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Geographic Distribution and Variation of Three Species of Ninespine Sticklebacks(Pungitius Tymensis,P.Pungitius and P.Sinensis)In Hokkaido**

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Geographic Distribution and Variation of Three Species of Ninespine Sticklebacks(Pungitius Tymensis,P.Pungitius and P.Sinensis)In Hokkaido** 魚 類 学 雑 誌 Jananese Journal of Ichthyology 31巻3号1984年 Vol.31,No.31984 北 海 道 に お け る トミヨ属 魚 類3種 の地 理 的 分 布 と形 態 変 異* 高 田 啓 介 ・後 藤 晃 ・濱 田 啓 吉 Geographic Distribution and Variation of Three Species of Ninespine Sticklebacks(Pungitius tymensis,P.pungitius and P.sinensis)in Hokkaido** Keisuke Takata,Akira Goto and Keikichi Hamada (Received March 29,1984) The patterns of geographic distribution and morphological variation of three species of nine spine sticklebacks(Pungitius tymensis,P.pungitius and P.sinensis)in Hokkaido,Japan,were in- vestigated.The range of geographic distribution of P.tymensis was essentially restricted to the following three areas,Teshio District of the northern part of Hokkaido,Ishikari District of the central part of Hokkaido and Kushiro-Nemuro District of the eastern part of Hokkaido.On the other hand,P.pungitius and P.sinensis were distributed more widely and continuously on Hokkaido,compared with the distribution of P.tymensis.P.pungitius was distributed continuous- ly in the rivers of the Pacific slope from Nemuro District to Iburi District of the central part of Hokkaido,though,discontinuously in the rivers along the Japan Sea and the Okhotsk Sea slopes. In most rivers facing the Nemuro Straits,P.pungitius was not collected.P.sinensis was dis- tributed in only a few rivers of the Pacific slope from Tokachi District to the Oshima Peninsula of the southern part of Hokkaido,but was continuously distributed in the rivers facing the Tsugaru Straits,the Okhotsk Sea and the Nemuro Straits. In three rivers examined(Osatsu,Fukunaga and Bettoga)P.tymensis exninntea more of an upstream distribution than did P.pungitius and/or P.sinensis.In two rivers(Fukunaga and Bettoga),P.pungitius and P.sinensis did not occur together along the lengths of the river,but were distributed with a different range respectively.The morphological characteristics of P. tymensis were quite different from the other two species and its meristic characters varied less than those of P.pungitius and P.sinensis.Difference in morphology between P.pungitius and P. sinensis was not significant except for number of lateral plates.The number of vertebrae,in both P.pungitius and P.sinensis decreased gradually from north to south on the Japan Sea slope,though the other meristic characters did not indicate such a geo-cline.The patterns of geographic vari- ations were most similar between the two species in the rivers of the Japan Sea slope. Comparison of morphological characteristics and distribution patterns suggest that P.tym- ensis is distinguished as an independent species from P.pungitius and P.sinensis.Although the comparative study does not necessarily show that P.pungitius and P.sinensis are different species from each other,it does not show that they are completely identical.The latter two species show different distribution patterns in the coexisting rivers and are distributed contiguously and allopat- ricaly in Hokkaido.It is supposed,therefore,that P.pungitius and P.sinensis in Hokkaido should be distinguished from each other at a lower taxonomic level than species. (Laboratory of Embryology and Genetics,Faculty of Fisheries,Hokkaido University,Hakodate 041,Japan) ト ミ ヨ(P.sinensis),ミ ナ ミ ト ミ ヨ(P.sinensis f.kai- 日 本 列 島 に 分 布 す る ト ミ ヨ 属 魚 類 に は,イ バ ラ ト ミ ヨ barae),エ ゾ ト ミ ヨ(P.tymensis)が 知 ら れ て お り(宮 (Pungitius pungitius),ム サ シ ト ミ ヨ(P.pungitius f.sp.), 地 ほ か,1976),北 海 道 に は,こ の うち ミナ ミ ト ミ ヨ と *北 海 道 の淡 水 魚 に 関す る研 究-V ム サ シ ト ミ ヨ を 除 く3種 が 分 布 す る. **Studies on the Freshwater Fish in Hokkaido, Japan-V 北 海 道 内 の3種 の 地 理 的 分 布 に つ い て は,池 田(1933), ―312― 高 田 ・後藤 ・濱 田:ト ミ ヨ属 の分 布 と変 異 小 林(1957,1959)の 研 究 が あ る.そ れ に よる とエ ゾ ト い て 再 検 討 した. ミヨは 北 海 道 西 部 の主 と して 日本 海 へ 注 ぐ水系 に,イ バ 調 査 地 点 の 概 要 お よ び 調査 方法 ラ トミヨは 日本 海 側 と太 平 洋 側 に注 ぐ水系 に,ト ミ ヨは 流 程 分 布 に 関 す る調 査 は3河 川 で 行 った.エ ゾ ト ミヨ 太 平 洋 側 と オ ホ ー ツ ク海 側 に注 ぐ水 系 に 分 布 す る.そ の と イバ ラ トミヨの2種 が共 存 す る道 央 地 方 の 石 狩 川 水系 後,石 城(1967)は,エ ゾ ト ミヨが 北 海 道 東 部根 釧 地 域 2次 支 流 の長 都 川 で は,1979年5月16日 お よび11月 に も分 布 す る こ とを 確 認 した.し か し,こ れ らの報 告 は 18日 か ら22日 まで の2回,ト ミ ヨを 含 め た3種 が 共 存 い ず れ も断 片 的 で あ り,北 海 道全 域 にわ た る3種 の 分 布 パ タ ー ンの特 徴 を 十 分 捉 え た とは言 いが た い. す る道 東 地 方 の 別 当 賀 川 と道 北 地 方 の天 塩 川 水 系2次 支 流 の福 永 川 で,そ れ ぞれ1982年8月8・9日 お よび1981 池 田(1933)は 日本 列 島 とサ ハ リ ンに おい て,本 属 魚 類 の 形態 に変 異 が 大 きい こ とを み い だ し,特 に 背 棘 数 と 年6月13日 か ら17日 に行 わ れ た(Fig.1). 鱗 板 数 を と りあ げ,3種 間 の 地理 的変 異 を比 較 した.そ 長 都 川 と福 永 川 で は,下 流 よ り上 流 に 向 って ほぼ 等 間 して,エ ゾ トミ ヨで は サ ハ リン産 と北 海 道 産 の鱗 板 数 に 隔 に9調 査 地 点(St.1～9),ま た 別 当 賀 川 で は7調 査 地 明 瞭 な 差 が 存在 す る こ と,イ バ ラ ト ミョ と トミヨの2種 点(St.1～7)を 設 け た.各 地 点 の河 川 形 態 を 可 児(1944) に おい て は そ の 鱗 板 数 と背 棘 数 が 南 方 へ行 くに従 い 減 少 に従 って 区分 す る と,長 都 川 で はSt.1はBc型,St. す る こ と,お よび イバ ラ トミヨ と ト ミヨの それ らの減 少 2～6はBb型 で あ り,St.7～9はAa-Bb移 行 型 で あ っ の 程 度 に は そ れ ぞ れ 特 異 性 が 認 め られ る こ とを示 した. た.ま た,福 永 川 では,ペ ン ケ沼 か らの流 出 お よび流 入 一 方 ,小 林(1959)は 人 為 交 配 に よ って 得 た エ ゾ トミ ヨ 地 点 に 位 置 す るSt.1とSt.2はBc型 で あ り,St.3は とイ バ ラ ト ミヨ,な らび に イ バ ラ ト ミヨ と トミヨのF1雑 Bb-Bc移 行型,St.4～7はBb型,St.8,9はAa-Bb 種 の形 態 が,北 海 道 の混 生 域 に お いて み い だ され る両 種 移 行 型 で あ った.別 当賀 川 の河 川形 態 は,St.1はBc の 中 間 型 の 形 態 とそ れ ぞ れ 類 似 して い る こ とか ら,そ れ 型,St.2～4はBb型,St.5～7はBb-Bc移 行 型 で あ らが 一 部 で 自然 交 雑 して い る可 能 性 を 指 摘 した.最 近, った.各 調 査地 点 で は追 い 込 み網(間 口1m,目 合3mm) 田 中(1982)は 東 北 地 方 の イバ ラ トミヨ と トミヨの形 態 を 用 い て く り返 し採 集 を 行 うこ とに よ って各 種 の 生 息 の を詳 し く調 査 し,混 生 域 に おい て,鱗 板 数 の 他 に幾 つか 有 無 を 確 め,ま た,採 集 され た 場 合 に は 全個 体 数 に 対 す の 計 数 的 形 質 の平 均 値 に も両 種 間 で 有 意 差 が あ る こ と, る各 種 の 個 体 数 比率 を求 めた. お よび 中 間型 は ご く少 数 しか出 現 しない こ とか ら,両 種 北 海 道 内 に お け る分 布 に つ い て は,1979年3月25日 の 間 に 何 らか の生 殖 的 隔 離 が存 在 して い る こ とを 示 唆 し か ら4月27日 に 道 央地 方,同 年8月6日 か ら8月15日 た. に道 北 ・道 南 地 方 の 日本 海 側,8月28日 か ら9月8日 この よ うに,ト ミヨ属3種 の形 態 的 特 徴 や そ の 変 異 に お よび1982年8月8日 か ら8月12日 に 太 平 洋側,1980 種 特 異 性 が あ る か否 か,及 び そ れ らの 間 の生 殖 的 隔 離 の 年7月26日 か ら8月4日 に道 東 地 方,1981年8月18日 程 度 に つ い て は,必 ず し も意 見 の一 致 を み てい な い.こ か ら21日 に道 北 地 方 とオ ホ ー ツ ク海 側 の各 河 川 で調 査 うした 未 解 決 の 問題 と関連 して,こ れ ら3種 の 分類 に 関 を行 った.各 調 査 河 川 で は追 い込 み 網 の他,投 網(網 丈 して も異 な る 見 解 が あ る.池 田(1933,1950),石 城 3m,目 合12mm)を 使 用 し,一 部 の河 川 の河 口部 で は (1967),宮 地 ほ か(1976)は3種 を それ ぞれ 独 立 した 種 小 型 地 曳 網 を用 い た.ま た,こ れ らの調 査 地 点 以 外 に, と した の に 対 し,Okada(1960),Manzing(1969), 各 方 面 か ら標 本 を 借 用 す る こ とに よ り,分 布 記 録 の不 備 Wootton(1976)は エ ゾ ト ミヨ,ト ミヨを イバ ラ トミヨ を補 った.
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