2013年7月13日 星期六

瘤蛾科主要支系間之親緣關係及台灣產物種分類歸屬之更新

文獻來源:Zahiri, R., J. D. Lafontaine, J. D. Holloway, I. J. Kitching, B. C. Schmidt, L. Kalia and N. Wahlberg. 2012. Major lineages of Nolidae (Lepidoptera, Noctuoidea) elucidated by molecular phylogenetics. Cladistics 1(2012): 1-23. [Abstract]

To elucidate the evolutionary relationships of the major lineages within the moth family Nolidae, we analysed a molecular dataset comprising eight independent gene regions (6.4 kbp), cytochrome c oxidase subunit I (COI) from the mitochondrial genome, and elongation factor-1α (EF-1α), ribosomal protein S5 (RpS5),carbamoylphosphate synthase domain protein (CAD), cytosolic malate dehydrogenase (MDH), glyceraldehyde-3-phosphate dehydrogenase (GAPDH), isocitrate dehydrogenase (IDH) and wingless genes from the nuclear genome, using parsimony and model-based evolutionary methods (maximum likelihood and Bayesian inference). Our analyses revealed a well-resolved phylogenetic hypothesis, again recovering the six previously recognized families within Noctuoidea (i.e. Oenosandridae, Notodontidae, Euteliidae, Erebidae, Nolidae and Noctuidae), and monophyly of the quadrifid Noctuoidea (i.e. Euteliidae, Erebidae, Nolidae and Noctuidae). The family Nolidae is diagnosed and characterized by two synapomorphies from morphology: construction of a ridged boat-shaped cocoon that bears a vertical exit slit at one end; and two other morphological character states: elongation of the forewing retinaculum into a bar-like or digitate condition and possession of a postpiracular counter-tympanal hood. We present a new phylogenetic hypothesis for Nolidae consisting of eight strongly supported subfamilies, two of which are erected here: Diphtherinae, Risobinae, Collomeninae subfam. nov., Beaninae subfam. nov., Eligminae, Westermanniinae, Nolinae and Chloephorinae. Where we are able, each monophyletic lineage is diagnosed by morphological autapomorphies and within each subfamily, monophyletic tribes and subtribes are circumscribed, most of which are also diagnosable by morphological apomorphies. We also describe two new taxa: Gelastocerini trib. nov. and Etannina subtrib. nov. The Neotropical subfamily Diphtherinae, here newly circumscribed, is considered to be the plesiomorphic sister lineage to the rest of Nolidae. Diphtherinae are characterized by loss of the proximal pair of metatibial spurs in males and by the presence of a frontal tubercle, which is presumably associated with a derived strategy of emergence from the cocoon.

接續2011年與2012年分別針對夜蛾總科、裳蛾科之內部分子親緣關係的研究,芬蘭土庫(Turku)大學博士生Zahiri與其共同研究者在這篇文章中專注於瘤蛾科進行分子親緣關係的釐清與形態特徵再檢討。
瘤蛾為夜蛾總科六個主要支系之一,全世界已知約有1861740種被描述,主要分布於舊熱帶區系,因部份類群(例如赭瘤蛾屬 Carea)之幼蟲胸部背方隆起而有此中文名之稱。此類群當中被視為主要之農業害蟲為鑽瘤蛾屬(Earias),其於印澳區與非洲區的物種常被視為棉花作物之柱莖性害蟲。
瘤蛾因包含了不少形態異質性高的類群,因此在過去的分類位階一直不明確,而部份成員特徵(例如聽器結構、前後翅翅脈相、幼蟲毛列序)介於夜蛾科與燈蛾科(現今之裳蛾科燈蛾亞科),亦散置於該些分類位階之下。雖然由Holloway (2003)為主的文章開始重新驗證了此科級的單系性,然而內部親緣關係尚未確認。
作者在材料上主要選取82個瘤蛾科各亞科、亞科下分類單元具代表性的物種,新世界之Afridinae與東方區之花布麗瘤蛾亞科(Camptolominae)則並未取樣到。方法上則選用粒腺體COI序列與七段獨立之核蛋白轉錄基因序列,作最簡約方法與模型化演化方法之親緣分析,並使用特徵最佳化分析(Mesquite ver. 2.75)來瞭解過去被視為在夜蛾總科具重建親緣意義特徵(例如三叉/四叉翅脈相、翅疆、聽器與作繭形式)的演化模式。

由重建親緣關係獲得以下主要結果:
  1. 瘤蛾科由下列形態共衍徵所支持:(1) 結船狀並具垂直羽化口的繭;(2) 前翅翅疆呈棒狀或指狀;(3) 鼓膜兜呈後氣孔式;
  2. 瘤蛾科包含六個具高支持度之枝系:中名未定之亞科(Diphtherinae), 長角瘤蛾亞科(Risobinae), 中名未定之亞科(Collomeninae), 中名未定之亞科(Beaninae), 旋瘤蛾亞科(Eligminae), 俊瘤蛾亞科(Westermanniinae), 瘤蛾亞科(Nolinae)與白襯瘤蛾亞科(Chloephorinae);其中Diphtherinae(DiphtheraLepidodes)Collomeninae為新世界特有的分類群;Diphtherinae因具有共衍徵(1)因此首次被視為瘤蛾科成員;Collomeninae因過去無清楚定義,在此文章中被正式描述為新亞科;印澳區廣布之Beana屬在親緣樹中與Eligminae, Westermanniinae, Nolinae之三叉支系成為姊妹群,因此被描述於新亞科Beaninae之下;
  3. Diphtherinae物種具有前額突出的結構,被視為成蟲與話時用來破繭用的功能性特徵,而雄蟲失去後脛節之近基部一對距的罕見雌雄異型特徵亦被重新提出來檢視;
  4. 白襯瘤蛾亞科所有屬級成員之雄蟲皆具有腹部基節的發聲器,此亞科下劃分鑽瘤蛾族(Eariadini)、赭瘤蛾族(Careini) 白襯瘤蛾族(Chloephorini)、皮瘤蛾族(Sarrothripini)與新族霜瘤蛾族(Gelastocerini),過去所定義之Ariolicini需後續研究。皮瘤蛾族下劃分皮瘤蛾亞族(Sarrothripina)、癬皮瘤蛾亞族(Blenina)與新亞族Etannina,後者具有附黑色鱗片的抱器突起之鑑識特徵。
  5. 花布麗瘤蛾類群雖未在此文研究中被取樣,然而與白襯瘤蛾族皆具有發達的發聲器結構,而兩者之幼蟲已知皆取食殼斗科,被推測具有潛在的近緣關係。
台灣產瘤蛾科物種於1992年出版之臺灣鱗翅目誌共收錄88(歸屬於夜蛾科皮瘤蛾亞科、白襯瘤蛾亞科與瘤蛾亞科),於此據Wan (1994; 1995), Inoue (2000), Fu & Tzuoo (2002), Holloway (2003), Fu & Hsu (2009), Kishida (2011); Sasaki (2011)Hu et al. (2013)之研究將臺灣產瘤蛾科名錄更新為6亞科107種並附備註如此連結

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