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  1. NTU Theses and Dissertations Repository
  2. 生命科學院
  3. 生態學與演化生物學研究所
Please use this identifier to cite or link to this item: http://tdr.lib.ntu.edu.tw/jspui/handle/123456789/91242
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???org.dspace.app.webui.jsptag.ItemTag.dcfield???ValueLanguage
dc.contributor.advisor陳國勤zh_TW
dc.contributor.advisorBenny Kwok-Kan Chanen
dc.contributor.author李旻憲zh_TW
dc.contributor.authorMin-Hsien Leeen
dc.date.accessioned2023-12-12T16:22:12Z-
dc.date.available2023-12-13-
dc.date.copyright2023-12-12-
dc.date.issued2023-
dc.date.submitted2023-10-16-
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dc.identifier.urihttp://tdr.lib.ntu.edu.tw/jspui/handle/123456789/91242-
dc.description.abstract茗荷藤壺 (Lepas) 為海洋表層的生物 (epipelagic organism) ,牠們的成體 (adult) 附著於海洋漂流物上,此特性使得茗荷藤壺得以隨著洋流漂流而成為廣泛分佈於世界各個海洋的生物。近年來有研究以COI、16S及18S三個遺傳標記所構成的分子證據指出,茗荷藤壺的族群因為陸地阻礙、洋流流向與洋流表層水溫等地理隔閡效應的影響而分化成區域性的分支 (biogeographic clades) 。然而,西北太平洋海域卻因為只有5隻L. anserifera 採集於日本,而不能有代表性地代表西北太平洋地區的多樣性。本研究以台灣周圍海域的茗荷藤壺標本做分子與形態的綜合分析,以探討茗荷藤壺在台灣周圍海域的多樣性模式。從四個DNA條碼基因 (COI, 16S, 18S and H3) 所構成的親緣關係樹 (phylogenetic trees) 中,本研究發現在台灣茗荷藤壺共可分成四個物種及五個區域性分支 (biogeographic clades)。之前被認為只在美國奧勒岡州 (Oregon) 發現之 L. anatifera (Oregon clade) 也出現在台灣海域,故本論文改稱之為L. anatifera北太平洋分支 (North-Pacific clade)。本論文也鑑定出L. pectinata於西北太平洋的一個新的分支 (NW-Pacific clade) 與一個未知的物種Lepas sp.。L. anatifera的北太平洋分支與L. pectinata的西北太平洋分支可能是受到北太平洋環流 (North Pacific gyre) 與赤道逆流 (Equatorial Countercurrent) 局限而分化。L. anatifera的全球性分支 (Global-clade) 與L. anserifera 也同樣出現在台灣海域,而L. anserifera是唯一未分化的全球性物種 (cosmopolitan species) 。以Assemble Species by Automatic Partitioning (ASAP) 和Bayesian implementation of the Poisson Tree Processes Model (bPTP) 兩種物種界定方法做出的結果也顯示,各個物種內大多數的區性分支之間的差異可能足以視為不同物種,而綜合分子與形態分析後,本研究認為L. anatifera 北太平洋分支與全球性分支可被視為不同物種。此外,未知物種L. sp.在過往曾以外觀特徵被認為是L. testudinata。然而,本研究經由分子證據佐以對過往的文獻記錄做形態上的比對後,認為L. sp.與L. testudinata為不同物種。L. testudinata模式地 (type locality) 於南非,故分佈於南半球的物種為L. testudinata,而本論文採得於西北太平洋的物種可被視為一個未知的新物種L. sp.。zh_TW
dc.description.abstractGooseneck barnacles of the genus Lepas are epipelagic organisms. Their adults attach to ocean-floating objects and can be dispersed by the ocean currents. Lepas barnacles thus have worldwide distributions. Previous studies studied the diversity of Lepas using COI, 16S, and 18S genes and showed that the populations were differentiated into several biogeographic clades which are influenced by vicariance effects of continental isolation, ocean currents, and sea surface temperature (SST). However, the diversity in the Northwest Pacific was understudied because the pattern addressed was based on five L. anserifera specimens collected from Japan. In the present study, extensive Lepas specimens were collected around Taiwan and were analyzed with DNA barcoding genes (COI, 16S, 18S, and H3 genes) and morphological approaches. Compared to previous studies, our study revealed the original Oregon clade of Lepas anatifera is also present in Taiwan waters and we named it as North-Pacific clade. We have identified one more clade of L. pectinata which is distributed in the northern hemisphere of the Pacific. We also identified one unknown, probably new species of Lepas in the NW Pacific. The differentiations of L. anatifera North-Pacific clade and L. pectinata NW-Pacific clade may be caused by the limited gene flow which is affected by the North Pacific gyre and Equatorial Countercurrent. The Taiwan waters also contain the Global clade of L. anatifera and L. anserifera. Only L. anserifera is undifferentiated and is cosmopolitan in the world. The species delimitation results of Assemble Species by Automatic Partitioning (ASAP) and Bayesian Implementation of the Poisson Tree Processes Model (bPTP) also suggested most regional clades of Lepas species may be considered as separate species. According to the combination of morphological analysis and molecular analysis, the North-Pacific clade and Global clade of L. anatifera can be considered as different species. In addition, the undescribed L. sp. identified from the present study was morphologically similar to L. testudinata. However, the present study shows they are different species according to the molecular evidence and historical taxonomic records. L. testudinata was first identified in South Africa, and therefore the L. testudinata in the southern ocean represent the true L. testudinata and the species identified in the Northwest Pacific (morphologically similar to L. testudinata) can be revealed as an undescribed new species.en
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dc.description.tableofcontents口試委員會審定書 i
Acknowledgements ii
摘要 iii
Abstract v
1. Introduction 1
1.1 Biogeography and phylogeography of barnacles 1
1.2 The pelagic stalked barnacle Lepas 4
1.3 History of taxonomic research in Lepas 4
1.3.1 First identification of the genus Lepas 4
1.3.2 Subgenera of Lepas 5
1.3.3 Valid species of Lepas 7
1.3.4 Species concept and species identities of Lepas 13
1.4 Molecular phylogeny and biodiversity of Lepas 19
1.5 Aim 19
2. Materials and methods 21
2.1 Sampling and preservation of specimens 21
2.2 DNA extraction, PCR, and sequencing 21
2.3 Phylogenetic analysis 22
2.4 Species delimitation 26
2.5 Morphological analysis 27
2.5.1 Specimen groupings 27
2.5.2 Multivariate analysis 28
2.5.3 Morphology of cirri, somatic body, and mouth parts. 29
3. Results 31
3.1 Phylogenetic analysis 31
3.1.1 18S nuclear gene phylogenetic pattern 31
3.1.2 COI, 16S mitochondrial phylogenies and concatenating analysis 32
3.1.3 H3 phylogeny and concatenating analysis of sequences from Taiwan 36
3.2 Species delimitation 38
3.3 Morphological analysis 40
3.3.1 External variances 40
3.3.2 Multivariate analysis 49
3.3.3 Observation of somatic body 50
4. Discussion 64
4.1 Diversity of Lepas 64
4.1.1 Diversity of Lepas in Taiwan 64
4.1.2 Genetic diversity of Lepas 65
4.2 Taxonomic issue of Lepas in Taiwan 72
4.2.1 Global clade and North-Pacific clade of L. anatifera 72
4.2.2 L. anserifera 78
4.2.3 L. pectinata NW-Pacific clade 79
4.2.4 L. sp. 83
Conclusions 87
Reference 89

List of Figures
Fig. 1. General characters of Lepas 110
Fig. 2. Sampling locations around the coastline of Taiwan and outlying islands. 111
Fig. 3. Parameters of multivariate analysis for comparing the differences in capitulum shape between between Global clade and North-Pacific clade of L. anatifera. 112
Fig. 4. 18S nuclear gene phylogeny. 113
Fig. 5. 16S mitochondrial phylogeny. 114
Fig. 6. COI and 16S combined mitochondrial phylogeny. 115
Fig. 7. Species delimitation with COI and 18S combined phylogeny. 116
Fig. 8. H3 mitochondrial phylogeny. 117
Fig. 9. COI, 16S, 18S, and H3 combined mitochondrial phylogeny. 118
Fig. 10. External variations of Global clade of L. anatifera. 119
Fig. 11. External variations of North-Pacific clade of L. anatifera. 120
Fig. 12. External variations of L. anserifera. 121
Fig. 13. External variations of L. sp. 122
Fig. 14. External variations of NW-Pacific clade of L. pectinata. 123
Fig. 15. 2-dimensional nMDS ordination plot generated from the 20 parameters displayed in Fig. 3. 124
Fig. 16. Filamentary appendages of L. anatifera Global clade 125
Fig. 17. Cirri I and II of L. anatifera Global clade 126
Fig. 18. Cirri III and IV of L. anatifera Global clade 127
Fig. 19. Cirri V and VI of L. anatifera Global clade 128
Fig. 20. Maxilla, maxillule and mandible of L. anatifera Global clade 129
Fig. 21. Mandible palp and labrum of L. anatifera Global clade 130
Fig. 22. Filamentary appendages of L. anatifera North-Pacific clade 131
Fig. 23. Cirri I and II of L. anatifera North-Pacific clade 132
Fig. 24. Cirri III and IV of L. anatifera North-Pacific clade 133
Fig. 25. Cirri V and VI of L. anatifera North-Pacific clade 134
Fig. 26. Maxilla, maxillule and mandible of L. anatifera North-Pacific clade 135
Fig. 27. Mandible palp and labrum of L. anatifera North-Pacific clade 136
Fig. 28. Filamentary appendages of L. anserifera 137
Fig. 29. Filamentary appendages of L. anserifera 138
Fig. 30. Cirri I and II of L. anserifera 139
Fig. 31. Cirri III and IV of L. anserifera 140
Fig. 32. Cirri V and VI of L. anserifera 141
Fig. 33. Maxilla, maxillule and mandible of L. anserifera. 142
Fig. 34. Mandible palp and labrum of L. anserifera 143
Fig. 35. Filamentary appendages of L. sp. 144
Fig. 36. Filamentary appendages of L. sp. 145
Fig. 37. Cirri I and II of L. sp. 146
Fig. 38. Cirri III and IV of L. sp. 147
Fig. 39. Cirri V and VI of L. sp. 148
Fig. 40. Maxilla, maxillule and mandible of L. sp. 149
Fig. 41. Mandible palp and labrum of L. sp. 150
Fig. 42. Filamentary appendages of L. pectinata NW-Pacific clade. 152
Fig. 43. Cirri I and II of L. pectinata NW-Pacific clade. 153
Fig. 44. Cirri III and IV of L. pectinata NW-Pacific clade 154
Fig. 45. Cirri V and VI of L. pectinata NW-Pacific clade 155
Fig. 46. Maxilla, maxillule and mandible of L. pectinata NW-Pacific clade. 156
Fig. 47. Mandible palp and labrum of L. pectinata NW-Pacific clade. 157
Fig. 48. Comparison of teeth numbers on mandibles of L. pectinata NW-Pacific clade. 158
Fig. 49. Comparison of setal types on distal region of Cirri I and endopodite of Cirri II in Global clade of L. anatifera. 160
Fig. 50. Comparison of setal types on distal region of Cirri I and endopodite of Cirri II in North-Pacific clade of L. anatifera. 162
Fig. 51. The updated diversity of Lepas. 163

List of Tables
Table 1. Nomenclature and type status of currently valid Lepas species 164
Table 2. Valid species and molecular information 165
Table 3. Used primers and annealing temperatures 168
Table 4. List of morphotypes and quantities of used specimens in multivariate analysis 169
Table 5. Pairwise distances of COI in Lepas 170
Table 6. Results of multivariate analysis based on parameters in Fig. 3. 171

List of Appendixes
Appendix 1. Sample information in the present study. 172
Appendix 2. Reference sample information from Schiffer & Herbig (2016). 186
Appendix 3. 16S phylogeny 194
Appendix 4 18S phylogeny 196
Appendix 5. H3 phylogeny 198
Appendix 6. COI+16S phylogeny 199
Appendix 7. COI+16S+18S phylogeny 201
Appendix 8. COI+16S+18S+H3 phylogeny 203
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dc.language.isoen-
dc.subject茗荷屬zh_TW
dc.subject鵝頸藤壺zh_TW
dc.subject茗荷屬zh_TW
dc.subject隱蔽種zh_TW
dc.subject地理隔離zh_TW
dc.subject親緣關係zh_TW
dc.subject形態zh_TW
dc.subject物種界定zh_TW
dc.subject生物地理學zh_TW
dc.subject生物地理學zh_TW
dc.subject物種界定zh_TW
dc.subject形態zh_TW
dc.subject親緣關係zh_TW
dc.subject地理隔離zh_TW
dc.subject隱蔽種zh_TW
dc.subject鵝頸藤壺zh_TW
dc.subjectvicariance effecten
dc.subjectbiogeographyen
dc.subjectspecies delimitationen
dc.subjectmorphologyen
dc.subjectphylogenyen
dc.subjectvicariance effecten
dc.subjectcryptic speciesen
dc.subjectgenus Lepasen
dc.subjectgoose barnacleen
dc.subjectbiogeographyen
dc.subjectspecies delimitationen
dc.subjectmorphologyen
dc.subjectgoose barnacleen
dc.subjectgenus Lepasen
dc.subjectcryptic speciesen
dc.subjectphylogenyen
dc.title台灣海域茗荷藤壺之隱蔽種多樣性zh_TW
dc.titleCryptic diversity of goose-neck barnacle Lepas in Taiwan watersen
dc.typeThesis-
dc.date.schoolyear112-1-
dc.description.degree碩士-
dc.contributor.oralexamcommittee施習德;町田龍二zh_TW
dc.contributor.oralexamcommitteeHsi-Te Shih;Ryuji Machidaen
dc.subject.keyword鵝頸藤壺,茗荷屬,隱蔽種,地理隔離,親緣關係,形態,物種界定,生物地理學,zh_TW
dc.subject.keywordgoose barnacle,genus Lepas,cryptic species,vicariance effect,phylogeny,morphology,species delimitation,biogeography,en
dc.relation.page203-
dc.identifier.doi10.6342/NTU202304335-
dc.rights.note同意授權(限校園內公開)-
dc.date.accepted2023-10-18-
dc.contributor.author-college生命科學院-
dc.contributor.author-dept生態學與演化生物學研究所-
dc.date.embargo-lift2028-10-16-
Appears in Collections:生態學與演化生物學研究所

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