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研究生:王大明
研究生(外文):John Ta-Ming Wamg
論文名稱:燈籠魚類系統分類之研究
論文名稱(外文):Systematic Studies on the Lanternfishes (Myctophiformes)
指導教授:陳哲聰陳哲聰引用關係莫顯蕎莫顯蕎引用關係
指導教授(外文):Che-Tsung ChenHin-Kiu Mok
學位類別:博士
校院名稱:國立海洋大學
系所名稱:漁業科學學系
學門:農業科學學門
學類:漁業學類
論文種類:學術論文
論文出版年:2001
畢業學年度:90
語文別:中文
論文頁數:169
中文關鍵詞:燈籠魚燈籠魚科新燈籠魚科燈籠魚目分佈類緣關係
外文關鍵詞:LanternfishesMyctophidaeNeoscopelidaeMyctophiformesDistributionPhylogenetic relationship
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  • 被引用被引用:6
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燈籠魚類全世界共有2科36屬約250種,直到1996年為止,台灣周邊海域共記載有2科6屬17種。為進一步了解台灣及東沙島周邊海域之燈籠魚(燈籠魚科及新燈籠魚科)種類及其分佈情形,以及燈籠魚目魚類之類緣關係,本研究乃於1991至1997年,利用台灣水產試驗所水試一號進行底拖網及IKMT(Issac-Kidd Midwater Trawl)之實驗調查,蒐集燈籠魚標本,作業範圍為北緯19至25度,東經114至123度之海域內,水深自50至1521公尺,共採集燈籠魚標本1622尾。經詳細鑑定得知共有16屬53種,其中11屬23種在該海域為新記錄種。新記錄種之燈籠魚為帶底燈魚(Benthosema fibulatum)、側上虹燈魚(Bolinichthys supralateralis)、燦爛眶燈魚 (Diaphus fulgens)、符氏眶燈魚 (D. fragilis)、黑潮眶燈魚 (D. kuroshio)、莫名眶燈魚 (D. problematicus)、華麗眶燈魚 (D. perspicillatus)、菲利眶燈魚(D. phillipsi)、高體電燈魚(Electrona risso)、近壯燈魚(Hygophum proximum)、萊氏壯燈魚(H. reinhardtii)、糙炬燈魚(Lampadena anomala)、日本葉燈魚(Lobianchia gemellarii)、黑體珍燈魚(Lampanyctus niger)、圖氏珍燈魚(L. turneri)、天紐珍燈魚(L. tenuiformis)、暗色燈籠魚(Myctophum asperum)、金焰燈籠魚(M. aurolaternatum)、閃光燈籠魚(M. nitidulum)、櫛棘燈籠魚(M. spinosum)、尖吻背燈魚(Notolychnus valdiviae)、尾棘背燈魚(Notoscopelus caudispinosus) 及 閃光背燈魚(N. resplendens)。其中側上虹燈魚(B. supralateralis)、高體電燈魚(E. risso)、圖氏珍燈魚(L. turneri)、糙炬燈魚(L. anomala)、尖吻背燈魚(N. valdiviae)及尾棘背燈魚(N. caudispinosus) 6種在南海也是新記錄種;而糙炬燈魚(L. anomala)更是全亞洲海域的新記錄種
在台灣東部、台灣西南及東沙島周邊海域中,各海域間之燈籠魚魚種組成,其相似度很接近。台灣東部及台灣西南兩海域間,其相似度指數為0.78。台灣西南及東沙島周邊兩海域為0.76。台灣東部及東沙島周邊兩海域,則為0.70稍低於前兩者。
在分析屬間類緣關係時,所採用的特徵形質有外部特徵與內部特徵兩大類。外部特徵主要有:牙齒型式、主上頜骨形狀、嗅囊大小、各鰭之相對位置、背鰭基與臀鰭基之大小、鰭條和棘之數目、發光器和發光腺之類型等。內部特徵主要有:脊椎骨數目、鰓耙數目、以及柔軟器官(soft organs)等。柔軟器官包括有:鰾之奇網和卵圓體之型式、胃之型式、幽門垂數目及腦之相對大小等。而上述柔軟器官特徵,乃首次被使用來闡明燈籠魚類屬間之類緣關係。
在新燈籠魚科中,其資料矩陣,乃以3屬4種和15個特徵所建立。以仙女魚科之仙女魚(Aulops japonicus)為外群,並用MacClade 4.0分析後,得到新燈籠魚屬)與軟犁燈籠魚屬,因同具翼骨齒之離徵,而類緣關係較近。
在燈籠魚科中,其資料矩陣,乃以33屬129種和56個特徵所建立。以新燈籠魚科之擬燈籠魚屬為外群,並用MacClade 4.0以無序原則(unordered) 分析後,得到真蜥燈魚屬及2個單系群。而此2個單系群有10個共同離徵。
在珍燈魚亞科中,有星燈魚族、裸燈魚族及珍燈魚族。由於同具3個離徵,本研究將其歸在珍燈魚亞科中。星燈魚族包括泰勒燈魚屬、克燈魚屬、羅燈魚屬、錦燈魚屬及星燈魚屬5屬。其中之克燈魚屬,因其和羅燈魚屬、錦燈魚屬及星燈魚屬,共同具有4個離徵,本研究將之歸在星燈魚族中。
珍燈魚屬和短鰭珍燈魚屬曾有許多學者將兩者同視為珍燈魚屬。在本研究中,因皆同具5個離徵,而在同一分支(類群)中;但亦因彼此各有6個不同的離徵,所以,兩者皆為有效的屬。
眶燈魚屬是燈籠魚科中種類最多的屬,本研究用46個種及41個特徵,經MacClade 4.0 以無序原則分析後,可分為兩個類群。其中一類群具So之離徵;而另一類群則具主上頜骨長度稍長,而在眼眶之後方的離徵。
燈籠魚類的鰾與發光器之構造上,是很獨特的。例如,燈籠魚類皆為單極型式的奇網,奇網是從鰾的前方或下前方進入鰾內,且有卵圓體。而巨口魚目其鰾之奇網乃是從鰾之後方進入鰾內,其奇網為雙極的型式。此外,新燈籠魚科魚類之奇網為五束,而所有燈籠魚科魚類之奇網為三束。故以鰾的形質特徵言,燈籠魚是相當獨特的。在發光器的部分,燈籠魚科發光器上有鱗片覆蓋,其鱗片有聚光之作用。而新燈籠魚科則無鱗片覆蓋在發光器上,而且,其發光器為不完整之圓形,不似燈籠魚為一完整之圓形。此外,光魚科魚類(Phosichthyidae)之發光器,則極易脫落,而與燈籠魚類不同。因此,許多的證據支持燈籠魚類為單系。
由以上可知,在台灣及東沙島周邊水域,共發現燈籠魚 2科16屬53種,其中有23種為新記錄。並且建立了該水域中,所有燈籠魚之簡明檢索表。此外對於全世界燈籠魚目之2科36屬,本研究亦建立各屬之簡明檢索表。在類緣關係中,燈籠魚科內除可分為,燈籠魚亞科及珍燈魚亞科兩類群外,本研究亦提出燈籠魚科內,屬間類緣關係之論說。而燈籠魚科內之珍燈魚屬及短鰭珍燈魚屬,兩者皆為獨立有效的屬。在眶燈魚屬內,該屬可分為兩個單系類群。最後,本研究所分析的特徵中,許多證據皆支持燈籠魚類為一單系群。

Lanternfishes which include the Myctophidae and Neoscopelidae are ubiquitous and speciose, with approximately 250 species in 36 genera. Until 1996, only 17 species in 6 genera had been recorded in the waters around Taiwan. This study aims to better elucidate the distribution of lanternfish around Taiwan and the Tungsha Islands, and to resolve the phylogenetic relationships of the genera in their 2 families. The area sampled for lanternfishes lies between 19°N and 25°N and 114°E and 123°E; lanternfish specimens were collected during 9 cruises from 1991 to 1997. They comprised 53 species belonging to 16 genera. Among these, 23 species are new to this area. These newly recorded species include: Benthosema fibulatum, Bolinichthys supralateralis, Diaphus fulgens, D. fragilis, D. kuroshio, D. problematicus, D. perspicillatus, D. phillipsi, Electrona risso, Hygophum proximum, H. reinhardtii, Lampadena anomala, Lobianchia gemellarii, Lampanyctus niger, L. turneri, L. tenuiformis, Myctophum asperum, M. aurolaternatum, M. nitidulum, M. spinosum, Notolychnus valdiviae, Notoscopelus caudispinosus, and N. resplendens. Among these, 6 species, including Bolinichthys supralateralis, Electrona risso, Lampanyctus turneri, Lampadena anomala, Notolychnus valdiviae, and Notoscopelus caudispinosus, are first records for the South China Sea, and the species, Lampadena anomala is a new record for Asian oceans.
The similarity indexes (SI) of lanternfish composition among the 3 areas were very close to one another. The SI between the southwestern and eastern waters of Taiwan was 0.78; it was 0.76 between southwestern waters and the Tungsha Islands. The SI between the Tungsha Islands and eastern Taiwan was 0.70, whereas the SI (0.78) between southwestern and eastern Taiwan showed the highest similarity among these 3 regions; this may be caused by the Kuroshio Current which passes along the east coast of Taiwan and its branch which passes off the southwestern coast.
The characters used in this study were composed of both external and internal characters. The external characters included mainly those from teeth, maxillary shape, olfactory shape, numbers of procurrent rays or spines, location of fins, comparison between dorsal and anal fin bases, and patterns of photophores and luminous tissue. The internal characters were comprised mainly of the vertebrae number, number of gill rakers, and information on the soft organs from the following parts: types of rete mirabile and oval in the swimbladder, shape of the stomach, number of pyloric caeca, and relative size of the brain. Many soft organ characters mentioned above help resolve the interrelationships of most myctophid genera.
In the Neoscopelidae, the data matrix is composed of 15 characters of 4 species (in 3 genera) with Aulops japonicus (Family: Aulopidae) as the outgroup. After the analysis using MacClade 4.0, by sharing the same apomorphic character of pterygoid teeth, the genus Neoscopelus was determined to be closer to the genus Solivomer than to the genus Scopelengys.
In the Myctophidae, the data matrix was composed of 56 characters of 129 taxa (in 33 genera). Scopelengys (Family: Neoscopelidae) was the outgroup. An analysis shows that Scopelopsis is the sister group of a large clade formed by 2 main monophyletic subgroups (the Myctophinae and Lampanyctinae). The latter clade is supported by 10 synapomorphic characters.
Three tribes, the Gonichthyini, Gymnoscopelini, and Lampanyctini, are in the subfamily Lampanyctinae. However, Paxton and Stiassny put the tribe Gonichthyini in the Myctophinae. The Gonichthyini was placed in the Lampanyctinae in the present study, because it shares 3 apomorphic characters with other lampanyctines. The Gonichthyini includes Loweina, Tarletonbeania, Gonichthys, Centrobranchus, and Krefftichthys. The genus Krefftichthys is now placed in the tribe Gonichthyini, which was allocated to the subfamily Myctophinae by Paxton and Stiassny. My treatment was based on the reasoning that this species shares the same apomorphic characters (e.g., olfactory organ almost equal with eye lens, pyloric caeca very close to cardiac region, the number of precaudal vertebrae less than 12, and 2 optic lobes parallel) with other Gonichthyini members (Loweina, Gonichthys, and Centrobranchus).
There are many polytomous states in the phylogenetic trees of Paxton's and Stiassny’s phylogenetic proposals which were derived from osteological and muscular characters. Such a polytomous phenomenon means that the relationships among genera of the Myctophidae remain unsettled. Unresolved interrelationships remain among genera of the Myctophidae in Paxton's analysis, and more polytomous states occurred in Stiassny's tree such as: Hygophum, Myctophum, and Metelectrona; Hintonia, Gymnoscopelus, and Lampichthys; Lampanyctodes, Idiolychnus, Lobianchia, and Diaphus; and Bolinichthys, Triphoturus, Stenobrachius, and Parvilux.
Because of sharing 5 apomorphic characters, the genera Lampanyctus and Naanobrachium are in 1 clade. Despite many scientists placing Naanobrachium in the genus Lampanyctus, synapomorphic characters for each of these 2 genera support their having valid generic status.
The genus Diaphus, the most speciose genus of the Myctophidae, can be divided into 2 groups by analysis of 41 characters in 46 taxa through MacClade 4.0. Among 46 species examined, 1 group possesses the apomorphic character of the photophore So, while the other group possesses the apomorphic character of the maxillary exceeding the eye orbital.
Much evidence supports lanternfish monophyly, such as structures of the swimbladder and photophores. The “unipolar” rete mirabile of the swimbladder in lanternfishes greatly differs from the “bipolar” rete mirabile of the swimbladder in Stomiiformes. The 3-strand unipolar rete mirabile of the swimbladder in the Myctophidae and the 5-strand unipolar rete mirabile of the swimbladder in the Neoscopelidae represent derived character states as the bipolar rete mirabile of the swimbladder in the Stomiiformes. Ovals of swimbladders are present in lanternfishes but are absent from the Stomiiformes. Photophores in the Myctophidae are covered by scales, but no scales cover the photophores in the Neoscopelidae. Furthermore, the “open-circle”shaped photophores in the Neoscopelidae differ from the “closed-circle”-type found only in the Myctophidae; however many differences exist with the Stomiiformes such as deciduous photophores in the family Phosichthyidae.
Fifty-three species of 16 genera in the families Myctophidae and Neoscopelidae were collected from waters surrounding Taiwan, including the Tungsha Islands. Among these, 23 species are new records to these areas. A key to the species is provided. And a key also to 36 genera, recorded from all over the world, in the 2 families is established in the present study. In phylogenetic relationships, the Myctophidae is comprised of 2 subfamilies: the Myctophinae and Lampanyctinae. We hypothesize about the generic interrelationships among the Myctophidae and Neoscopelidae. And Lampanyctus and Naanobrachium are valid genera. The genus Diaphus can be divided into 2 subgroups. Characters analyzed in this study provide additional evidence to support the monophyly of the Myctophiformes.

目 錄
壹、中文摘要--------------------------------------------------- 2
貳、英文摘要------------------------------------------------------ 5
參、謝辭 ---------------------------------------------------------- 9
肆、前言 ----------------------------------------------------------- 10
伍、材料與方法 -------------------------------------------------- 14
陸、結果 ---------------------------------------------------------- 21
一、分類體系及新記錄種之描述 ---------------------------- 21
1. 全球燈籠魚目及屬之檢索表---------------------------- 21
2. 台灣及東沙島周邊海域燈籠魚之新記錄種之描述
並附本省產種之檢索表 ---------------------------------- 25
3. 台灣及東沙島周邊海域燈籠魚之分佈概況 ---------- 49
二、燈籠魚類緣關係 ------------------------------------------ 52
柒、討論 ---------------------------------------------------------- 55
捌、引用文獻 ----------------------------------------------------- 68
玖、附表 ---------------------------------------------------------- 75
拾、附圖 ---------------------------------------------------------- 85
拾壹、附錄-------------------------------------------------------- 139
1. 燈籠魚科資料矩陣---------------------------------- 139
2. 眶燈籠魚屬之資料矩陣----------------------------- 148
3. 新燈籠魚科之資料矩陣----------------------------- 150
4. 形質特徵說明--------------------------------------- 151
5. 標明特徵之類緣關圖------------------------------- 157
6. 燈籠魚科刪除發光器特徵之類緣關圖------------- 163
7. 燈籠魚科僅用發光器特徵之類緣關圖------------- 165
8. 燈籠魚科所有特徵以有序原則之類緣關圖------- 167
9. 眶燈籠魚屬所有特徵以有序原則之類緣關圖---- 168
10. 附Paxton及Stiassny之類緣關圖---------------- 169

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