࢭࣥࢱ࣮㣴Ṫ◊✲ᡤ ࡢ୰ᮧᬛᖾ༤ኈࡣ㸪⌧ᆅ࡛ࡢࢰ࣮ࢽࣥࢢᑟධࡘ࠸࡚ࡈᣦᑟ࠸ࡓࡔ࠸ࡓ㸬 㔝ᕝ⁺ᴗ༠ྠ⤌ྜࡢ㔝⬶⏕Ặ㸪㜿༡ಇᘯẶ㸪㤳⸨ㄔ୍Ặ㸪 ࠕ㔮ලࢭࣥࢱ࣮➉⏣ࠖ ௦⾲ࡢ㧗 ₩Ặ㸪㐟⁺ᅋయࠕ33Clubࠖࡢྜྷ㔝ဴஓẶ㸪࠾ࡼࡧ୕⏣ ᪼Ặࡣ㸪 ⚗⁺༊タ⨨㛵ࡍࡿࡈពぢࢆୗࡉࡗࡓ㸬㔮ලᗑࠕWalkerࠖ௦⾲ࡢᏳ㒊ಇᘯẶ㸪 㣴㨶ሙࠕỈⱌࠖ௦⾲ࡢ㊊❧ᚭಙẶࡣ㸪ㄪᰝᮇ㛫୰ࡢ࣐ࢦ⛶㨶ࡢᨺὶ㛵 ࡋ࡚౽ᐅࢆᅗࡗ࡚࠸ࡓࡔ࠸ࡓ㸬㔝ᕝ⁺ᴗ༠ྠ⤌ྜࡢబ⏝ ኖඖ௦⾲⌮⤌ྜ㛗 ࡣ㸪⁺ᴗᶒ⾜つ๎㐟⁺つ๎ࡢᨵṇⱥ᩿ࢆୗࡉࢀࡓ㸬ᙜศ┴⁺ᴗ⟶⌮ ㄢࡢఀᮾᜤ㑻Ặࡣ㸪⁺ᴗㄪᩚつ๎ࡢᨵṇ࡛ኚ࠾ୡヰ࡞ࡗࡓ㸬ఀᮾẶࡢ ࡈᨭࡀ࡞ࡅࢀࡤ㸪Ἴᮌྜᕝࡢ࣓࣡⚗⁺༊ࡢୖ㝈᧔ᗫࡣᐇ⌧ࡋ࡞ࡗࡓ㸬Gࢢ࣮ࣟࣈ௦⾲ࡢỤ୍ཱྀ⏕Ặࡣ㸪ㄪᰝᮇ㛫୰ࡢࢻࣛࢫ࣮ࢶࡢ⿵ಟࡘ࠸࡚ ኚ࠾ୡヰ࡞ࡗࡓ㸬 ศ┴㎰ᯘỈ⏘◊✲ᣦᑟࢭࣥࢱ࣮Ỉ⏘◊✲㒊ࡢ⚟⏣ ✨༤ኈ㸪 ྜྷᒸ᐀భẶ㸪㧗㔝㡰ᏊẶࡣ㸪➹⪅ࡢ༤ኈྕྲྀᚓࢆࡈᨭୗࡉࡗࡓ㸬ࡇࢀࡽࡢ᪉ࠎ ᚰࡼࡾ࠾♩⏦ࡋୖࡆࡿ㸬 106 ᘬ⏝ᩥ⊩ Aarts, B. G. W. and P. H. Nienhuis. 2003. Fish zonations and guilds as the basis for assessment of ecological integrity of large rivers. Hydrobiologia, 500: 157–178. Barton, K. 2015. Package ‘MuMIn’: multi-model inference. R package version 1.13. 4. Bisson, P. A., J. L. Nielson, R. A. Palmason and L. E. Grove. 1982. A system of naming habitat types in small streams, with examples of habitat utilization by salmonids during low streamflow. Pages 62–73 in N. B. Armantrout, ed. Acquisition and utilization of aquatic habitat inventory information. Proceedings of a symposium, Portland, OR, USA. Bisson, P. A., J. M. Buffington and D. R. Montgomery. 2006. Valley segments, stream reaches, and channel units. Pages 23–50 in F. R. Hauer and G. A. Lamberti, eds. Methods in stream ecology (2nd edition). Academic Press, London. Bolker, B. M., M. E. Brooks, C. J. Clark, S. W. Geange, J. R. Poulsen, M. H. H. Stevens and J. S. S. White. 2009. Generalized linear mixed models: a practical guide for ecology and evolution. Trends Ecol. Evol., 24: 127–135. Bolker, B., H. Skaug, A. Magnusson and A. Nielsen. 2012. Getting started with the glmmADMB package. Available at glmmadmb. r-forge. r-project. org/glmmADMB. pdf. Bozek, M. A. and F. J. Rahel. 1991. Assessing habitat requirements of young Colorado River cutthroat trout by use of macrohabitat and microhabitat analyses. Trans. Am. Fish. Soc., 120: 571–581. Burnham, K. P. and D. R. Anderson. 2002. Model selection and multimodel inference: a practical information-theoretic approach, second ed. Springer, New York. xxvi+488 pp. Clement, M., D. Posada and K. A. Crandall. 2000. TCS: a computer program to estimate gene genealogies. Mol. Ecol., 9: 1657–1659. Crandall, K. A., O. R. Bininda-Emonds, G. M. Mace and R. K. Wayne. 2000. 107 Considering evolutionary processes in conservation biology. Trends Ecol. Evol., 15: 290–295. Einum, S., A. G. Finstad, G. Robertsen, K. H. Nislow, S. McKelvey and J. D. Armstrong. 2012. Natal movement in juvenile Atlantic salmon: a body size-dependent strategy? Popul. Ecol., 54: 285–294. Einum, S., G. Robertsen, K. H. Nislow, S. McKelvey and J. D. Armstrong. 2011. The spatial scale of density-dependent growth and implications for dispersal from nests in juvenile Atlantic salmon. Oecologia, 165: 959–969. 㐲⸨㎮࣭ᆤ₶୍࣭ᒾ⏣ᬛஓ㸬2006㸬Ἑᕝᕤస≀ࡀ࣡ࢼ࣐ࢦࡢಶయ⩌ Ꮡ⥆ཬࡰࡍᙳ㡪㸬ಖ⏕ែᏛ◊✲㸪11: 4–12㸬 Excoffier, L. and H. E. Lischer. 2010. Arlequin suite ver 3.5: A new series of programs to perform population genetics analyses under Linux and Windows. Mol. Ecol. Resour., 10: 564–567. Excoffier, L., P. E. Smouse and M. Quattro. 1992. Analysis of molecular variance inferred from metric distances among DNA haplotypes: application to human mitochondrial DNA restriction data. Genetics, 131: 479–491. Fagan, W. F., R. S. Cantrell and C. Cosner. 1999. How habitat edges change species interactions. Am. Nat., 153: 165–182. Fausch, K. D. 1984. Profitable stream positions for salmonids: relating specific growth rate to net energy gain. Can. J. Zool., 62: 441–451. Fausch, K. D., C. L. Hawkes and M. G. Parsons. 1988. Models that predict standing crop of stream fish from habitat variables: 1950–85. Gen. Tech. Rep. PNW-GTR-213. U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, Portland, OR, USA. 52 pp. Foldvik, A., A. G. Finstad and S. Einum. 2010. Relating juvenile spatial distribution to breeding patterns in anadromous salmonid populations. J. Anim. Ecol., 79: 501–509. Fraser, D. J., L. K. Weir, L. Bernatchez, M. M. Hansen and E. B. Taylor. 2011. Extent and scale of local adaptation in salmonid fishes: review and meta-analysis. Heredity, 108 106: 404–420. Frissell, C. A., W. J. Liss, C. E. Warren and M. D. Hurley. 1986. A hierarchical framework for stream habitat classification: viewing streams in a watershed context. Environ. Manage., 10: 199–214. Garcia de Leaniz, C., I. A. Fleming, S. Einum, E. Verspoor, W. C. Jordan, S. Consuegra, N. Aubin-Horth, D. Lajus, B. H. Letcher, A. F. Youngson, J. H. Webb, L. A. Vøllestad, B. Villanueva, A. Ferguson and T. P. Quinn. 2007. A critical review of adaptive genetic variation in Atlantic salmon: implications for conservation. Biol. Rev., 82: 173–211. Geist, D. R. and D. D. Dauble. 1998. Redd site selection and spawning habitat use by fall chinook salmon: the importance of geomorphic features in large rivers. Environ. Manage., 22: 655–669. Girard I. L., J. W. A. Grant and S. Ó. Steingrímsson. 2004. Foraging, growth, and loss rate of young-of-the year Atlantic salmon (Salmo salar) in relation to habitat use in Catamaran Brook, New Brunswick. Can. J. Fish. Aquat. Sci., 61: 2339–2349. Hall, H. J. and L. W. Nawrocki. 1995. A rapid method for detecting mitochondrial DNA variations in the brown trout, Salmo trutta. J. Fish Biol. 46: 360–364. ᮏⲮ㕲ኵ㸬1977㸬࣐ࢦࡢቑ㣴Ṫ㛵ࡍࡿᇶ♏ⓗ◊✲㸬ᒱ㜧┴Ỉ⏘ヨ㦂ሙ◊✲ ሗ࿌㸪22: 1–103. ර⸨๎⾜࣭㛵 Ὀኵ࣭ሯᮏᕭ࣭▮┿▱Ꮚ࣭ஂಖஂ┤㸬1992㸬ຍᕝ࠾ ࡅࡿᢳỈ᳜≀⩌ⴠࡢ⏕ែⓗ౯್㸬᪂₲┴ෆỈ㠃Ỉ⏘ヨ㦂ሙㄪᰝ◊✲ሗ࿌㸪18: 1–20. 㣤⏣ 㐶࣭୰ᮧᬛᖾ㸬2009㸬⁺༠ࡢᙺ㔮ࡾேࡢ⩏ົ㸬୰ᮧᬛᖾ࣭㣤⏣ 㐶 㸦⦅㸧 㸪pp.35–40. Ᏺࡿ࣭ቑࡸࡍὶ㨶㸬㎰ᒣ⁺ᮧᩥ༠㸪ᮾி㸬 ୖᖿ⏕㸬2013㸬㨶㢮㸬ᕝ㑣㒊ᾈဢ࣭Ỉ㔝ಙᙪ㸦┘ಟ㸧 㸪୰ᮧኴኈ㸦⦅㸧 㸪pp. 123– 144. Ἑᕝ⏕ែᏛ㸬ㅮㄯ♫㸪ᮾி㸬 Inoue, M and M. Nunokawa. 2002. Effects of longitudinal variations in stream habitat structure on fish abundance: an analysis based on subunit-scale habitat classification. 109 Freshwater Biol., 47: 1594–1607. Inoue, M. and S. Nakano. 1999. Habitat structure along channel-unit sequences for juvenile salmon: a subunit-based analysis of in-stream landscape. Freshwater Biol., 42: 597–608. Inoue, M., S. Nakano and F. Nakamura. 1997. Juvenile masu salmon (Oncorhynchus masou) abundance and stream habitat relationships in northern Japan. Can. J. Fish. Aquat. Sci., 54: 1331–1341. ᯈ㝯ᙪ㸬1976㸬ዉⰋ┴㧗ぢᕝࡢࣈࣛࣁࣖᒓ㸦Phoxinus㸧㨶㢮ࡢ 2 ᆺ—ࡑࡢ ᙧែⓗ⏕ែⓗ≉ᚩࡘ࠸࡚㸬㟼ᒸዪᏊᏛ◊✲⣖せ㸪10: 201–220. ᯈ㝯ᙪ㸬1979㸬㟼ᒸ┴℩ᡞᕝỈ⣔࠾ࡅࡿࣈࣛࣁࣖᒓ㸦Phoxinus㸧㨶㢮ࡢ 2 ᆺ—I㸬ὶࢀἢࡗࡓศᕸࡘ࠸࡚㸬㟼ᒸዪᏊᏛ◊✲⣖せ㸪13: 153–175. Jackson, D. A., P. R. Peres-Neto and J. D. Olden. 2001. What controls who is where in freshwater fish communities—the roles of biotic, abiotic, and spatial factors. Can. J. Fish. Aquat. Sci., 58: 157–170. ᬒᖹ┿࣭᫂ᚨගಇ㸬2003㸬ᮍ⏕ᜥୖὶᇦࡢ࣓࣡ࡢ⛣᳜ᨺὶຠᯝ㸬ศᾏ Ỉ◊ㄪ◊ሗ㸪4: 71–76. Kanaiwa, M. and Y. Harada. 2010. Theoretical study of fitness reduction in stream-dwelling fish by the fixation of slightly harmful mutations under habitat fragmentation. J. Agric. Sci., Tokyo Univ. Agric., 55: 10–18. ⚄⏣ၿఙ㸬2012㸬EZR ࡛ࡸࡉࡋࡃᏛࡪ⤫ィᏛ㹼EBM ࡢᐇ㊶ࡽ⮫ᗋ◊✲ࡲ࡛ 㹼㸬୰እ་Ꮫ♫㸪ᮾி㸪365 pp. ⎔ቃᗇ㸬1987㸬ศ┴㸦4㸧㔝ᕝ㸬pp. 43–64. ➨ 3 ᅇ⮬↛⎔ቃಖᇶ♏ㄪᰝἙ ᕝㄪᰝሗ࿌᭩㸪ᕞ࣭Ἀ⦖∧㸬ⶶ┬༳ๅᒁ㸪ᮾி㸬 ྍඣ⸨ྜྷ㸬1978㸬ὶ᳇᪻ࡢ⏕ែ㸬pp. 3–91. ྍඣ⸨ྜྷ㞟㸪୍ᕳ㸪ᬑཬ∧㸬 ᛮ⣴♫㸪ᮾி㸬 Kano, Y., T. Kondou and Y. Shimizu. 2010. Present status and conservation of the markless forms of stream-resident masu salmon Oncorhynchus masou (the so-called ‘iwame’) in Japanese mountain streams. Ichthyol. Res., 57: 78–84. 110 Kano, Y., Y. Shimizu and K. Kondou. 2006. Sympatric, simultaneous, and random mating between markless trout (iwame; Oncorhynchus iwame) and red-spotted masu salmon (amago; Oncorhynchus masou ishikawae). Zool. Sci., 23: 71–77. ⢑㇂ⱥ୍㸬2012㸬㞳ᩓⓗࢹ࣮ࢱ㐣ศᩓ㸬㔠 ᫂ဴ㸦⦅㸧 㸪pp. 81–102㸬R ࡛Ꮫ ࡪࢹ࣮ࢱࢧ࢚ࣥࢫ 10 ୍⯡⥺ᙧࣔࢹࣝ㸬ඹ❧ฟ∧㸪ᮾி㸬 ∦㔝 ಟ࣭୰ᮧᬛᖾ࣭ᒣᮏ⚈୍㑻࣭㜿㒊ಙ୍㑻㸬2004㸬㛗㔝┴ᾆ㔝ᕝ࠾ࡅࡿ 㨶㢮ࡢ✀⤌ᡂ㣗≀㛵ಀ㸬᪥ᮏỈ⏘Ꮫㄅ㸪70: 902–909. ຍ⸨᠇ྖ㸬1990㸬࣐࣓࣭࣐ࣖࢦ㸪ࡑࡢ⏕ែ㔮ࡾ㸬ࡘࡾே♫㸪ᮾி㸬195 pp㸬 Kawamura, K., M. Kubota, M. Furukawa and Y. Harada. 2007. The genetic structure of endangered indigenous populations of the amago salmon, Oncorhynchus masou ishikawae, in Japan. Conserv. Genet., 8: 1163–1176. ⴛሙ♸୍࣭ᒸ⏣ᬛᖾ࣭బࠎᮌⰋᾈ࣭ᑠᕝ㭯ⶶ࣭㇂ၨᖾ࣭㧗ᮌⱱ▱࣭ᯘ ᑦ㸬 2003b㸬⌧ᆅほ ᇶ࡙ࡃ◁㕲ᕝ࠾ࡅࡿ㨶㢮⏕ᜥሙᡤࡢᢕᥱண 㸬Ἑᕝᢏ ⾡ㄽᩥ㞟㸪9: 433–438. ⴛሙ♸୍࣭༓ⴥṊ⏕࣭ຊᒣ ᇶ࣭Ἑཱྀὒ୍࣭ᑿ⃝༟ᛮ㸬2002㸬ࣁࣅࢱࢵࢺࢭ ࢫ࣓ࣥࢺ࠾ࡅࡿἙ㐨ෆᚤᆅᙧࡢᢕᥱᡭἲ㛵ࡍࡿ◊✲㸬Ἑᕝᢏ⾡ㄽᩥ㞟㸪 8: 191–196. ⴛሙ♸୍࣭༓ⴥṊ⏕࣭ຊᒣ ᇶ࣭ᑿ⃝༟ᛮ㸬2003a㸬୰ᑠἙᕝ୰ὶᇦ࠾ࡅࡿ㨶 㢮⏕ᜥሙᡤࡢศᕸᵓ㐀㸬Ἑᕝᢏ⾡ㄽᩥ㞟㸪9: 421–426. ⴛሙ♸୍㸬2013㸬Ἑᕝᆅᙧࡢ≉ᚩࡑࡢศ㢮㸬ᕝ㑣㒊ᾈဢ࣭Ỉ㔝ಙᙪ㸦┘ಟ㸧 㸪 ୰ᮧኴኈ㸦⦅㸧 㸪pp. 13–33. Ἑᕝ⏕ែᏛ㸬ㅮㄯ♫㸪ᮾி㸬 Kikko T., Y. Harada, D. Takeuchi and Y. Kai. 2008. Interpopulation variation in egg size of fluvial white-spotted charr Salvelinus leucomaenis. Fish. Sci., 74: 935–937. Kikko T., Y. Kai, M. Kuwahara and K. Nakayama. 2008. Genetic diversity and population structure of white-spotted charr, Salvelinus leucomaenis, in the Lake Biwa water system inferred from AFLP analysis. Ichthyol. Res., 55: 141–147. ᮌᮏᆂ㍜㸬2008㸬ኳ↛⏕⏘ຊࡀపࡃ✀ⱑᨺὶࡀྍḞ࡞ὶᇦ࠾ࡅࡿࢰ࣮ ࢽࣥࢢᑟධ㝿ࡋ࡚ࡢㄢ㢟ᢕᥱ㸬⊂❧⾜ᨻἲேỈ⏘⥲ྜ◊✲ࢭࣥࢱ࣮୰ኸỈ 111 ⏘◊✲ᡤ㸦⦅㸧 㸪pp. 69–91㸬ὶᇦ⟶⌮యไᵓ⠏ᴗሗ࿌᭩㸬Ỉ⏘ᗇ㸪ᮾி㸬 ᮌᮏᆂ㍜࣭ᬒᖹ┿࣭᫂␁ᆅஂ࣭⚟⏣♸୍࣭㛗⃝ஓ㸬2013㸬ᕞࡢ୍ὶ ࠾ࡅࡿ࣐ࢦᾋୖ⛶㨶ࡢὶ⛬ศᕸ㸬㨶㢮Ꮫ㞧ㄅ㸪60: 15–26. ᮌᮏᆂ㍜࣭ᬒᖹ┿࣭᫂␁ᆅஂ࣭㛗⃝ஓ㸬2015㸬ᕞᮾ㒊ࡢἙᕝୖὶᇦ ࠾ࡅࡿὶ⛬ἢࡗࡓ㨶㢮⩌㞟ࡢኚ㸬㨶㢮Ꮫ㞧ㄅ㸪62: 1–12㸬 ᮌᮏᆂ㍜࣭ᬒᖹ┿࣭᫂␁ᆅஂ࣭㛗⃝ஓ㸬2015㸬ᕞᮾ㒊Ἑᕝࡢ࣐ࢦ⏕ ᜥᇦୗὶഃ࠾ࡅࡿᾋୖ⛶㨶ࡢ⏕ᜥሙᡤ⏝㸬㨶㢮Ꮫ㞧ㄅ㸪62㸦༳ๅ୰㸧 㸬 Kimoto, K., T. Mekata, H. Takahashi and K. Nagasawa. 2015. Genetic structure of the amago and iwame forms of the red-spotted masu salmon (Oncorhynchus masou ishikawae) in the upper Ono River, northeastern Kyushu, southern Japan. Aquacult. Sci., 63 (in press). ᮌᮧΎᮁ㸬1959㸬♽ẕ࣭ഴᒣ⩌ࡢ࢚ࣀࣁ㸬ຍ⸨ᩘຌ࣭❧▼ᩄ㞝㸦⦅㸧 㸪pp. 109– 119. ♽ẕᔂᒣ⩌㸬ࡋࢇࡘࡃࡋᒣᓅ㸪⚟ᒸ㸬 ᮌᮧΎᮁ㸬1972㸬࣐࣓ࣖࡢ⏘༸⩦ᛶࡘ࠸࡚㸬㨶㢮Ꮫ㞧ㄅ㸪19: 111–119. ᮌᮧΎᮁ㸬2004㸬ὶ㨶ࡢ㐺ṇ࡞⏝ቑṪࡢࡓࡵ㸬ᮌᮧΎᮁ㸦┘ಟ㸧㸪pp. 243– 252. ࣡ࢼ㸪࣐࣓ࣖ㸪࣐ࢦࡢቑṪ⟶⌮㸬ᅜෆỈ㠃⁺ᴗ༠ྠ⤌ྜ㐃ྜ㸪 ᮾி㸬 Kimura, S. 1989. The yamame, land-locked masu salmon of Kyushu Island, Japan. Physiol. Ecol. Japan, Spec., 1: 77–92. Kimura, S. and M. Nakamura. 1961. A new salmonid fish, Oncorhynchus iwame sp. nov., obtained from Kyushu, Japan. Bull. Biogeogr. Soc. Japan, 22: 69–74. 㔠 ᫂ဴ㸬2007㸬R ࡼࡿࢹ࣮ࢱࢧ࢚ࣥࢫ—ࢹ࣮ࢱゎᯒࡢᇶ♏ࡽ᭱᪂ᡭἲ ࡲ࡛㸬᳃ฟ∧㸪ᮾி㸬307 pp. ᕝ࠼ࡳ࣭ᫍ㔝ኵ࣭ᒸᓮⓏᚿኵ࣭ᕝᛅ⏕㸬2004㸬ศ┴ศᕝỈ⣔ࡽ࠼ ࡽࢀࡓࢩ࣐ࢻࢪࣙ࢘ࡑࡢ⏕≀ᆅ⌮ⓗ㉳※㸬㨶㢮Ꮫ㞧ㄅ㸪51: 117–122. ᕝ࠼ࡳ࣭୰ᓥ ῟࣭ᫍ㔝ኵ࣭ᕝᛅ⏕㸬2009㸬ᕞᮾ㒊࠾ࡅࡿࢩ࣐ࢻ ࢪࣙ࢘ᒓ㨶㢮ࡢศᕸࣃࢱ࣮ࣥࡑࡢᡂ❧㐣⛬㛵ࡍࡿ⪃ᐹ㸬㨶㢮Ꮫ㞧ㄅ㸪56: 7–19. 112 ㏆⸨༟ဢ࣭➉ୗ┤ᙪ㸬2005㸬࣓࣡ࡢ⏕ែʊ⛣ື㸪ᡂ㛗࠾ࡼࡧ⏘༸㸬∦㔝 ಟ࣭ ᳃ ㄔ୍㸦⦅㸧㸪pp. 232–239. ᕼᑡῐỈ㨶ࡢ⌧ᅾᮍ᮶ʊ✚ᴟⓗಖࡢࢩࢼࣜ ࢜ʊ㸬ಙᒣ♫㸪ᮾி㸬 Kramer, D. L., R. W. Rangeley and L. J. Chapman. 1997. Habitat selection: patterns of spatial distribution from behavioural decisions. Pages 37–80 in J.-G. J. Godin, ed. Behavioural ecology of teleost fishes. Oxford University Press, Oxford. ஂಖ㐩㑻㸬1980㸬ᾏ㐨ࡢࢧࢡ࣐ࣛࢫࡢ⏕άྐ㛵ࡍࡿ◊✲㸬ᾏ㐨ࡉࡅ࣭ࡲ ࡍࡩሙ◊✲ሗ࿌㸪34: 1–95. ஂಖ⏣ோᚿ࣭୰ᮧᬛᖾ࣭ᒣ㝯࣭Ώ㑔⢭୍㸬2001㸬ᑠᨭὶ࠾ࡅࡿ࣡ࢼ㸪ࣖ ࣐࣓ᙜṓ㨶ࡢ⏕ᜥᩘ㸪⛣ືศᩓ࠾ࡼࡧᡂ㛗㸬᪥ᮏỈ⏘Ꮫㄅ㸪67: 703–709. Kubota, H., T. Doi, S. Yamamoto and S. Watanabe. 2007. Genetic identification of native populations of fluvial white-spotted charr Salvelinus leucomaenis in the upper Tone River drainage. Fish. Sci., 73: 270–284. Lasne, E., B. Bergerot, S. Lek and P. Laffaille. 2007. Fish zonation and indicator species for the evaluation of the ecological status of rivers: example of the Loire basin (France). River Res. Appl., 23: 1–14. ๓ᕝගྖ㸬2000㸬ὶ㨶ࡢ⏕ែ◁㜵ᕤࡢᙳ㡪㸬ኴ⏣⊛ᙪ࣭㧗ᶫ๛୍㑻㸦⦅㸧㸪 pp. 89–105. ὶ⏕ែ◁㜵Ꮫ㸬ᮾிᏛฟ∧㸪ᮾி㸬 ๓ᕝගྖ࣭୰㔝 ⦾㸬1994㸬㐳Ἑᅇ㐟ࡽἙᕝ⏕ά㸬ᚋ⸨ ࣭ሯᮏᕭ࣭ ๓ᕝගྖ㸦⦅㸧 㸪pp. 206–220. ᕝᾏࢆᅇ㐟ࡍࡿῐỈ㨶ʊ⏕άྐ㐍ʊ㸬ᮾ ᾏᏛฟ∧㸪ᮾி㸬 ┿ᒣ ⣫㸬1992㸬ࢧࢡ࣐ࣛࢫ Oncorhynchus masou (Brevoort) ࡢῐỈᇦࡢ⏕ά࠾ ࡼࡧ㈨※ᇵ㣴㛵ࡍࡿ◊✲㸬ᾏ㐨ࡉࡅ࣭ࡲࡍࡩሙ◊✲ሗ࿌㸪46: 1–156. ┿ᒣ ⣫㸬1999㸬༓ṓᕝ࠾ࡅࡿࢧࢡ࣐ࣛࢫᗂ㨶࠾ࡼࡧࣈࣛ࢘ࣥࢺࣛ࢘ࢺࡼ ࡿᾋୖᮇࢧࢡ࣐ࣛࢫ⛶㨶ࡢᤕ㣗㸬ࡉࡅ࣭ࡲࡍ㈨※⟶⌮ࢭࣥࢱ࣮◊✲ሗ࿌㸪2: 21–27. McRae, C. J., K. D. Warren and J. Shrimpton. 2012. Spawning site selection in interior Fraser River coho salmon Oncorhynchus kisutch: an imperiled population of 113 anadromous salmon from a snow-dominated watershed. Endang. Species Res., 16: 249–260. Mercado-Silva, N., J. Lyons, E. Díaz-Pardo, S. Navarrete and A. Gutiérrez-Hernández. 2012. Environmental factors associated with fish assemblage patterns in a high gradient river of the Gulf of Mexico slope. Rev. Mex. Biodiv., 83: 117–128. Miller, L. M., T. Close and A. R. Kapuscinski. 2004. Lower fitness of hatchery and hybrid rainbow trout compared to naturalized populations in Lake Superior tributaries. Mol. Ecol., 13: 3379–3388. Mizoiri, S., N. Takeshita, S. Kimura and O. Tabeta. 1997. Geographical distributions of two bagrid catfishes in Kyushu, Japan. Suisanzoshoku, 45: 497–503. Ỉ㔝ಙᙪ࣭ᚚໃஂྑ⾨㛛㸬2000㸬Ἑᕝࡢ⏕ែᏛ㸪⿵ゞ∧㸬⠏ᆅ᭩㤋㸪ᮾி㸬247 pp. Moore, K. M. S. and S. V. Gregory. 1988. Response of young-of-the-year cutthroat trout to manipulation of habitat structure in a small stream. Trans. Am. Fish. Soc., 117: 162–170. ᳃⏣ኴ㑻࣭ᒣᮏ⚈୍㑻㸬2004㸬ࢲ࣒ᵓ⠏ࡼࡿἙᕝศ᩿ࡀࡶࡓࡽࡍࡶࡢ— ᕝࡣ᳃ᾏࢆࡘ࡞ࡄ㐨—㸬๓ᕝගྖ㸦⦅㸧㸪pp. 281–312. ࢧࢣ࣭࣐ࢫࡢ⏕ែ 㐍㸬ᩥ୍⥲ྜฟ∧㸪ᮾி㸬 Mull, K. E. and M. A. Wilzbach. 2007. Selection of spawning sites by coho salmon in a northern California stream. N. Am. J. Fish. Manage., 27: 1343–1354. Nagata, M. 2002. Ecological studies on the dispersal of newly emerged masu salmon fry, Oncorhynchus masou. Sci. Rep. Hokkaido Hatchery, 56: 1–87. Nagata, M. and S. Yanai. 2002. Changes in habitat use and preference by newly emerged masu salmon fry, Oncorhynchus masou, in Hokkaido streams. Fish. Sci., 68 (Suppl. 1): 45–48. ྡ㉺ ㄔ࣭୰㔝 ⦾࣭ᚨ⏣ᖾ᠇㸬1988㸬ὶᇦ࠾ࡅࡿ࣐ࢦࡢᡂ㛗క࠺⏕ ᜥሙᡤ࠾ࡼࡧ㣗≀⏝ࡢኚ㸬᪥ᮏỈ⏘Ꮫㄅ㸪54: 33–38. Naiman, R. J., H. Décamps, J. Pastor and C. A. Johonston. 1988. The potential 114 importance of boundaries to fluvial ecosystems. J. N. Am. Benthol. Soc., 7: 289– 306. ୰ᆓᚭḟ㸦⦅㸧 㸬2013㸬᪥ᮏ⏘㨶㢮᳨⣴ʊ✀ࡢྠᐃʊ㸦➨ 3 ∧㸧㸬ᮾᾏᏛฟ ∧㸪⛙㔝㸪2428 pp. ୰ᮧಇභ㸬1997㸬ὶἣㄪᰝ㸬⋢ಙ⾜࣭Ỉ㔝ಙᙪ࣭୰ᮧಇභ㸦⦅㸧㸪pp. 244–251. Ἑᕝ⏕ែ⎔ቃᕤᏛ㸬ᮾிᏛฟ∧㸪ᮾி㸬 ୰ᮧኴኈ㸬1989㸬㔝እ⛉Ꮫ࠾ࡅࡿࢫࢣ࣮ࣝㄽ: ✵㛫ၥ㢟ࡢᩚ⌮㸬ᾏ㐨 Ꮫ㎰Ꮫ㒊₇⩦ᯘ◊✲ሗ࿌㸪46: 287–313. ୰ᮧᬛᖾ㸬1999a㸬㨣ᛣᕝୖὶ࠾ࡅࡿ࣡ࢼ㸪࣐࣓ࣖࡢ⏘༸ᗋࡢ❧ᆅ᮲௳ࡢẚ ㍑㸬᪥ᮏỈ⏘Ꮫㄅ㸪65: 427–433. ୰ᮧᬛᖾ㸬1999b㸬ேᕤ⏘༸ሙ࠾ࡅࡿ࣡ࢼࡢ⏘༸⏘╔༸ࡢࡩ㸬᪥ᮏỈ⏘ Ꮫㄅ㸪65: 434–440. ୰ᮧᬛᖾ㸬2001㸬⪺ࡁྲྀࡾㄪᰝࡼࡿ࣡ࢼᅾ᮶ಶయ⩌ࡢ⏕ᜥศᕸ᥎ᐃ㸬◁㜵 Ꮫㄅ㸪53: 3–9. ୰ᮧᬛᖾ㸬2005㸬࣡ࢼࡢ⛶㨶࡛ࡣ㸪ᨺὶࡉࢀࡓᆺ㣴Ṫ㨶ࡀᑠᆺኳ↛㨶ࡢ⏕ ᜥᩘࢆῶࡽࡋ㸪ᡂ㛗ࢆᢚไࡍࡿ㹼ᨺὶ㨶ࡼࡿኳ↛㨶ࡢࠕ⏕ែⓗᨩࠖ㹼㸬 ࢡࢿࢵࢺ㸪(86): 44–47. ୰ᮧᬛᖾ㸬2009㸬ὶ㔮ࡾሙࡢࢰ࣮ࢽࣥࢢ⟶⌮㸬୰ᮧᬛᖾ࣭㣤⏣ 㐶㸦⦅㸧㸪pp. 47–56. Ᏺࡿ࣭ቑࡸࡍὶ㨶㸬㎰ᒣ⁺ᮧᩥ༠㸪ᮾி㸬 ୰ᮧᬛᖾ㸬2011㸬㨣ᛣᕝࡢ㯮㒊ࢲ࣒ୗὶ࠾ࡅࡿἙᗋࡢ㟢┙㸬ᅵᮌᏛㄽᩥ 㞟 B1㸦ỈᕤᏛ㸧 㸪67: 54–59. ୰ᮧᬛᖾ࣭ᚨ⏣ᖾ᠇࣭㧗ᶫ๛୍㑻㸬2009㸬ேᕤ⏘༸Ἑᕝ࠾ࡅࡿ࣡ࢼࡢ⏘༸ ᙜṓ㨶ࡢືែ㸬ᛂ⏝⏕ែᕤᏛ㸪12: 1–12. Nakamura, T. and T. Doi. 2014. Do stocked hatchery-reared juveniles ecologically suppress wild juveniles in Salvelinus leucomaenis? J. Fish Biol., 84: 1289–1299. ୰㔝 ⦾࣭ୖᖿ⏕࣭᱓ཎ⚞▱࣭㇏ᓥ↷㞝࣭ᲄ ඖ࣭⸨ᡞỌᚿ࣭ᮡᒣ ᘯ࣭ ዟᒣ ᝅ࣭➲㈡୍㑻㸬1995㸬ᾏ㐨Ꮫኳሷ࣭୰ᕝᆅ᪉₇⩦ᯘ࠾ࡼࡧ㞄᥋ᆅ ᇦ࠾ࡅࡿῐỈ㨶㢮┦ᒣ࣭◁㜵ࢲ࣒ࡀศᕸཬࡰࡍᙳ㡪㸬ᾏ㐨Ꮫ㎰ 115 Ꮫ㒊₇⩦ᯘ◊✲ሗ࿌㸪52: 95–109. ୰㔝 ⦾࣭⏣ཱྀⱱ⏨࣭ᰘ⏣ຬ࣭ྂᕝဴኵ㸬1993㸬ࢧࢶ࣐࢟ࢫ࣭࣐ࢦ㸬ᕝ㑣 㒊ᾈဢ࣭Ỉ㔝ಙᙪ㸦⦅㸧 㸪pp. 169–179. ᪥ᮏࡢῐỈ㨶㸬ᒣ㇂♫㸪ᮾி㸬 Nakano, S. and M. Nagoshi. 1985. Density regulation and growth of a redspot masu-trout, Oncorhynchus rhodurus, in a mountain stream. Physiol. Ecol. Japan, 22: 1–16. Nakano, S., T. Kachi and M. Nagoshi. 1990. Restricted movement of the fluvial form of red-spotted masu salmon, Oncorhynchus masou rhodurus, in a mountain stream, central Japan. Japan. J. Ichthyol., 37: 158–163. ᰿ṇ㸬1990㸬ศᏊ㐍㑇ఏᏛ㸬ᇵ㢼㤋㸪ᮾி㸬vii+433 pp. Nielsen, E. E., M. M. Hansen and L. A. Bach (2001) Looking for a needle in a haystack: discovery of indigenous Atlantic salmon (Salmo salar L.) in stocked populations. Conserv. Genet., 2: 219–232. ᑿ⏣ᫀ⣖㸬2011㸬⍇⍈†ὶධἙᕝጜᕝ࠾ࡅࡿࣅ࣐࣡ࢫࡢ⏘༸⛶㨶ࡢᾋୖʊ ᬮక࠺ἙᕝỈ ୖ᪼ࡢᙳ㡪ゎᯒʊ㸬ᛂ⏝⏕ែᕤᏛ㸪13: 149–154. ᓥṇ‶㸬1957㸬ᕞࡅࡿ࣐࣓࣐ࣖࢦࡢศᕸ㸬ື≀Ꮫ㞧ㄅ㸪66: 21–24. ᒸ㒊ṇஓ࣭ᑠᯇ❶༤㸬2012㸬㧗▱┴ዉ༙ᕝỈ⣔࠾ࡅࡿᅾ᮶࣐ࢦࡢ㆑ู ಶయ⩌ᵓ㐀ࡢ᥎ᐃ㸬Ỉ⏘ቑṪ㸪60: 89–97. Oksanen, J. 2013. Multivariate analysis of ecological communities in R: vegan tutorial: http://cc.oulu.fi/~jarioksa/opetus/metodi/vegantutor.pdf. Oksanen, J. 2014. Cluster analysis: tutorial with R. University of Oulu, Oulu: http://cc.oulu.fi/~jarioksa/opetus/metodi/sessio3res.pdf. Oksanen, J., F. G. Blanchet, R. Kindt, P. Legendre, P. R. Minchin, R. B. O’Hara, G. L. Simpson, P. Solymos, M. Henry, H. Stevens and H. Wagner. 2013. Package ‘vegan’. Community ecology package Version 2.0-10: http://vegan.r-forge.r-project.org/, http://cran.r-project.org. Palumbi, S., A. Martin, S. Romano, W. O. McMillan, L. Stice and G. Grabowski .1991. The Simple Fool’s Guide to PCR. Version 2. University of Hawaii Zoology 116 Department, Honolulu, Hawaii, 46 pp. Primmer, C. R. 2011. Genetics of local adaptation in salmonid fishes. Heredity, 106: 401–403. R Development Core Team. 2005. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria, URL http://www.R-project.org. Rahel, F. J. and W. A. Hubert. 1991. Fish assemblages and habitat gradients in a Rocky Mountain–Great Plains stream: biotic zonation and additive patterns of community change. Trans. Am. Fish. Soc., 120: 319–332. Raymond, M. and F. Rousset. 1995. An exact test for population differentiation. Evolution, 49: 1280–1283. బᕝᚿᮁ࣭୰ᮧኴኈ㸬2010㸬Ἑᕝᨵಟ㨶㢮ࡢಖ㸬㔝⏕⏕≀ಖㆤᏛ㸦⦅㸧 㸪 pp. 268–279. 㔝⏕ື≀ಖㆤࡢ㸬ᮅ᭩ᗑ㸪ᮾி㸬 Sakai, H., C. Yamamoto and A. Iwata. 1998. Genetic divergence, variation and zoogeography of a freshwater goby, Odontobutius obscura. Ichthyol. Res., 45: 363– 376. 㜰⏣ᘯ㸬1993㸬⌫☻ᕝὶᇦ࠾ࡅࡿ࣐࣓ࣖ⏕ᜥᐦᗘἙᗋᙧែࡢ㛵ಀ㸬 Ỉ⏘ቑṪ㸪41: 27–33. Sakata, K., T. Kondou, N. Takeshita, A. Nakazono and S. Kimura. 2005. Movement of the fluvial form of masu salmon, Oncorhynchus masou masou, in a mountain stream in Kyushu, Japan. Fish. Sci., 71: 333–341. Sato, T., T. Demise, H. Kubota, M. Nagoshi and K. Watanabe. 2010. Hybridization, isolation, and low genetic diversity of Kirikuchi char, the southernmost populations of the genus Salvelinus. Trans. Am. Fish. Soc., 139: 1758–1774. Schlosser, I. J. 1982. Fish community structure and function along two habitat gradients in a headwater stream. Ecol. Monogr., 52: 395–414. ᚿ㈡ྐග࣭ᕝ㔝⏣ᐇኵ㸬1977㸬㔝ᕝỈ⣔ࡢỈ㉁㸬ศᏛᩍ⫱Ꮫ㒊㸦⦅㸧 㸪pp. 62–87. 㔝ᕝ㸸⮬↛࣭♫࣭ᩍ⫱㸬ศᏛᩍ⫱Ꮫ㒊㸪ศ㸬 117 ᓥ㔝Ᏻ㞝࣭ᡞḟᩥኵ㸬1994㸬ศ┴ࡢྡỈʊ⏨ụỈ࣭➉⏣Ỉ⩌࣭ⓑᒣᕝ㸬 ᪥ᮏᆅୗỈᏛ㸦⦅㸧㸪pp. 262–276㸬ྡỈࢆ⛉Ꮫࡍࡿ㸬ᢏሗᇽฟ∧㸪ᮾி㸬 ⓑ▼ⰾ୍࣭㕥ᮌ႐୕㑻࣭⋢⏣㑻㸬1957㸬୕㔜┴㤿㔝ᕝࡢ࣐ࢦ㛵ࡍࡿỈ⏘ ⏕≀Ꮫⓗ◊✲㸬➨ሗ㸬⏘༸⩦ᛶ㛵ࡍࡿ◊✲㸬ῐỈ༊Ỉ⏘◊✲ᡤ◊✲㈨ᩱ㸪 14: 1–17. Steingrímsson, S. Ó. and J. W. A. Grant. 2003. Patterns and correlates of movement and site fidelity in individually tagged young-of-the-year Atlantic salmon (Salmo salar). Can. J. Fish. Aquat. Sci., 60: 193–202. Stockwell, C. A., A. P. Hendry and M. T. Kinnison. 2003. Contemporary evolution meets conservation biology. Trends Ecol. Evol., 18: 94–101. ᮡⱝᆂ୍࣭➉ෆᕭ࣭㕥ᮌ◊୍࣭Ọ⏣ග༤࣭ᐑᮏ┿ே࣭ᕝᮧὒྖ㸬1999㸬ཌ⏣ ᕝ࠾ࡅࡿࢧࢡ࣐ࣛࢫ⏘༸ᗋࡢศᕸᵓ㐀㸬ᾏ㐨❧Ỉ⏘Ꮴሙ◊✲ሗ࿌㸪 53: 11–28. ❧ᕝ ற࣭ᮏⲮ㕲ኵ㸬1976㸬Ἑᕝᨺὶ㸬ᅜ†Ἑᕝ㣴Ṫ◊✲㣴㫟㒊㸦⦅㸧 㸪 pp. 123–137. 㣴㫟ࡢ◊✲㸬⥳᭩ᡣ㸪ᮾி㸬 ⏣ᓥᩄ⾜㸬1965㸬ኳ↛グᛕ≀㹙♽ẕᒣ⣔㝣ᑒ㨶࣓࣡ࠖᣦᐃ⏦ㄳ᭩㸬➉⏣ᕷ㸪 ศ㸬10 pp. 㧗℈⚽ᶞ࣭㣕℩㯞⨾࣭ᯇᑿᩄ⏕࣭Ụ⸨❳࣭▼⏣ ῟㸬2006㸬ศ┴ࡢἙᕝ ⏕ᜥࡍࡿࢩ࣐ࢻࢪࣙ࢘ Cobitis biwae ࣐ࣖࢺࢩ࣐ࢻࢪࣙ࢘ Cobitis matsubarai ࡢᆅ⌮ⓗศᕸ㸬ศᏛᩍ⫱⚟♴⛉Ꮫ㒊◊✲⣖せ㸪28: 155–160. 㧗ᶫ๛୍㑻㸬1988㸬ὶ⎔ቃࡢಖ㛵ࡍࡿ◁㜵Ꮫⓗ◊✲㸬ᾏ㐨Ꮫ㎰Ꮫ㒊 ₇⩦ᯘ◊✲ሗ࿌㸪45: 371–453. 㧗ᶫ๛୍㑻㸬2000㸬ὶ࠾ࡅࡿ◁㜵ᕤ㸬ኴ⏣⊛ᙪ࣭㧗ᶫ๛୍㑻㸦⦅㸧 㸪pp. 41– 50. ὶ⏕ែ◁㜵Ꮫ㸬ᮾிᏛฟ∧㸪ᮾி㸬 㧗ᶫ๛୍㑻࣭ኴ⏣⊛ᙪ㸬2000㸬ὶ⎔ቃࡢ≀⌮ⓗᵓ㐀㸬ኴ⏣⊛ᙪ࣭㧗ᶫ๛୍㑻 㸦⦅㸧 㸪pp. 7–16. ὶ⏕ែ◁㜵Ꮫ㸬ᮾிᏛฟ∧㸪ᮾி㸬 Ṋᒃ ⸅㸬2009㸬Ⓨ║༸ᨺὶࡢ᪉ἲ㸬୰ᮧᬛᖾ࣭㣤⏣ 㐶㸦⦅㸧 㸪pp. 66–71. Ᏺ ࡿ࣭ቑࡸࡍὶ㨶㸬㎰ᒣ⁺ᮧᩥ༠㸪ᮾி㸬 118 ㇂ཱྀ⩏๎࣭୰㔝 ⦾㸬2000㸬ᆅ⌫ ᬮᒁᡤⓗ⎔ቃᨩࡀῐỈ㨶㢮⩌㞟ཬ ࡰࡍ」ྜⓗᙳ㡪㸸࣓࢝ࢽࢬ࣒㸪ண ࡑࡋ࡚Ἴཬຠᯝ㸬㝣ỈᏛ㞧ㄅ㸪61: 79–94. Taniguchi, Y., F. J. Rahel, D. C. Novinger and K. G. Gerow. 1998. Temperature mediation of competitive interactions among three fish species that replace each other along longitudinal stream gradients. Can. J. Fish. Aquat. Sci., 55: 1894–1901. Taylor, E. B. 1991. A review of local adaptation in Salmonidae, with particular reference to Pacific and Atlantic salmon. Aquaculture, 98: 185–207. Teichert, M. A. K., A. Foldvik, T. Forseth, O. Ugedal, S. Einum, A. G. Finstad, R. D. Hedger and E. Bellier. 2011. Effects of spawning distribution on juvenile Atlantic salmon (Salmo salar) density and growth. Can. J. Fish. Aquat. Sci., 68: 43–50. Thompson, J. D., D. G. Higgins and T. J. Gibson. 1994. CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Res., 22: 4673–4680. Thurow, R. F. 1994. Underwater methods for study of salmonids in the Intermountain West. general technical report INT-GTR-307, USDA Forest Service, Intermountain Research Station, Ogden, Utah, USA. 28 pp. ᚫཎဴஓ࣭ᓊ ᙌ࣭ཎ ᚭ࣭⇃ᓮ ༤㸬2010㸬Ἑᕝᨺὶࡋࡓ㣴Ṫ࣐ࢦᡂ⇍ ぶ㨶ࡢ⏘༸ᗋ❧ᆅ᮲௳༸ࡢⓎ║⋡㸬᪥ᮏỈ⏘Ꮫㄅ㸪76: 370–374. ᚨගಇ㸪2004㸬ศ┴㸬ᮌᮧΎᮁ┘ಟ㸪࣡ࢼ㸪࣐࣓ࣖ㸪࣐ࢦࡢቑṪ⟶ ⌮ʊ᭷⏝㈨※⏕ែ⣔⟶⌮ᡭἲ㛤Ⓨᴗሗ࿌᭩ʊ㸬pp. 221–236. ᅜෆỈ㠃⁺ ᴗ༠ྠ⤌ྜ㐃ྜ㸪ᮾி㸬 ᚨගಇ㸬2003㸬ῐỈ⏕≀ቑṪᢏ⾡㛤Ⓨᴗ ࣓࣐࣡ࢦࡢ㓄㸬ᖹᡂ 14 ᖺᗘศ┴ෆỈ◊ᴗሗ࿌㸪pp. 12–13. Torgersen, C. E., C. V. Baxter, H. W. Li and B. A. McIntosh. 2006. Landscape influences on longitudinal patterns of river fishes: spatially continuous analysis of fish-habitat relationships. Am. Fish. Soc. Symp., 48: 473–492. Tymchuk, W. E., L. F. Sundström and R. H. Devlin. 2007. Growth and survival 119 trade-offs and outbreeding depression in rainbow trout (Oncorhynchus mykiss). Evolution, 61: 1225–1237. ୖ㔝ᓅᙪ࣭⏣୰ᶞ࣭ᒣ 㝯㸬2009㸬ᑠᨭὶ࠾ࡅࡿ࣡ࢼ㸪࣐࣓ࣖ⛶㨶ࡢ ᐃⅬ㸪⾜ືᅪ࠾ࡼࡧᦤ㣵㢖ᗘᑐࡍࡿ୧✀ᡂ㨶ࡢᙳ㡪㸬᪥ᮏỈ⏘Ꮫㄅ㸪 75: 802–809. Vannote, R. L., G. W. Minshall, K. W. Cummins, J. R. Sedell and C. E. Cushing. 1980. The river continuum concept. Can. J. Fish. Aquat. Sci., 37: 130–137. ⱝᯘ ㍤࣭୰ᮧᬛᖾ࣭ஂಖ⏣ோᚿ࣭ᒣ 㝯㸬2003㸬୰⚙ᑎ†ὶධἙᕝ⏕ᜥ ࡍࡿࢧࢣ⛉㨶㢮 2 ✀ࡢᙜṓ㨶ࡢ⏕ᜥ⎔ቃ㸬㨶㢮Ꮫ㞧ㄅ㸪50: 123–130. Watanabe, K. 1998. Parsimony analysis of the distribution pattern of Japanese primary freshwater fishes, and its application to the distribution of the bagrid catfishes. Ichthyol. Res., 45: 259–270. Watanabe, K. and M. Nishida. 2003. Genetic population structure of Japanese bagrid catfishes. Ichthyol. Res., 50: 140–148. ᒣᮏ ⪽㸬1991㸬࣡ࢼ㸪ࡑࡢ⏕ែ㔮ࡾ㸬ࡘࡾே♫㸪ᮾி㸬203 pp㸬 Yamamoto, S., K. Morita, I. Koizumi and K. Maekawa. 2004. Genetic differentiation of white-spotted charr (Salvelinus leucomaensis) populations after habitat fragmentation: spatial-temporal changes in gene frequencies Conserv. Genet., 5: 529–538. ᒣෆ 㸬1982㸬࣓࣡ࡢேᕤᏤ࣓࣡×࣐ࢦࡢ㓄ᐇ㦂ࢆ⤊ࢃࡗ࡚㸬 ᮌᮧⱥ㐀㸦⦅㸧pp. 119–124. ῐỈ㨶ቑห ࣐࣓࣭࣐ࣖࢦ≉㞟㸬㈈ᅋἲேῐỈ 㨶ಖㆤ༠㸪㜰㸬 ᾷ ኴᡃ࣭ᒣୗ๛ྖ࣭బᅵဴஓ࣭Ṋᮧ Ἠ࣭ᮌᮧΎᚿ㸬2001㸬୕㔜┴ᚿᦶᆅ᪉ ࡢἙᕝ㨶㢮┦㸬㨶㢮Ꮫ㞧ㄅ㸪48: 27–40. Yoshimura, C., T. Omura, H. Furumai and K. Tockner. 2005. Present state of rivers and streams in Japan. River Res. Appl., 21: 93–112. 120
© Copyright 2024 ExpyDoc
