琵琶湖におけるニゴロブナ Carassius Auratus Grandoculis 種苗の 放流水域としてのヨシ帯の重要性と放流事業の課題

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琵琶湖におけるニゴロブナ Carassius Auratus Grandoculis 種苗の 放流水域としてのヨシ帯の重要性と放流事業の課題 Nippon Suisan Gakkaishi 77(5),822833 (2011) 琵琶湖におけるニゴロブナ Carassius auratus grandoculis 種苗の 放流水域としてのヨシ帯の重要性と放流事業の課題 藤原公一,1,2a臼杵崇広,1a 根本守仁,1 松尾雅也,3 竹岡昇一郎,3 田中 満,3 北田修一2 (2011 年 2 月 23 日受付,2011 年 5 月 15 日受理) 1滋賀県水産試験場,2東京海洋大学海洋科学部,3琵琶湖栽培漁業センター The importance of the reed zone as a release region for larval and/or juvenile nigorobuna Carassius auratus grandoculis in Lake Biwa, and problems for stock enhancement KOH-ICHI FUJIWARA,1,2aTAKAHIRO USUKI,1a MORIHITO NEMOTO,1 MASAYA MATSUO,3 SYOICHIRO TAKEOKA,3 MITSURU TANAKA3 AND SHUICHI KITADA2 1Shiga Prefecture Fisheries Experiment Station, Hikone, Shiga 5220057, 2Faculty of Marine Science, Tokyo University of Marine Science and Technology, Minato, Tokyo 1088477, 3Stock Enhancement Center of Lake Biwa, Kusatsu, Shiga 5250005, Japan EŠective release strategies for nigorobuna Carassius auratus grandoculis in Lake Biwa were investigated by release-recapture experiments of otolith marked nigorobuna with ALC. As a result, nigorobuna arrived at the juvenile stage exceeding approximately 16 mm in standard length and released to the Phragmites australis reed zone showed a rapid increase in survival rate. A remarkable decrease in the survival rate of stocked ˆsh in the lake was observed in cases of 1) releasetothereed-freelakeshore,2) reduction of the depth of ‰ooded reed zone ac- cording to draw down of water level by artiˆcial manipulation, 3) growthretardationofreedinspringbyexces- sive cutting management of reed in winter, and 4) excessive release of nigorobuna larvae to a limited reed zone. Larvae-release techniques which overcome the problems and reed zone-independent release methods for nigorobuna should be developed in the future. キーワードALC 耳石標識,再捕率,生残率,ニゴロブナ,標識放流,琵琶湖,放流効果,ヨシ帯 ニゴロブナ Carassius auratus grandoculis は,コイ科 ことが明らかになった。6) また,ヨシ帯奥部は本種仔魚 フナ属に属する琵琶湖の固有亜種13)で,滋賀県の食文 が好んで摂餌する微小甲殻類(ミジンコ亜綱)69)が豊富 化を代表するフナズシ4,5)の材料に用いられる。このた で,貴重な摂餌の場となっているが,溶存酸素濃度が著 め本種は琵琶湖漁業にとって非常に重要な漁獲対象とな しく低く,一般には魚類が生息しにくい環境にあること っており,琵琶湖ではその資源を積極的に増やすため, が判った。6) しかし本種仔魚は生理的にそのような貧酸 琵琶湖栽培漁業センターと滋賀県漁業協同組合連合会に 素環境に適応しており,ヨシ帯奥部は本種仔魚にとって よって種苗放流事業が行われている。著者らは,この事 捕食者からの退避地として機能していると考えられ 業を効果的に進めるうえで必要な種苗放流適地について た。6,10) それでは,本種の仔魚や稚魚が琵琶湖で生き残 知見を得るため,本種の養成親魚から得た仔稚魚をよく るうえでヨシ帯は不可欠であろうか。この疑問を解決す 発達したヨシ帯に放流して初期生態を調査した。その結 るため著者らは,ALC(アリザリンコンプレクソン) 果,本種は仔魚期にはヨシ帯の岸辺付近(以下,奥部) で耳石に標識(以下,ALC 標識)11,12)を付けて識別した へ積極的に集まり,その水域を発育の場として利用する ニゴロブナ種苗を琵琶湖のヨシ帯,近隣の砂浜およびそ Tel81749281611.Fax81749252461.Emailfujiwara-koichi@pref.shiga.lg.jp a 現所属滋賀県水産課(Shiga Prefecture Fisheries Management Division, Otsu, Shiga 5208577, Japan) ニゴロブナ種苗放流の適地と課題 823 れらの沖合水域へ放流し,琵琶湖で再捕調査を行い,本 県水産試験場(以下,滋賀水試。滋賀県彦根市)へ搬入 種仔稚魚が琵琶湖で生き残るうえでヨシ帯が非常に重要 し,琵琶湖水を流したパンライト製または FRP 製の水 な役割を果たしていることを示した。また,ヨシ帯への 槽に収容管理して,数日後にふ化仔魚を得た。これら ALC 標識種苗の放流調査により,高い放流効果が期待 ふ化仔魚は藤原13)の手法に従って飼育した。種苗各群 できる種苗サイズ(発育段階)を明らかにし,併せて, の生産概要は Table 1 にまとめた。 ヨシ帯が持つ本種仔稚魚のナーサリー機能は,治水を目 標識 種苗の耳石(礫石)に藤原11)および藤原ら12) 的とした琵琶湖の人為的な水位低下やヨシ帯の管理を目 の方法で ALC 標識を付けた。なお,放流後に再捕され 的とした冬季のヨシ刈り取り,焼き払い,およびヨシ帯 る種苗の放流時の礫石の長径(以下,耳石長)を推定す への過度の種苗放流によって低下することを明らかにし るため,原則として最終の標識操作は各群とも放流する た。 前日から放流直前までの 24 時間かけて行い,放流時の 種苗の耳石外縁を ALC で染めた。Table 1 には種苗各 材料および方法 群へ付けた ALC 標識についても併せて示した。 供試魚 1992 年から 1995 年にかけて,毎年原則と ヨシ帯の重要性の検討 ニゴロブナの種苗放流適地と してニゴロブナの産卵盛期の 5 月上旬に,琵琶湖栽培 してのヨシ帯の重要性を検討するため,Fig. 1 に示した 漁業センター(以下,栽培漁業センター。滋賀県草津市) 滋賀県長浜市湖北町海老江(以下,海老江)の琵琶湖沿 のニゴロブナ親魚養成池へ人工産卵藻(キンラン,川島 岸のヨシ帯(Site E),近隣の同町尾上(以下,尾上) 商事)を設置して,ニゴロブナに自然産卵させた。産卵 の砂浜(Site O)およびそれらの沖合(竹生島の南方) 翌日には,卵が付着した人工産卵藻を活魚輸送車で滋賀 の水深約 80 m の水域(Site CS)へ本種種苗 94E 群, Table 1 Details of the ALC marked1 nigorobuna Carassius auratus grandoculis released in Lake Biwa Standard length Group Spawning Hatching Release Number Release Age at release ALC mark type2 name date date date released site 3 Mean±SD N DAH4 mm 92M11 5/11/1992 5/17/1992 5/19/1992 2 Dot 5.53±0.32 124 1,074,000 M 92M12 5/11/1992 5/17/1992 5/27/1992 10 Dot and ring 7.61±0.95 100 295,000 M 92M13 5/11/1992 5/17/1992 6/ 6/1992 20 Dot and 2 rings 9.73±1.43 135 49,000 M 92M14 5/11/1992 5/17/1992 6/16/1992 30 Dot and 3 rings 15.12±2.47 113 26,000 M 92M15 5/11/1992 5/17/1992 6/26/1992 40 Dot and 4 rings 16.06±2.80 140 29,300 M 92M21 6/11/1992 6/15/1992 6/17/1992 2 Dot 5.74±0.42 112 313,000 M 92M22 6/11/1992 6/15/1992 6/23/1992 8 Dot and ring 6.99±0.47 129 193,000 M 92M23 6/11/1992 6/15/1992 7/ 1/1992 16 Dot and 2 rings 8.07±0.92 142 199,000 M 92M24 6/11/1992 6/15/1992 7/ 9/1992 24 Dot and 3 rings 10.67±1.48 180 91,000 M 92M25 6/11/1992 6/15/1992 7/17/1992 32 Dot and 4 rings 14.76±2.72 124 25,800 M 94E 5/ 9/1994 5/15/1994 6/26/1994 42 5 rings 16.63±3.65 250 132,000 E 94O 5/ 9/1994 5/15/1994 6/26/1994 42 4 rings 15.47±3.52 250 121,000 O 94CS 5/ 9/1994 5/15/1994 6/26/1994 42 3 rings 17.62±4.31 250 98,000 CS 94M 5/ 9/1994 5/15/1994 6/28/1994 44 3 rings 18.14±4.50 240 95,000 M 95M 4/29/1995 5/ 5/1995 7/ 6/1995 62 3 rings 20.62±3.62 244 30,000 M 95C1 5/25/1995 5/31/1995 7/ 6/1995 36 Dot and 2 rings 16.80±3.69 289 84,000 C 95C2 4/29/1995 5/ 5/1995 7/ 6/1995 62 2 rings 20.62±3.53 257 32,000 C 93E 5/12/1993 5/18/1993 7/ 1/1993 44 4 rings 16.22±3.30 203 194,000 E 95E 5/15/1995 5/20/1995 7/ 4/1995 45 3 rings 17.28±3.34 323 37,000 E 95CN1 5/15/1995 5/20/1995 8/ 1/1995 73 3 rings 23.30±6.06 218 17,000 CN 95CN2 5/4/1995 5/10/1995 8/ 1/1995 83 6 rings 25.52±6.14 200 66,000 CN 95H 5/15/1995 5/20/1995 7/11/1995 52 4 rings 15.97±3.37 314 120,0005 H 95A 5/15/1995 5/20/1995 7/11/1995 52 3 rings 16.47±3.71 323 109,0006 A 1 Refer to the text for ALC marking. 2 Except for the mark of only a dot, ALC marks were mutually discriminable from the size, interval, etc. of rings. 3 Refer to Fig. 1. 4 Days after hatching. 5 Non-marked ˆsh of 81,000 (23.21±SD3.70 mm in standard length) were released with marked group 95H. 6 Non-marked ˆsh of 506,000 (20.13±SD3.19 mm in standard length) were released with marked group 95A. 824 藤原,臼杵,根本,松尾,竹岡,田中,北田 94O 群および 94CS 群を,ともに 1994 年 6 月 26 日に 放流した。沿岸部への放流種苗(94E 群,94O 群)は, 滋賀水試から活魚輸送車で群毎に放流水域付近まで運搬 し,バケツリレーして放流した。沖合への放流種苗 (94CS 群)は,尾上にある朝日漁港で活魚輸送車から 漁船の活魚水槽へ移し替えて沖合まで運搬して放流し た。なお,放流時に各群から 250 尾を無作為抽出して 冷凍保存し,後日,解凍後,これら全個体の標準体長 (以下,体長)を測定して群毎に平均値と SD を計算す るとともに,体長 2mm間隔の相対度数を求め,同群の 放流尾数にこれを乗じて,体長 2mm間隔毎の放流尾数 を推定した。同年 11 月 5 日から翌年 4 月 15 日には, 琵琶湖の北湖盆(北湖)で小型機船底曳き網を用いてニ ゴロブナ標本を 13,525 尾採集した。その標本を冷凍保 存し,後日解凍して体長と体重を測定した後,耳石を調 べて標識魚を特定し,さらに各群を確定して,群毎の再 捕尾数を求めた。また,各群の耳石の最外部の ALC リ ングの長径(=放流時の耳石長)を蛍光顕微鏡(OP- TIPHOT,XTEFD,ニコン)の接眼マイクロメータ Fig. 1 Release sites of nigorobuna Carassius auratus を用いて計測し,藤原らの方法12)で群毎の耳石長体長 grandoculis. Nigorobuna with diŠerent ALC mark shown in Table 1 were released in each site. 関係式から個体毎に放流時の体長を推定し(以下,この 体長を推定放流体長という。),推定放流体長 2mm間隔 で再捕尾数を求めた。これらのデータから,群毎および 定,標識確認を行い,92M12~92M15 群と 92M22~ 各群の推定放流体長 2mm間隔の再捕率(再捕尾数/放 92M25 群の群毎の再捕率を算出した。 流尾数×100,)を計算した。 また,これら放流種苗のうち,92M10 シリーズの放 ヨシ帯内での放流サイズ毎の生残率の推定 藤原ら6) 流魚は 92M14 群を放流した 8 日後の 1992 年 6 月 24 日 はヨシ帯におけるニゴロブナの初期生態を知るため,滋 以降に,92M20 シリーズの放流魚は 92M24 群を放流 賀県近江八幡市牧町(以下,牧,Fig. 1)の琵琶湖沿岸 した当日の 7 月 9 日以降に,同ヨシ帯の両端と中央部 に自生するヨシ帯(幅が概ね 150~160 m,奥行きが概 に岸辺から沖合にかけて設置した 3 枚のネットの先端 ね 45~50 m)に,ALC 標識を付けた 10 群のニゴロブ に設置したマス網で捕獲されるようになった。92M11 ナ仔稚魚を放流した。この内,5 群は 1992 年 5 月 17 ~92M14 群と 92M21~92M22 群は,それまでヨシ帯 日にふ化したもので,滋賀水試内で継続飼育しながら異 内部でタモ網で採集されたが,マス網では採集されなか なるパターンの ALC 標識を付けて,2 日齢,10 日齢, ったことから,それぞれ 6 月 23 日以前および 7 月 8 日 20 日齢,30 日齢および 40 日齢の時点で Table 1 に示 以前はそのヨシ帯内に限定して生息していたと考えられ したとおり順次放流した(本報では,それぞれ 92M11 ている(藤原ら6))。そこで,6 月 24 日時点で未放流の 群,92M12 群,92M13 群,92M14 群および92M15 92M15 群を除く 92M10 リーズおよび 7 月 9 日時点で 群。以下,92M10 シリーズ)。また,他の 5 群は同年 6 未放流の 92M25 群を除く 92M20 シリーズの各群の牧 月 15 日にふ化したもので,同様に ALC 標識を付けて 2 のヨシ帯内での生残率を,これらの採集尾数のデータを 日齢,8 日齢,16 日齢,24 日齢,32 日齢の時点で放流 用いて田中14),北田15)にならい,Fig. 2 に示したモデル した(それぞれ92M21 群,92M22 群,92M23 群, にしたがって推定した。ここで Ri は,i 番目に放流した 92M24 群および 92M25 群。以下,92M20 シリーズ)。 92M10 シリーズまたは 92M20 シリーズの種苗の放流 このうち,92M11 群と 92M21 群の標識はともに卵段 尾数,Ei+1 は(i+1)番目に放流した 92M10 シリーズ 階で付けた耳石(礫石)の核部分が点状に発光する標識 または 92M20 シリーズの種苗の放流時点(i+1)にお であるため両者を識別ができないが,他の群の標識はそ ける i 番目の放流群の生残尾数である。ここで i 番目に れぞれ識別できる。そこで,今回の調査では 1992 年 10 放流した種苗と(i+1)番目に放流した種苗の(i+1) 月 15 日から翌年 4 月 6 日にかけて琵琶湖北湖で小型機 時点以降の生残率は等しく,それら種苗のマス網による 船底曳き網を用いて 21,459 尾のニゴロブナ標本を採集 捕獲割合も等しいと仮定して,i 番目に放流した 92M10 し,得られた標本を上記と同様に保存して,後日,測 シリーズまたは 92M20 シリーズの種苗の次の群の放流 ニゴロブナ種苗放流の適地と課題 825 Fig. 2 Schematic diagram of survival estimation of larval and juvenile nigorobuna Carassius auratus grandoculis in the reed zone (Site M in Fig. 1). 時点(i+1)の生残尾数を次式で求めた。なお,ci と た。 n ci+1 は,それぞれ 92M10 シリーズ(6 月 24 日以降)ま ÂS T, n+1=_ Âq i, i+1, (n=2, 3) たは 92M20 シリーズ(7 月 9 日以降)の i 番目と(i+ i=1 1)番目の放流群のマス網で再捕された尾数である。 この分散は,同様にデルタ法によって 2 2 Ri+1ci V ( ÂS T,3)q 23V ( Âq 12)+〈Âq 12V ( Âq 23) ÂE i+1= ci+1 +2 Âq 12 Âq 23 Cov ( Âq 12, Âq 23) これらの生残尾数から,i 番目に放流した 92M10 シ および 2 2 2 2 リーズまたは 92M20 シリーズの種苗の次の群の放流時 V ( ÂS T,4) Âq 23 Âq 34V ( Âq 12)+ Âq 12 Âq 34V ( Âq 23) 〈 2 2 2 点(i+1)の生残率 Âq i, i+1(以下,群間生残率)を次式 + Âq 12 Âq 23V ( Âq 34)〈+2 Âq 12 Âq 23 Âq 34 Cov ( Âq 12, Âq 23) 2 で計算した。 +2 Âq 12 Âq 23 Âq 34 Cov〈( Âq 12, Âq 34) 2 ÂE i+1 Ri+1ci +2 Âq 12 Âq 23 Âq 34 Cov ( Âq 23, Âq 34) Âq i, i+1= = Ri Rici+1 と推定される。ここでは,異なる期間の生残は互いに独 なお,群間生残率の漸近分散はデルタ法(例えば北田16) 立と仮定し,共分散の項を無視して分散を計算した。 を参照)によって一次の項まで展開して次式で推定した。 1992 年 5 月 11 日には,同ヨシ帯の中央において, 2 2 R i+1 V (ci) c i V (c i+1) 岸辺から沖合方向への距離と水深を実測し,その後の水 V ( Âq i, i+1) 2 2 + 4 R i [ c i+1 c i+1 位の変動に伴う水に浸かったヨシ帯(以下,水ヨシ帯) 〈 ci の奥行きの変化を計算した。 -2 3 Cov (ci, ci+1) c i+1 ] 効果的放流サイズの検討 琵琶湖での高い放流効果が ここで,V (ci)=Ripâ i(1-pâ i), pâ i=ci/Ri,〈V (ci+1)=Ri+1 期待できる種苗サイズを検討するため,Fig.
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