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本論文・Full Paper 56(1): 7 －19 魚類学雑誌 本論文・Full Paper 56(1): 7 －19 1 〒631–8505 奈良市中町3327–204 近畿大学農学部 2 〒631–0061 奈良市富雄元町1–21 おさかな総合研究所 3 〒819–0396 福岡市西区元岡744 九州大学大学院工学研究院環境都市部門 4 〒870–0802 大分市高崎山下海岸 大分マリーンパレス水族館「うみたまご」 （2008年5月12日受付；2008年12月20日改訂；2008年12月26日受理） キーワード： mtDNA，PCR-RFLP，生物地理，古瀬戸内川 Emi Kitagawa, Jun Nakajima, Kazuo Hoshino and Tadao Kitagawa*. 2009. Geo- graphic distribution and biogeographical origin of the spined loach genus Cobitis Japanese Journal of in north-eastern Kyushu Island, Japan. Japan. J. Ichthyol., 56(1): 7–19. Ichthyology © The Ichthyological Society of Japan 2009 Abstract In order to clarify the distribution pattern of three Japanese spined loach species’ complexes in the genus Cobitis (‘yamato’, biwae, and striata com- plexes) in north-eastern Kyushu Island, 106 specimens obtained from 19 drainage systems were morphologically and genetically examined. Although that region has been previously considered occupied only by the ‘yamato’ complex, some popula- tions of the biwae and striata complexes were confirmed, the distribution patterns being explained by the paleo-Setouchi drainage pattern during the most recent Pleistcene glacial periods. Distinctive populations within the middle race of striata complex included 2 new local forms in Fukuoka Prefecture (the Setouchi form, Hakata form and Onga form). *Corresponding author: Faculty of Agriculture, Kinki University, Nara 631–8505, Japan (e-mail: [email protected]) 本産シマドジョウ属魚類Cobitisの分類は歴 シマドジョウとスジシマドジョウのグループをタイ 日史的に混乱しており未だに決着を見ていない リクシマドジョウ群とまとめ，それぞれをヤマト が，形態学的分類（池田, 1936, 1937a; Okada and シマドジョウ亜群，スジシマドジョウ亜群に区分 Ikeda, 1939; Mizuno, 1970; Saitoh and Aizawa, 1987; している．しかし，遺伝学的研究からヤマトシマ Suzawa, 2006）および核型・アロザイム・ DNA分 ドジョウのグループはシマドジョウとスジシマド 析等による遺伝学的研究結果 (Ueno and Ojima, ジョウのグループの個体間の交雑を起源とする異 1976; Saitoh et al., 2000; Kitagawa et al., 2001, 質 4 倍体で，このグループの名称の根拠となった 2003a,b, 2005; Shimizu et al., 2004) を総合すると， ヨーロッパ産のタイリクシマドジョウCobitis taenia 従来，シマドジョウ，ヤマトシマドジョウ（また とは全く異なるものであること (Ueno and Ojima, はタイリクシマドジョウ），スジシマドジョウ，イ 1976; Sezaki et al., 1994)，さらにmtDNA分析によ シドジョウとよばれてきた4グループに区分でき， りスジシマドジョウのグループとタイリクシマド それぞれのグループが複数の独立種を含んでいる ジョウC. taeniaが近縁関係にないことが明らかに ことが明らかになっている．斉藤 (2002) では，池 なっていることから (Kitagawa et al., 2005)，タイリ 田 (1936, 1937a) と整合させるため，従来のヤマト クシマドジョウ群として両者をまとめることは不 8 北川えみほか 適当である．本報告では，Saitoh et al. (2000) にお （ 宮崎県版レッドデータブック作成検討委員会, ける英語表記を基本として，それぞれの和名，学 2000）．以上の結果から，九州にはシマドジョウ種 名をシマドジョウ種群 Cobitis biwae complex，スジ 群が生息していないとの見解が近年まで受け入れ シマドジョウ種群 Cobitis striata complex，ヤマト られてきた（君塚，2002）．九州における多くのシ シマドジョウ種群 Cobitis sp. ‘yamato’ complex，イ マドジョウ種群の生息情報が否定される一方で， シドジョウ種群 Cobitis takatsuensis complexとよ 池田 (1937b) やOkada and Ikeda (1939) が報告した ぶ．なお，Kitagawa et al. (2005) によるmtDNA分 大分県大分川水系におけるシマドジョウ種群の分 析の結果に基づくと，アジア大陸東部に生息して 布は再検証されないままになっていたが，近年， いるCobitis lutheriとCobitis tetralineataもスジシマ 北川ほか (2004) が形態およびmtDNA分析に基づ ドジョウ種に含めることが適当である．これら日 き大分川水系においてシマドジョウ種群（4倍体） 本産4 種群のうちイシドジョウ種群を除く3 種群 が生息していることを確認した．これに続き高濱 は，いずれも河川中・下流に生息し互いに形態的 ほか (2006) により，大分川水系に隣接する大野川 に類似しており，同定の困難さが本分類群内にお 水系からもシマドジョウ種群（4 倍体）の生息が ける分類の混乱の主たる要因となってきた． 報告されている． これまで，九州地方におけるシマドジョウ類の 一方，スジシマドジョウ種群についても，過去 分布に関する情報に混乱が生じており，特にシマ にヤマトシマドジョウ種群の2倍体種族とされてい ドジョウ種群に関する分布情報の混乱は著しかっ た有明海側流入河川の集団（Ueno and Ojima, た．その主たる原因は，Temminck and Schlegel 1976; 上野ほか，1980）が，スジシマドジョウ種 (1846) において記載されたシマドジョウC. biwae 群に属することが判明し，現在ではスジシマド （原記載ではC. taenia japonica）の模式産地が，長 ジョウ種群小型種九州型として扱われている（斉 崎県とされていたことにある（澤田・相澤，1983）． 藤，2002）．また近年，今川水系よび近隣河川か また，池田 (1936) の報告したシマドジョウの標本 ら同種群中型種の分布が報告されている（福岡県 に福岡県産の個体が含まれていたこと，池田 環境部自然環境課，2001; 中島ほか, 2006）．さら (1937b) およびOkada and Ikeda (1939) において大 に，同県の遠賀川水系から新たにスジシマドジョ 分県大分川水系にシマドジョウの分布が報告され ウ種群小型種（型不明）が記録され（福岡県環境 ていたこと，宮崎県大淀川水系からシマドジョウ 部自然環境課，2001）， この集団は中島ほか が報告されたこと（皆森，1951），1970年代まで (2006) においてシマドジョウ属不詳種Aとして記 の図鑑においてシマドジョウの分布域に九州東部 録されている．また同じく中島ほか (2006) におい が含まれていること（中村，1963; 宮地ほか， て，福岡市を流れる河川からスジシマドジョウ種 1976），宮崎県川内川水系上流部の個体がシマド 群小型種に類似するさらなるシマドジョウ属不詳 ジョウとして扱われたこと（山根，1969）など， 種集団の存在が報告され，シマドジョウ属不詳種 九州におけるシマドジョウ種群の分布を示唆する Bとして記録されている． 情報が複数存在していた．しかし，池田 (1937a) このように，九州，特に北東部においてシマド およびOkada and Ikeda (1939) において福岡県筑後 ジョウ類の分布に関する新たな情報が蓄積されつ 川水系や長崎県下などの九州北西部に生息してい つあり，より詳細な再検証が必要である．そこで るのはシマドジョウ種群ではなくヤマトシマドジョ 本研究では，現在もシマドジョウ類の分布情報が ウ種群であることが報告され，Kimizuka and 混乱している九州北東部から得られた標本を形態 Kobayashi (1983) も九州北西部地域からはヤマト 学的・遺伝学的に精査し，同地域に生息する3 種 シマドジョウ種群しか得られなかったことを報告 群の分布について整理するとともに，その成立起 している．また，Kimizuka and Kobayashi (1983) 源について考察した．また，先に述べた遠賀川水 は，山根 (1969) の標本を再調査し，これがヤマト 系と福岡市の河川の2 つのシマドジョウ属不詳種 シマドジョウ種群であること，さらに同産地から 集団について，これがスジシマドジョウ中型種の 採取した個体についても形態学的，核学的見地か 地方型に該当すると考えられたため，保全上の理 らすべてヤマトシマドジョウ種群と同定している． 由から仮の呼称を与えたので，これについても報 皆森 (1951) による大淀川水系におけるシマドジョ 告する． ウ種群の分布についても，その後の調査研究にお いてヤマトシマドジョウ種群しか得られていない 九州北東部のシマドジョウ属の分布 9 Table 1. Comparison of character conditions of three Japanese species’ complexes of Cobitis from eastern Kyushu. Specimens with identical character status in each locality grouped in single line No. Composition Position mtDNA*4 Drainage Lamina Color Identifica- in n Sex*1 of of dorsal Ploidy (Hha I, system circularis pattern tion*5 map vertebrae*2 fin*3 Taq I) Fukuoka 1 Naka 1 ? circle spotted A, B, C 15–17 (y, y) yamato 1 / spotted A, B, C 15–17 (y, y) yamato 1 / spotted A, B, C 15–17 (s, s) striata 2 Mikasa 1 ? circle spotted A, B, C 15–17 tetraploid (y, y) yamato 2 / spotted A, B, C 15–17 tetraploid (y, y) yamato 3Tatara 1 / spotted A, B, C 15–17 (s, s) striata 1 / spotted A, B, C 15–17 diploid (s, s) striata 1 ? circle striped C 15–17 diploid (s, s) striata 4 Daikon 2 / spotted A, B, C 15–17 tetraploid (y, y) yamato 5 Saigo 1 ? circle spotted A, B, C 15–17 tetraploid (y, y) yamato 2 / spotted A, B, C 15–17 tetraploid (y, y) yamato 6 Onga 1 ? circle spotted B, C 15–17 diploid (s, s) striata 5 ? circle spotted A, B, C 15–17 tetraploid (y, y) yamato 1 ? circle spotted A, B, C 15–17 (yamato or striata) 2 / spotted A, B, C 15–17 diploid (s, s) striata 5 / spotted A, B, C 15–17 tetraploid (y, y) yamato 1 / spotted A, B, C 15–17 diploid (s, s) striata 2 / spotted A, B, C 15–17 — 1— spotted tetraploid (y, y) yamato 7 Murasaki 2 / spotted A, B, C 15–17 tetraploid (y, y) yamato 1— spotted tetraploid (y, y) yamato 8 Ima 1 ? circle striped A, B, C 15–17 diploid (s, s) striata 1 / spotted A, B, C 15–17 tetraploid (y, y) yamato 1 / striped A, B, C 14 diploid (s, s) striata 3 / striped A, B, C 15–17 diploid (s, s) striata 2 / spotted A, B, C 15–17 — 9 Harai 1 / spotted A, B, C 15–17 tetraploid (y, y) yamato 1 / striped A, B, C 15–17 (s, s) striata 1 / spotted A, B, C 15–17 (y, y) yamato 1— spotted (y, y) yamato 1 / spotted A, B, C 14 (yamato or striata) 3 / spotted A, B, C 15–17 — 10 Otonashi 1 / spotted A, B, C 15–17 tetraploid (y, y) yamato 11 Kii 1 ? circle striped A, B, C 15–17 (s, s) striata 2 / spotted A, B, C 15–17 (y, y) yamato Oita 12 Yamaguni 4 / spotted A, B, C 15–17 — 13 Iroha 1 / spotted A, B, C 13 (striata) 1 / spotted B, C 15–17 (biwae or striata) 2 / spotted A, B, C 15–17 — 14 Yakkan 6 / spotted A, B, C 15–17 — 15 Yorimo 2 / spotted A, B, C 15–17 (y, y) yamato 1— spotted (y, y) yamato 16 Yasaka 1 ? circle spotted A, B, C 15–17 tetraploid (y, y) yamato 3 / spotted A, B, C 15–17 — 17 Oita 1 ? beak like spotted B 18 tetraploid (b, b) biwae Oita 17 2 ? beak like spotted B 18 tetraploid (b, b) biwae 10 北川えみほか よび神奈川県立生命の星・地球博物館 (KPM-NI) に登録・保管されており，中島ほか (2006) により シマドジョウ類標本の精査方法は北川ら (2004) 福岡県下から得られたシマドジョウ属不詳種の登 に従った．すなわち，形態学的分析には相澤 録個体（登録番号に*を示した）も含まれている． (1981) および澤田・相澤 (1983) によって日本産シ 採集地，登録番号，個体数は以下のとおりである． マドジョウ属の有効な分類形質とされている4 形 なお，各集団の個体数保全を考慮し，ここでの採 質（雄の胸鰭基部骨質盤の形状，体側部の斑紋， 集地の記載は県名と水系のみにとどめる．福岡県 腹椎・尾椎数の構成および背鰭の位置）の精査を， 那珂川水系 (FKUN 033589, 033590, 033806*, 倍数性の判定にはSezaki and Kobayasi (1978) に基 nϭ3)，同御笠川水系 (FKUN 033725-033727, nϭ3)， づく血球サイズの計測を，ミトコンドリアDNA 同多々良川水系 (FKUN 033758, 033759, 033807*, (mtDNA) による遺伝学的分析にはKitagawa et al. nϭ3), 同大根川水系 (FKUN 033728, 033529, nϭ2)， (2003a,b, 2005) に基づく ND1遺伝子領域を対象と 同西郷川水系 (FKUN 0337730-033732, nϭ3)，同 した制限断片長多型 (RFLP) 分析を行った． 遠賀川水系 (FKUN 033553-033555, 033587*, 1991年 8 月から2005年4 月までの期 033588*, 033733*, 033734*, 033735 –033745, 間に九州北東部の河川19水系 (Table 1; Fig. 1) か nϭ18)，同紫川水系 (KPM-NI 9148–9150, nϭ3)， ら著者らおよび研究協力者が採集した106個体を 同今川水系 (FKUN 033556, 033557, 用いた．また，シマドジョウ類の分布情報に乏し 033583–033586, 033746, 033747, nϭ8)，同祓川水 い地域である九州南東部の宮崎県一ツ瀬川からも 系 (FKUN 033558–033561, 033748–033750, 033752, 2 個体を得たので，これも分析対象に加えた．こ nϭ8)，同音無川水系 (FKUN 033553, nϭ1)，同城 れらの標本は近畿大学農学部水産学科 (FKUN) お Table 1. (Continued) No. Composition Position mtDNA*4 Drainage Lamina Color Identifica- in n Sex*1 of of dorsal Ploidy (Hha I, system circularis pattern tion*5 map vertebrae*2 fin*3 Taq I) Oita 17 2 ? beak like spotted B, C 15–17 tetraploid (b, b) biwae 1 ? beak like spotted B, C 15–17 tetraploid (b, b) biwae 1 ? beak like spotted B, C 15–17 (b, b) biwae 2 / spotted B 15–17 tetraploid (b, b) biwae 3 / spotted A, B, C 15–17 tetraploid (b, b) biwae 1 / spotted B, C 15–17 (b, b) biwae 1 / spotted B, C 15–17 (biwae or striata) 1— spotted tetraploid (b, b) biwae 18 Ono 2 ? beak like spotted B 18 tetraploid (b, b) biwae 1 ? beak like spotted B, C 15–17 tetraploid (b, b) biwae 1 / spotted B 18 tetraploid (b, b) biwae 3 / spotted B 15–17 tetraploid (b, b) biwae 1 / spotted B, C 15–17 tetraploid (b, b) biwae 1 / spotted B 18 (b, b) biwae 19 Banjo 1 ? circle spotted A, B, C 15–17 tetraploid (y, y) yamato 5 / spotted A, B, C 15–17 tetraploid (y, y) yamato Miyazaki 20 Hitotsuse 1 ? circle spotted A, B, C 15–17 (y, y) yamato 1 / spotted A, B, C 15–17 (y, y) yamato *1 Specimens less than 40 mm SL regarded as immature (Ϫ) based on Ikeda (1932). *2 Letters indicate range of vertebral composition shown in Fig. 3. *3 Represented by vertebral count of before insertion of
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