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研究生: 林容諄
Lin, Rung-Juen
論文名稱: 蠶蛾科之起源、演化及其生物學研究
The Biology, Evolution and Origin of Silkworms and their Relatives (Lepidoptera: Bombycidae)
指導教授: 徐堉峰
Hsu, Yu-Feng
口試委員: Michael F. Braby
Michael F. Braby
陳韋仁
Chen, Wei-Jen
李壽先
Li, Shou-Hsien
林思民
Lin, Si-Min
徐堉峰
Hsu, Yu-Feng
口試日期: 2022/01/12
學位類別: 博士
Doctor
系所名稱: 生命科學系
Department of Life Science
論文出版年: 2022
畢業學年度: 110
語文別: 英文
論文頁數: 211
中文關鍵詞: 蠶蛾科親緣關係系統生物學生物地理特徵演化
英文關鍵詞: Bombycidae, character evolution, historical biogeography, phylogeny, systematics
研究方法: 實驗設計法比較研究田野調查法
DOI URL: http://doi.org/10.6345/NTNU202200311
論文種類: 學術論文
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  • 馴化已久的家蠶(Bombyx mori)(Lepidoptera: Bombycidae),是鱗翅目的模式物種之一,為第一個完成全基因組序列定序(genome sequence)的鱗翅目物種。由於家蠶結繭的蠶絲及蠶絲的副產品擁有重要的經濟價值,根據文獻記載早在5000年前就開始被人類馴養利用,目前有超過1000種以上的品系。近年結合分子數據證據透過馴化品系之間的基因組關係研究顯示,家蠶(B. mori)是由野蠶(B. mandarina)所馴養而來,與歷史文獻紀錄吻合,透過絲路分別傳往世界各地。儘管家蠶研究深入,然而蠶蛾科內其他與蠶蛾近緣物種的研究資料十分缺稀,例如多數的蠶蛾科物種沒有完整的生活史資料,蠶蛾科內不同屬之間的關係尚未釐清,幼蟲取食桑科(Moraceae)桑屬(Morus)或榕屬(Ficus)植物的演化情形未知,且蠶蛾科的分布起源仍然充滿疑問。本研究取樣所有文獻記錄地區之蠶蛾科樣本,囊括亞洲、南美洲、非洲、澳洲及東南亞地區,研究結果簡述如下:1) 採集、飼養蠶蛾科物種、記錄其生活史資料包含寄主植物、卵的排列方式、初齡幼蟲之原生毛序、繭的結構與色型等;2) 以6個基因(1個粒線體基因與5個核基因)序列資料建構可靠的蠶蛾科親緣關係,結果顯示蠶蛾科可分為南美洲新世界亞科Epiinae與亞洲古北區舊世界亞科Bombycinae兩個亞科,屬級關係中除Bivincula與Gunda為併系群,其餘屬為單系群;3) 根據幼蟲的寄主植物記錄與分子數據建構的親緣關係樹進行特徵演化模擬,推測利用桑科榕屬做為寄主植物為祖徵,而利用桑科桑屬植物則為較晚近演化出來的結果;4) 利用新建構之分子親緣關係樹結合化石、地質年代與分子鐘估算物種分化時間,重建蠶蛾科之生物地理歷史事件,估算蠶蛾科約在6千1百萬年前於岡瓦那大陸(Gondwana)出現,約在5千1百萬年前分化為新世界與舊世界兩大亞科;分析顯示,Bombycinae的祖先約在4千6百萬年前經由一次澳洲至亞洲的擴散事件(dispersal)形成現今物種在東南亞及亞洲的分布;而後又再經由一次的擴散事件,從亞洲擴散至非洲地區。

    The silkeworm, Bombyx mori (Lepidoptera: Bombycidae) is an important model species, which is well studied, representing the first species in which the whole genome sequence was completed in the order of Lepidoptera. According to previous studies, there are over 1,000 inbred strains of silkworms which were domesticated at least 5,000 years ago for producing valuable silk. Recently, it was demonstrated that silkworms were domesticated from wild silkworm B. mandarina by resequencing 40 genomes of domesticated and wild silkworms, and that economically valuable silkworms were spread worldwide through the Silk Road. Although there have been many studies on silkworm, there is a lack of information on other species in Bombycidae, such as life histories, evolutionary relationships, the evolution of life history characters and their hostplants, and their origin. This study investigates the morphology, biology, character evolution, origin and historical biogeography of silkworms and their relatives in Asia, East-Palearctic, Australia, South America, and Africa. The main findings and results are summarized as follows: 1) comparative illustrations of the immature stages, morphology, male, and female genitalia are provided for 13 species in 11 genera; 2) the well-supported phylogeny based on a six gene data set indicates the family Bombycidae comprises two subfamilies, Epiinae and Bombycinae; most of genera were monophyletic, although Bivincula and Gunda were polyphyletic; 3) ancestral trait reconstruction suggests Moraceae is the ancestral host plant family, with feeding on the genus Ficus likely the most recent common ancestor; 4) reconstruction of the biogeographical history of Bombycidae suggests an origin in Southern Gondwana (Australia–Antarctica–South America) during the Paleocene (c. 61 Mya); the family then split vicariantly into its two subfamilies in the Eocene (c. 51 Mya) as follows: Bombycinae (Australia) + Epiinae (South America). Asia was subsequently colonized by dispersal out of Australia (c. 46 Mya). The other continents and land masses in Africa and East-Palaearctic were later colonized by dispersal or range expansion from Asia.

    Acknowledgements i Abstract ii List of Tables vi List of Figures vii Chapter 1: General Introduction 1 Silk and silkworm 1 Bombycidae 1 Chapter 2: Systematics and Biology of Bombycidae 6 Introduction 6 Systematics of Bombycidae 6 Checklist of Bombycidae of the World 10 Checklist of Bombycidae of Taiwan 10 Materials and Methods 12 Taxon sampling and field observations 12 Laboratory rearing 13 Molecular data 14 Sequence alignment and phylogenetic analysis 15 Results 16 Molecular phylogeny 16 Immature stage biology 18 Discussion 39 Chapter 3: Character Evolution in Bombycidae 46 Introduction 46 Host plants 47 Egg-laying behavior 49 Larval color pattern 50 Cocoons 51 Materials and Methods 53 Taxon sampling and data collection 53 Molecular work 53 Life history characters 54 Character evolution 56 Phylogenetic signal 57 Results 58 Host plants 59 Egg cluster size and deposition pattern 59 Larval color pattern 60 Cocoon color, structure and pupation sites 60 Sensitivity of the phylogenetic signal test 61 Discussion 63 Host plant shifts 63 Egg cluster size and orientation 64 Larval color pattern 66 Cocoon color, structure and pupation sites 68 Chapter 4: Historical Biogeography of Bombycidae 70 Introduction 70 Materials and Methods 74 Taxon sampling 74 DNA extraction, amplification, and sequencing 75 Phylogenetic analyses 75 Divergence times 76 Biogeographic analyses 77 Results 77 Phylogenetic relationships 77 Divergence times and molecular dating 78 Biogeographic reconstruction 79 Discussion 80 Chapter 5: Conclusion 86 Summary 86 Limitations of available data 88 References 90 Tables 104 Figures 120 Appendixes 186

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