透翅蛾科之系統分類地位研究以及臺灣產物種之分類修訂與生物學研究

Systematics of Sesiidae (Lepidoptera: Sesioidea), with special reference to taxonomy and biology of species in Taiwan

梁家源(Liang Jia-Yuan)

指導教授 ： 徐堉峰

國立臺灣師範大學/理學院/生命科學系/博士(2017年)

https://doi.org/10.6345/NTNU202202053

摘要

透翅蛾科為透翅蛾總科中數量及物種多樣性最高的類群，目前全世界已被鑑定命名的透翅蛾科種類約有160屬、1454種和51亞種。透翅蛾科於全球幾乎都有採集記錄，其中以東洋區的多樣性最高。透翅蛾科幼蟲為鑽蛀性，會蛀食寄主植物的樹幹、樹皮、嫩枝、根部和果實等組織，或者在寄主植物的莖和枝條上形成蟲癭，並造成農業和當地植物的危害。透翅蛾科的擬蜂形態特徵使其過去一直受到許多鱗翅目研究者矚目，最近的研究中雖已證實透翅蛾科為一個單系群，但在透翅蛾科中除了少數幾個屬間的親緣關係曾被探討過之外，透翅蛾科內的系統發育關係至今缺乏有系統的分析，對暸解這一多樣性很高的類群的起源、食性演化、生物地理都是很大的障礙。本研究目的為(1)用形態及分子序列之證據來探討透翅蛾科的親緣關係，檢測目前使用的不同亞科及族級分類單元是否為單系群，藉以作出分類上的調整，以及進一步探討成蟲擬蜂特徵起源及幼生期食性演化之問題。(2) 重新修訂處理臺灣產透翅蛾科的分類問題，並記錄幼生期寄主植物、取食特性和雌蟲產卵行為等生物學資料。材料上選取透翅蛾科中75個屬96個物種作為研究內群，以木蠹蛾科、蝶蛾科和短軀蛾科作為外群。本研究共計使用4科84屬105個物種，其中78屬80種使用16個幼蟲、9個蛹、19個成蟲非生殖器、16個雄生殖器與6個雌生殖器外部構造特徵，以最大簡約分析重建形態親緣關係樹；38屬60種使用粒線體DNA COⅠ、COII序列以及核DNA EF-1α、wingless序列資料，以最大或然分析和貝式分析來重建分子親緣關係樹。結果顯示透翅蛾科為單系群，其短軀蛾科為最近的姊妹群；在亞科分類層級中，Sesiinae亞科為單系群，Tinthiinae亞科呈併系群；在族級分類層級中，分子資料支持各族皆為單系群，Sesiinae亞科內各族之間的關係以 ((Melittiini + Sesiini) + (Paranthrenini (Osminiini + Cissuvorini))) + Synanthedonini呈現，形態資料支持Tinthiini、Paraglosseciini、Pennisetiini、Similipepsini、Melittiini、Cissuvorini、Osminiini、Paranthrenini和Synanthedonini族各為單系群。其成蟲擬態蜂類外觀特徵的出現，為類群或物種各自的平行演化。幼生期宿主利用特徵起始階段重建分析顯示，透翅蛾科祖先的幼生期寄主利用形態應為蛀莖性，從蛀莖轉為造癭的演化過程為多次的獨立演化。在臺灣產透翅蛾科的物種分類修訂上，本研究進行1個新組合與5個同物異名處理；臺灣產透翅蛾科物種共計24屬43種，其中包含2個新紀錄屬、8個新種、1新紀錄種，並記錄15種物種的生活史資料。

關鍵字

透翅蛾 ； 姬透翅蛾亞科 ； 透翅蛾亞科 ； 親緣關係 ； 寄主植物

並列摘要

The Sesiidae (clearwing moths) are the most diverse and numerous moths within the superfamily Sesioidea, with 1454 species (with addition of 51 subspecies) in 160 genera already described worldwide. The family reaches the highest species diversity in the Oriental Region. The larvae of clearwing moths are mostly borers in tree trunk, bark, twig, root, and fruits, and some species are known to induce galls, causing damages to crops in agriculture and trees in forestry. The appearance of wasp/bee-mimicry of clearwing moths in external morphology has been a focus of interest to biologists. Although recent phylogenetic studies on higher classification of Lepidoptera based suggested Sesiidae is a monophyletic group, the infra-family systematic of clearwing moths have not been worked out, forming obstacles on our understanding of their origin, evolution of feeding habits, and biogeography. The aim of the present study focuses on the Sesiidae (1) to examine monophyly and phylogenetic relationship of infra-family taxa (tribes) of clearwing moths, and explore the questions with regard to origin of these wasp/bee-mimicry and feeding habits; (2) to comprehensively revise the Taiwanese fauna and documents host associations and immature biology. In this study, 75 genera and 96 species of Sesiidae ingroup taxa plus Cossidae, Castniidae and Brachodidae as outgroups for phylogenetic reassessment. Totally 4 family 84 genera and 105 species were collected. 78 genera and 80 species intended maximum parsimony analysis of reconstructing morphological phylogeny based on morphological characters from all development stages (19 larva, 9 pupa, 19 adult, 16 male genitalia, 6 female genitalia). 38 genera and 60 species intended maximum likelihood analysis and Bayesian inference analysis of reconstructing molecular phylogeny based on COI, COII, EF-1α and wingless combined sequences. The result shows the Sesiidae is a monophyletic group, and Brachodidae as sistergroup. At subfamily level, Sesiinae is a monophyletic group, but Tinthiinae is a paraphyletic group. At tribe level, all tribes were retrieved as monophyletic groups based on molecular data within Sesiinae, the relationship ((Melittiini + Sesiini) + (Paranthrenini (Osminiini + Cissuvorini))) + Synanthedonini. Tribes Tinthiini, Paraglosseciini, Pennisetiini, Similipepsini, Melittiini, Cissuvorini, Osminiini、Paranthrenini and Synanthedonini are all retrieved as monophyletic groups with inter-tribal relationship umresolved based on morphology. The appearance of mimicry of apparent characters of wasp in adults is parallel evolution in groups or species. The host use ASR in larval stage, that the ancestral state of the Family Sesiidae is stem-boring feeding habits. The transformation of stem-boring to gall-making is the results of many independent evolution. In taxonomic accounts, 24 genera and 43 species are recorded in Taiwanese fauna, including 2 newly recorded genera, 8 newly species, 1 newly recorded species, 5 new synonymies and 1 new combination. Host associations and immature biology are documented for 15 species in Taiwan.

並列關鍵字

clearwing moths ； Tinthiinae ； Sesiinae ； phylogeny ； host plants

參考文獻

松村松年 (1911) 續日本千蟲圖解：第3卷。警醒社。東京。147頁。

松村松年 (1931) 日本昆蟲大圖鑑。刀江書院。東京。1479頁。

Ahmad M., R. Mishra & J. Ahmad (2001) Insect pest spectrum of poplar in India. Indian Forester. 127(12): 1353-1366.

Areekul B. & D. L. J. Quicke (2006) The use of colour characters in phylogenetic reconstruction. Biological Journal of the Linnean Society. 88:193-202.

Arita Y. & O. G. Gorbunov (1995) A review of the genus Macrotarsipus Hampson, [1893] (Lepidoptera, Sesiidae) of the Oriental region. Transactions of the Lepidopterological Society of Japan. 6(2): 103-111.