2009年11月22日 星期日

以分子序列初步重建鉤蛾總科親緣關係


A pilot study on the molecular phylogeny of Drepanoidea (Insecta: Lepidoptera) inferred from the nuclear gene EF-1a and the mitochondrial gene COI

文獻出處: Wu CG, Han HX, Xue DY. 2009. A pilot study on the molecular phylogeny of Drepanoidea (Insecta: Lepidoptera) inferred from the nuclear gene EF-1α and the mitochondrial gene COI. Bulletin of entomological research. 2009 Jul 7: 1-10 [Link]

簡介
本研究為中國科學院動物學組鱗翅目系統學研究組的武春光博士研究生、韓紅香與薛大勇博士,對鉤蛾總科類群首次進行分子親緣關係的重建,藉以初步釐清此總科內所含類群的單系性、各亞科間的關係,並以現有資訊討論長久具爭議性的鳳蛾科的分類歸屬。

由過去形態證據的研究中,鉤蛾、尺蛾、鳳蛾與燕蛾的分類地位與歸屬一直都有爭議。依據現今最被廣為使用的分類架構,鉤蛾總科包含了鳳蛾科與鉤蛾科,前者僅含少數屬與物種,而鉤蛾科中含鉤蛾亞科, 大鉤蛾亞科以及波紋蛾亞科共約700種描述種,其中鉤蛾亞科的Oreta屬常被提升為族級或亞科級地位。由於形態親緣關係的不穩定與研究者對不同特徵來定義分類階層的權重不一,近幾年來夜蛾科與尺蛾科使用多分子序列來進行高階親緣重建的方法也被應用在鉤蛾與鳳蛾分類群之上。

研究材料主要來自於舊北區中國產類群,包含鉤蛾總科的鳳蛾科2屬2種, 鉤蛾科11屬11種作為內群,外群選用夜蛾科與尺蛾科各2屬2種,表列於備註一並加入台灣產相關屬級或種級物種的網路影像連結。

方法上以核DNA EF-1
α序列以及結合EF-與粒腺體DNA COI序列所分別獲得的親緣樹顯示:
  1. 鳳蛾類群方面,兩屬兩物種Epicopeia hainesiPsychostrophia nymphidiaria形成單系群,此枝系在以EF-序列建構的樹型中與其他枝系關係不清楚,但在結合兩序列的樹型中與同為內群的鉤蛾科加上尺蛾科的枝系形成姊妹群,也就是在此分析結果中,過去鳳蛾歸屬於鉤蛾總科的處理並不適當。此結果在形態上可由鳳蛾並沒有鉤蛾科尺蛾的腹部聽器結構,以及此三分類群在前翅M脈系統、後翅Sc + R1脈與Rs脈關係之間的差異來支持。然而鳳蛾類群真正的分類地位可能需要涵蓋都多的尺蛾與燕蛾分類群與更多的分子序列來進行討論。
  2. 結合EF-與CII序列的樹型顯示廣義的鉤蛾科中Hypsomadius insignisOreta vatama形成單一枝系並與其他鉤蛾類群形成姊妹群,此結果對應形態上的特徵為HypsomadiusOreta的體型較壯碩呈黃褐色至棕色、下唇鬚短寬而密生毛束、口喙與翅刺(frenulum)不發達、後足脛節僅有一排等長的距刺;相對的其他鉤蛾姊妹群體修長呈白黃色、下唇鬚細長而分節明顯、口喙與翅刺皆發達、後足脛節有兩排距刺,兩枝系類群的生殖器結構亦有差異。本研究據此將過去提出的Oretinae亞科恢復使用,使現今鉤蛾總科包含四個亞科,然而作者承認所選取的分類群僅侷限於中國產物種,未來Oretinae類群與姊妹群間的關係尚須更多的分類群取樣來釐清。此外,馬達加斯加特產的 Nidarini(備註二)(包含五描述種)與其他鉤蛾亞科類群的關係應該也是未來待解決的問題。
  3. 鉤蛾亞科大鉤蛾亞科以及波紋蛾亞科在兩顆樹中都各自形成單系群,鉤蛾與大鉤蛾亞科形成姊妹群,以及此枝系與姊妹群大鉤蛾的樹型被高度的支持。此結果與Minet (2002)所提出的形態證據關係相符,他提出大鉤蛾相對波紋蛾與鉤蛾擁有較小且具有背方骨化區的聽器腔,此腔室的形態在大鉤蛾類群呈極度狹窄而至消失皆有。
備註一
Outgroup
Geometridae
Ennominae
Odontopera bilinearia coryphodes [Odontopera bilinearia subarida in Taiwan]
Geometrinae
Tanaorhinus viridiluteata [Tanaorhinus in Taiwan]
Noctuidae
Helicoverpa armigera [Species in Taiwan]
Catocala fraxini
[Catocala armandi shirozui in Taiwan]
Ingroup
Epicopeiidae
Epicopeia hainesi [Epicopeia mencia in Taiwan]
Psychostrophia nymphidiaria

Drepanidae
Cyclidiinae
Cyclidia substigmaria

Drepaninae
Callidrepana patrana [Species in Taiwan]
Ditrigona conflexaria
[D. triangularia in Taiwan]
Hypsomadius insignis [Species in Taiwan]
Macrocilix maia
[Species in Taiwan]
Macrocilix mysticata [M. mysticata flavotincta in Taiwan]
Microblepsis leucosticta
[M. violacea in Taiwan]
Oreta vatama
[O. loochooana in Taiwan]
Tridrepana fulvata
[T. flava in Taiwan]
Thyatirinae
Gaurena fletcheri

Habrosyne conscripta
[H. fraterna chekiangensis in Tawian]
Parapsestis lichenea


備註二
Minet與Scoble (1999)提出的鉤蛾亞科可分為三個族級分類群: 鉤蛾族 Oretini族以及Nidarini族

鉤蛾總科與相關參考文獻
1. Holloway JD. 1998. The Moths of Borneo: Families Castniidae, Callidulidae, Drepanidae and Uraniidae. The Malayan Nature Journal 52: 1–155.
2. Minet J. 2002. The Epicopeiidae: Phylogeny and a redefinition, with the description of new taxa (Lepidoptera: Drepanoidea). Annales de la Societe Entomologique de France 38: 463–487.
3. Minet J and Scoble MJ. 1999. The drepanoid/geometroid assemblage. pp. 301–320 in Kristensen NP (Ed.) Handbook of Zoology, Vol. IV. Arthropoda: Insecta. Part 35. Lepidoptera, Moths and Butterflies. Berlin & New York, Walter de Gruyter.
4. Nakajima H. 1970. A contribution to the knowledge of the immature stages of Drepanidae occurring in Japan. Tinea 8: 167–184.

Abstract
A molecular phylogenetic study of the Drepanoidea based on the EF-
sequences and combined EF-1α and COI sequences was carried out in order to infer higher classification at and above the subfamily level. The sample contained 14 taxa representing 13 genera recognized in the Drepanoidea. The results revealed that the Drepaninae, Thyatirinae and Cyclidiinae respectively form monophyletic groups. The sister relationship between the Drepaninae and the Thyatirinae was validated. The monophyly of the Cyclidiinae with the Drepaninae+Thyatirinae was supported robustly. Hypsomadius insignis and Oreta vatama within the traditional definition of the Drepaninae formed an individual clade with robust support (100%) and constitutes a sister relationship to a clade containing the rest of the Drepaninae in all the topologies, which means that the subfamily Oretinae of the Drepanidae should be restored. The family Drepanidae is divided into four subfamilies: Drepaninae, Oretinae, Thyatirinae and Cyclidiinae in this work. The family Epicopeiidae formed a monophyly with high bootstrap values. The result of combined analysis of EF- and COI showed that the Epicopeiidae have a closer phylogenetic relationship with the Geometridae than with the Drepanidae and belong to neither the Drepanoidea nor the Geometroidea.

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