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Zeon PDF Driver Trial 中文摘要 綜合1997 年3月至1999年12月期間於高雄及小琉球附近海域 採得樣本的分析結果，共發現浮游性仔稚魚 75科119 屬182 種，其 中以1公尺網行表層拖網部份共計 42科72屬93種；而多層網部份 則發現 41科48屬73種。採集樣本中日本鯷(Engraulis japonicus)為 最優勢的種類，且全年皆有出現的記錄，其它的種類則包括了沿岸 小型魚種(如天竺鯛科及雀鯛科)、沿岸洄游魚種(如鯖科)、大洋性表 層魚種(如鯷科)、大洋中表層性魚種(如燈籠魚科)、河口域仔稚魚種 (如�虎科及�科)及小型底棲性魚種(如沙�科及合齒科)等。而大鵬 灣內僅發現 2科3屬3種，皆為小型的底棲種類。 高雄及小琉球附近海域由於受陸源水及高屏海底峽谷地形的影 響，水文環境較為複雜，仔稚魚相也較為豐富，種類組成及豐度在 季節間的變化較大；相對的，大鵬灣內由於為半封閉的水域環境， 且常年受人為的污染影響，水文環境變化不大，因而仔稚魚的種類 組成較為單純且季節性的變化較不明顯。 整體看來，高雄及小琉球附近海域浮游性仔稚魚的豐度以 6月 份(濕季時)為最高，平均豐度為www.zeon.com.tw108±116 ind./100m3。而2月份時 E. japonicus 在小琉球附近的測站大量出現，推測 1~2月應為該海域 E. japonicus 的繁殖季節，而其他幾種優勢種亦有明顯季節性消長的現 象；另外，在高雄第一港口外以及高屏溪口外的測站 1、4、5、6 仔 稚魚豐度有較高的情形Zeon PDF。而在日夜垂直分布方面 Driver，小琉球附近海域 Trial 浮游性仔稚魚日、夜間的豐度變化並不明顯；除了少數種類以外， 大多數種類日間及夜間的分布皆以 100 米以淺水層為主。 Abstract: There were 75 families, 119 genera and 182 species of ichthyoplankton found in the adjacent areas of Kaohsiung and Liuchiu Yu Island. Fifty families, 82 genera and 111 species were collected in the surface waters, and 37 families, 54 genera and 68 species were collected in oblique tows to 100m with an open 1 m net, while 41 families, 48 genera and 73 species were collected by a multiple opening-closing net. Engraulis japonicus was the most dominant species, and was found all year round; other dominant species included coastal fish species (Apogonidae and Pomacentridae), anadromous species (Scombridae), oceanic species (Engraulidae and Myctophidae), and demersal species (Gobiidae). There were only 3 species of ichthyoplankton belonging to 3 genera and 2 families found in the Tapong Bay, all these species are benthic species. The coastal waters of Kaohsiung and Liuchiu Yu Island might be influenced both by the river and the topography of Kaping Trench, and thus the hydrological conditions of this area were more complex, and the ichthyoplankton was more diverse. On the other hand, the hydrological conditions of the semi-closed Tapong Bay changed less than that of the estuary of Kaoping river, thus the species composition of larval fish was simple and less diverse. Overall, the highestwww.zeon.com.tw abundance of the ichthyoplankton in the adjacent areas of Kaohsiung and Liuchiu Yu Island was found in the wet season(June), with an average of 108± 116 ind./100 m3. E. japonicus was the most abundant in February, revealed that might be the breeding season of this species. Other dominant species (eg. Bregmaceros japonica andZeon Tridentiger sp.) alsoPDF showed significant Driver seasonal variation. Trial Higher abundance of ichthyoplankton was usually found in the entrance of Kaohsiung first harbor(St.1) and the estuary of Kaoping river(ie. St.4~6). No significant diel vertical migration of the ichthyoplankton was observed in this study. Most ichthyoplankton were found in the water column above 100 m during both day and night. 前言: 台灣四面環海，地處亞熱帶地區，沿岸富有多種經濟性魚類(李 等，1985)。但近年來由於魚類資源的過渡開發，已造成某些魚類資 源幾近枯竭。對於魚類資源應如何有效的管理、保育及合理的開發 利用，已成為今後漁業發展之重要課題。為了瞭解魚類資源的動態， 則需先對各種魚類的生活習性及生棲環境有充分的認識。沿岸海域 由於有機質豐富，孕育了豐富的浮游生物，為許多仔稚魚種良好的 孵育場所;其間分佈的仔稚魚數量、種類及變動情形已成為 今後水產資源研究的重要課題(曾等，1985)。 浮游性仔稚魚其實就是發育初期的魚類，分類上屬於脊索動物 門(Phylum Chordata)，有顎上綱(Superclass Gnathostomata)、硬骨魚 綱(Class Osteichthyes)(沈，1984)。至於軟骨魚綱(Class Chondrichthyes) 則因幼生時期不行浮游生活，並不包括在浮游性仔稚魚的範疇之內。 硬骨魚類自卵孵化後大都有一段幼生時期是行浮游性生活，也就是 所謂的魚類仔、稚魚時期；此一時期的魚類因為其個體微 小，器官發育未完全與運動能力不足，大多隨波逐流，賴攝食水中 之微小浮游動、植物為生(李等，1985)。另外，由於仔稚魚缺乏完 整的運動器官，所以其分布深受水團www.zeon.com.tw、海流及其它環境因子的影響； 因此，我們亦可以將浮游性仔稚魚當成偵測海洋水團現象的指標生 物(biological indicators)(Chiu and Wang,1989)。目前全球約有 2萬種 現生種魚類(Okiyama,1988)，而台灣目前已鑑出近 900 種本土性的 仔稚魚Zeon(丘，1999)。 一般而言PDF，魚類的發育過程可以分為八個階段 Driver Trial(曾 等，1985)，而仔稚魚期的型態特徵也會因魚種及不同發育階段而有 所不同，這八個發育階段包括:(1) 卵(Eggs):包括未受精卵及孵化前 的受精卵。(2) 卵黃囊期仔魚(Yolk-sac larvae):魚類孵化後至卵黃囊 消失為止的階段稱之。(3) 脊索末端上屈前仔魚期(Preflexion larvae): 卵黃囊消失至脊索末端開始上屈的階段。(4) 脊索末端上屈中仔魚 期(Flexion larvae):脊索末端開始上屈至上屈完成的階段。(5) 脊索末 端上屈後仔魚期(Postflexion larvae):脊索末端上屈完成至各鰭部發育 完成之階段。(6) 稚魚期(Juvenile):各鰭部發育完成至成魚特徵出現 之階段。(7) 幼魚及未成魚期(Young and Immature):成魚特徵出現， 但是生殖腺並未成熟的階段。(8) 成魚期(Adult):生殖腺完全成熟， 型態特徵不再變化的階段等。其中，由卵黃囊期仔魚至稚魚期的五 個階段，我們稱之為魚類的仔、稚魚時期。 不同種類及生長階段的仔稚魚在形態構造上也會有很大的不同， 這些仔稚魚由於形態構造特異，所以在鑑種的過程中很容易就被鑑定 出來(丘，1999)；它們有的體型特別，如鰻鱺科(Anguillidae)及鯙科 (Muraenidae)的柳葉型幼生、左鰈科(Bothidae)的側扁體型等；有的則 形成特殊的構造及器官，例如大眼鯛科(Priacanthidae)頭部上方的棘 刺、奇棘魚科(Idiacanthidae)延長的眼柄及蝦虎魚科(Gobiidae)大而明 顯的鰾等。有關於仔稚魚研究，主要是以魚類卵黃囊期仔魚至稚魚期 的階段為主要研究範圍，由於卵黃囊期仔魚的種類特徵辨識較為困 難，加上坊間對此一階段仔魚的鑑種圖鑑並不多見www.zeon.com.tw，一般而言，關於 仔稚魚的研究還是以脊索末端上屈前仔魚期至稚魚期為主，此一階段 的仔稚魚不但各微細構造都有專業圖鑑可供辨識，對於許多個別、或 特殊魚種，無論其發育階段或形態特徵，更不乏專業且深入的資料可 供查詢Zeon。 PDF Driver Trial 一般而言，仔稚魚的研究調查工作大多以下列幾種為主：(1)不同 仔稚魚種外部形態，生理構造及發育過程的研究。(2)地區性的仔稚魚 種類分布及豐度調查。(3)特殊魚種行為模式的研究及與環境間相互關 係的研究等。與其他浮游生物一樣，仔稚魚的行為模式也是很複雜的， 它們不但有著不同的生活模式及生棲場所，也有著不同的水平及垂直 遷移情形發生(Leis, 1991; Palomera, 1991)，而這一點又與仔稚魚種、 不同的成長階段以及特殊習性有關(Leis, 1991)。浮游性仔稚魚之所以 表現出複雜的行為模式及改變其分布的水層，可能是因為下列幾種因 素:覓食(Fortier and Leggett, 1984)、躲避掠食者(Gray, 1998)、保持體 內能量(Gray, 1998)、對環境的生理需求(Gray, 1998)、規則化的自發性 運動(Fortier and Leggett, 1982)等。以往的報告指出，全球各地不同種 類仔稚魚的垂直及水平分布都與該處的水文環境條件有著密切的關係 ( Loeb, 1980; Hamann et al., 1981; Palomera, 1991)；Tzeng et al.(1997) 在台灣東北部鹽寮灣海域進行仔稚魚調查研究亦發現，近岸水域仔稚 魚的發生、豐度及種類組成常常會依不同的產卵季節與環境因子而有 所改變。 在國外，有關仔稚魚及魚類初期生活史的調查研究可謂不勝枚 舉，然而在我國卻較為少見(曾等，1985)。在台灣仔稚魚的研究工作 則發展的較晚，最早是由台灣大學海洋研究所在 1970 年對中國南海 進行的仔稚魚初步調查(Hu, 1974)。而早期仔稚魚的研究多以分類及 型態描述作為探討主題，例如胡等www.zeon.com.tw(1970)、Chen and Tan(1973)對台灣 鰹鮪類仔稚魚所作的研究報告，其中完整地記錄了台灣附近海域鰹鮪 類仔稚魚的形態特徵及鑑種依據；其它魚種的研究還有烏魚(Tung, 1973)及飛魚類(Chen, 1978)等。直到最近的二十年，才有較多的學者 投入有關於仔稚魚生態及分布方面的研究Zeon PDF Driver，其中又以行政院農委會於 Trial 民國七十二年結合全國各級學術研究單位，為期三年的台灣沿岸海域 仔稚魚苗研究調查工作，將台灣北部、東部、西部及南部沿岸及河口 區的仔稚魚的種類組成及分布做了初步較完整的調查，為台灣本土仔 稚魚研究工作推動之先驅(袁，1985)。台灣在仔稚魚方面的研究可以 說才剛剛起步，研究方向還是以地方性的仔稚魚資源調查為主，其中 又以台灣北部及東部的研究較為普遍，例如 Chiu(1993)、劉(1985)、 曾等(1985)、莊等(1985)對台灣北部各河口及沿岸海域仔稚魚做的調 查以及 Yeh(1993)、Tzeng(1989)等對東部黑潮水域仔稚魚的調查報告 等；至於台灣西部以及南部海域的仔稚魚研究除了陳(1985)對高屏及 濁水溪附近仔稚魚之出現做過較深入的調查研究之外，其它則較為少 見。 本研究的目的，是利用不同的實驗設計及採樣方式於高雄及小琉 球一帶海域進行採樣，以瞭解台灣西南沿近海域浮游性仔稚魚的種類 組成、季節性分布及日、夜垂直遷移的情形，並探討其與地緣環境因 子間可能的相互關係，俾便對該海域仔稚魚種類組成、分布及漁業資 源評估有更進一步的瞭解及貢獻。 材料與方法: 一、採樣時間、地點及方法 本研究因不同的實驗需要及目的而有不同的採樣點及採樣方式www.zeon.com.tw ，分別敘述如下: (1) 仔稚魚在高雄及小琉球海域測站間及季節性的分布: 本研究是利用海研三號於 1997 年3月22日、6月7日、9月8 日及Zeon12月26日等4 個航次PDF，在高雄及小琉球附近海域的 Driver 6Trial個測站(圖 1)進行日間表水層(0~5 公尺)之水平拖網，使用的網具為 1公尺網(One meter net,網口直徑一公尺，網身長度 4.5公尺，網目大小為 330μm)。 採樣時在網口處加裝流量計(flow meter)以估算流經網口之水量，拖 網時船速維持在 2節左右(約1m/sec)，每次拖網時間為 10分鐘。採 得的樣品在船上直接以 5~10 %的中性福馬林(pH=7.4~7.6)溶液固 定，裝入 1000ml 的樣本瓶中保存並記錄基本資料如採樣日期、測站 名及流量計讀數等，以便攜回實驗室做進一步的鑑種、計數與資料 分析。 另外，為了比較在 100 公尺以淺水層與表水層中仔稚魚之種類 組成及豐度變化的情形，我們在 1997 年9月8日及12月26日的兩 個航次當中，於離岸的測站 2、3、4、5與表層拖網同步進行 100 公 尺斜拖的採樣工作，採樣網具仍然使用 1公尺網，而拖網方式改為’V’ 型拖，每次採樣時間也是 10分鐘。 (2) 仔稚魚之日夜垂直分布: 本研究是由海研三號在 1997 年5月2~3 日、7月7~8 日、10月 17~18 日、12月26~27 日及1998 年2月20~21 日的航次，在第 4測 站(如圖1)以多層式浮游動物網(HydroBios multiple nets, 網框為 0.5 ×0.5 公尺的四方形，網目大小 330μm)進行日間(12:00AM~3:00PM) 及夜間(10:00PM~1:00AM)的分層採樣;採樣深度分別為表水層(0~5 公尺)、50公尺、100 公尺、150 公尺及 200 公尺。採樣時在每個網 口加裝流量計以估算濾過之水體積www.zeon.com.tw，每一水層拖網時間為 25~30 分 鐘。在5月2日的採樣過程中，因操作疏失不慎將 150m 深的網具 流失，因而缺乏該水層之樣品。另外，7月份採樣時因為多層網網 具故障，故臨時以北太平洋標準網(NorPac net,網口直徑 0.45 公尺， 網長Zeon1.8 公尺，網目大小為 PDF330μm) 加裝閉鎖器來Driver代替原本的採樣網 Trial 具。採樣深度與之前一樣，仍為水表層、50、100、150 以及200 公 尺，每一水層的拖網時間為 10分鐘。 (3) 大鵬灣內及高屏溪口附近海域乾、溼兩季仔稚魚種之分布: 為了對台灣西南沿海仔稚魚種類組成與分布有更進一步的了 解，我們亦利用海研三號 1999 年6月24日及12月24日的航次， 於高屏海域另外7個測站(圖2)以1公尺網(One meter net,網目大小330 μm)進行仔稚魚的水平拖網採樣，每次拖網的時間為 10分鐘，網口 並加裝流量計以估算濾過之水體積。另外，亦配合海研三號航次， 同日於大鵬灣內 3個測站(圖2)租用塑膠管筏進行樣本採集工作，因 受限於水域較為狹窄且水淺，而改以北太平洋標準網(NorPac net,網 口直徑 0.45 公尺，網長 1.8 公尺，網目大小為 330μm)進行水表層(0~1 米)之拖網採樣，拖網時間為 5分鐘。 二、海水水文資料的收集 在每一測站的採樣前或採樣後，均利用海研三號上的溫鹽深儀 (SBE 9/11 CTD; SeaBird Electronics Inc., Washington D.C., USA)進行 水文環境參數資料的偵測及收集，所收集的資料包括不同深度水體 之溫度、鹽度、螢光值、透光度及 pH值，這些資料經過電腦軟體 (Seasoft V4.202; SeaBird Electronics Inc., Washington D.C., USA)的轉 換處理後，用來進行水文環境之分析；另外，於採樣的同時亦隨時 記錄當時的海況及天候狀況等相關資料www.zeon.com.tw。而大鵬灣內每一測站採樣 前則以自攜式溫鹽儀測量表水的溫鹽資料。 三、樣本的處理 Zeon採得的樣品攜回實驗室後 PDF，先於解剖顯微鏡 Driver(Askmnia; Trial SLG Chemnitz, Co., Germany)下進行鏡撿的工作，因為樣品中仔稚魚的數 量有時會很少，所以除非有必要，盡量以全部的樣品直接進行分析。 若樣品中仔稚魚量太多，則視樣品的多寡先以分樣器(Folsom plankton splitter)分為二分之一至四分之一不等再行鏡檢。 所有樣品中的仔稚魚挑出後，改以 75 %的酒精浸泡保存，並儘 速進行鑑種及計數工作。仔稚魚鑑種的工作則以較高倍率及高解析 度之解剖顯微鏡下進行;種類的鑑定方面主要是依據多篇圖鑑以及 Okiyama (1988)、丘(1999)、曾等(1985)、陳(1985)、莊等(1985)、黃 (1985)、Leis(1983、1989)等人所描述之特徵以及屏東科技大學陳正 修老師、中央研究院王友慈老師的協助及指導。仔稚魚的鑑種方式 主要是依照其外部特徵，這些外部特徵包括下列幾種(Okiyama,1979 ；曾等，1985；丘，1999): 1.體型: 呈長條型、側扁、縱扁或柳葉型、球形等。 2.體型比例: 體高/體長的比例。 3.頭部比例: 頭部/體長的比例。 4.肛門位置: 肛門位於魚體中央偏前或偏後，或是位於第幾肌節處。 5.鰭部特徵: 各鰭部形狀、鰭條數、背鰭數目、是否有脂鰭等。 6.眼部形狀: 圓形或是橢圓形、不規則；是否有眼柄等。 7.鰓蓋棘與眼眶上棘: 是否有鰓蓋棘、眼眶上棘以及數目、形狀等。 8.體表突起物: 頭部有無棘狀突起物、有無鬚及前、後隅角下突起。 9.腸道形式: 腸道平直或曲折www.zeon.com.tw，以及曲折形式等。 10.色素胞形態特徵: 體表及肌肉是否有色素胞分布，分布位置、形 態、數目及色澤。 11.肌節數: 計算各階段仔稚魚肌節的數目來鑑別種類。 12.Zeon其他特殊構造: 如發光器PDF(例:某些燈籠科魚種 Driver)；體表具硬質骨 Trial 板狀構造(例:飛海蛾)等。 初學鑑種時依照仔稚魚不同的形態特徵，將其拍照或利用解剖 顯微鏡將特徵描繪下來，並詳細記錄其微細構造如肌節數、色素胞 位置及鰭條數等，以期對仔稚魚能有更進一步的了解。待各部特徵 的辨識較熟練後，再於解剖顯微鏡下正式進行樣品的鑑種工作，鑑 種時將採得仔稚魚樣本盡量鑑別至最低的分類階層。 四、資料處理及分析 為瞭解浮游性仔稚魚類群聚內之種類豐富程度(Species richeness) 及個體數在種間分配是否均勻，本研究以 Shannon-Weaver 之種歧異 度指數(Index of species diversity, H’)來進行估算分析，其公式如下: n H’ = -Σ(Pi)(log2 Pi) i=1 Pi : 第i種生物之個體數在總個體數(所有種類之個體數總和)的 比值。 為了進一步瞭解仔稚魚群聚組成及數量在不同採樣測站及季節 間的變異程度；亦利用主成份分析法(Principal Component Analysis, PCA)進行分析，再將分析所得之前兩個主成份軸之轉載值(Loading score)以群聚分析法(Cluster analysis, CA)繪製成樹狀圖來加 以探討。另外，亦以 Bray-Curtis Similarity(PRIMER; Clarke and Warwick, 1994)的分析方法來探討仔稚魚之種群及其分布特徵www.zeon.com.tw 。 為暸解仔稚魚群聚豐度在深度、季節、日夜以及測站間的差異 情形，將採樣測站劃分為近、遠岸及高屏溪口、高雄港外兩條測線， 利用變方分析統計法(ANOVA)來測試其是否有顯著的差異存在；再 以ZeonDuncan’s多變距分析法加以分析比較 PDF Driver；另外，亦以線性回歸分析 Trial 來探討仔稚魚豐度與環境因子的相關性，以期能瞭解影響 該海域浮游性仔稚魚時空分布的可能原因。 結果: 本研究的結果依不同的研究目的及採樣方式分為四個部份加以 論述，分別為水文環境因子的探討、仔稚魚群聚在測站間及季節性 的變化、仔稚魚之日夜垂直分布以及大鵬灣內外海域於乾、濕兩季 仔稚魚之種類組成及分布等。分別敘述如下: (ㄧ)水文環境因子的探討 由1997 年3月至12月期間於高雄及小琉球附近六個測站(圖1) 測得水表層的溫鹽度結果(表1)得知，海域表水溫度以 9月時最高(平 均溫度 28.7±0.30C)；12月時最低(平均溫度 24.2±0.40C)。至於測站 間的水溫差異則以
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