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  • 學位論文

帶魚總科(真骨魚類:鯖形目)之分類、親緣關係及身體延長與偶鰭退化之間的演化相關性之研究

Taxonomy, phylogeny and evolutionary correlation between body elongation and paired fin reduction in Trichiuroidea (Teleostei: Scombriformes)

指導教授 : 陳韋仁

摘要


帶魚總科(Trichiuroidea)為真骨下綱(Teleostei)棘鰭類(Acanthomorpha)鯖形目(Scombriformes)之魚類,由帶魚科(Trichiuridae)和帶鰆科(Gempylidae)兩科組成,目前全球共包含26個屬及72個種。該類群為水層或底層習性之海水魚類,身體形狀呈現由梭形至皮帶狀之一系列分布。生態與漁業研究將具有資源管理與保育的重要性,其需要堅實的分類學基礎。然而,漁業的重要種類,例如:帶魚屬之物種(Trichiurus spp.),其親緣關係假說、生物分類、物種之多樣性以及型態鑑定至今仍存在著爭議。此研究使用五個核基因序列,為帶魚總科重建分子親緣關係,解決長期存在的問題,包括帶魚總科在鯖形目此一單系群之中的親緣關係位置、帶魚總科之內的親緣關係,以及各屬是否為單系群。本研究結果支持帶魚總科和其他五個科組成一單系群(包括長鰭金眼鯛科[Caristiidae]、長鰭帶鰆科[Scombrolabracidae]、烏魴科[Bramidae]、圓鯧科[Nomeidae]和鯖科[Scombridae])。帶鰆科之中的鱗網帶鰆屬(Lepidocybium)和單系群的帶魚科形成姊妹群。帶魚總科的大多數的屬被確認為單系群。帶魚屬則包括了沙帶魚屬而為一併系群。這些發現提供了帶魚總科未來分類更動之依據。為了檢驗物種假說並探索物種之多樣性,我使用包含了687個粒線體COI基因序列之資料矩陣(共包含176個新序列,與511個取自公開資料庫的序列),並以此利用整合性的分類方法來進行物種界定。總共有56個假定種被界定,並揭露了多個屬之中被低估之多樣性,包括紫金魚屬(Promethichthys)、短帶鰆屬(Rexea)、無耙帶鰆屬(Nesiarchus)、隆頭帶魚屬(Tentoriceps),以及帶魚屬。針對帶魚屬的物種,我統整物種界定的結果和深入的全魚型態檢視結果,建立有效的檢索表,以鑑定西北太平洋和熱帶印太平洋區域之中已知、已被發現的物種。 此研究也利用建立的親緣關係系統,檢驗帶魚總科是否出現身體延長和附肢退化(胸鰭和腹鰭退化)之間的相關性,此著名現象曾在陸地四足類和底棲魚類上被發現過。該現象長期被認為是適應挖掘或縫隙躲藏的演化結果,而這兩個因素都可在行水層性或底層性生活的帶魚總科上被排除。親緣關係迴歸比較分析方法(PICs, PGLS, PGLMM)顯示身體延長程度和胸鰭/腹鰭的長度之間呈現負的演化相關性。為了找到可能導致此相關性的因子,我探討了運動方式和生態可能造成的影響。

並列摘要


Trichiuroidea is a superfamily of acanthomorph fishes classified under the Scombriformes (Teleostei). A total of 72 species in 26 genera are currently recognized in this superfamily which comprises cutlassfishes from Trichiuridae and snake mackerels from Gempylidae. They are marine pelagic or demersal fishes, exhibiting a spectrum of body shape from fusiform to elongated form. While prospective work on ecology and fisheries of this group of fishes is important for resource management and conservation, this requires a solid taxonomic basis. However, controversies still remain for phylogenetic hypothesis, taxonomic status, species-level diversity and morphological diagnosis of trichiuroids, even for species of high interest in fisheries (e.g., Trichiurus spp.). In this study, a comprehensive multi-nuclear gene phylogeny of the Trichiuroidea was newly reconstructed to firstly answer long-lasting questions regarding the placement of the Trichiuroidea within the monophyletic Scombriformes sensu Nelson et al. (2016), relationships within the superfamily and the monophyly of currently recognized genera. My phylogenetic results supported monophyly of Trichiuroidea and five of the examined families (Caristiidae, Scombrolabracidae, Bramidae, Nomeidae and Scombridae) expect Gempylidae. One of gempylid genus (Lepidocybium) was found to be the sister group of the Trichiuridae. Within the Trichiuroidea, most of the included genera were confirmed to be monophyletic. Trichiurus was found to be paraphyletic with respect to Lepturacanthus. These findings provided a basis for future revisions for the classification of trichiuroid taxa. To further test the species hypothesis and to explore the species-level diversity, a mitochondrial COI gene dataset with 687 samples (176 newly obtained; 511 retrieved from public sources) was constructed for the species delimitation analyses through an integrated approach in taxonomy. A total of 56 putative species were finally determined. In several genera such as Promethichthys, Rexea, Nesiarchus, Tentoriceps and Trichiurus, hidden diversity was detected. For Trichiurus species, by integrating the results of species delimitation analyses and a close morphological examination on the voucher specimens, I herein established an effective identification key to the known and discovered Trichiurus species occurring in the tropical Indo-West Pacific and Northwest Pacific. With the established phylogenetic framework of the Trichiuroidea, this study also tested the hypothesis of evolutionary correlation between body elongation and reduction of appendages (i.e., pectoral and pelvic fins in fishes), a famous phenomenon discovered on terrestrial tetrapods and benthic fishes. This phenomenon has long been considered as a consequence of fossorial or crevice-sheltering adaptation in which the conditions are ruled out for the pelagic and demersal fishes like trichiuroids. The results from the analyses of phylogenetic regression-based comparative methods (PICs, PGLS, PGLMM) for the trichiuroids showed a negative evolutionary correlation between body elongation levels and lengths of pectoral/pelvic fins. To understand potential causes of this correlation, aspects of locomotion and ecology were discussed.

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