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Kobe University Repository : Thesis Kobe University Repository : Thesis Phylogeography of the subfamily Scaphidiinae (Coleoptera、 学位論文題目 Staphylinidae) in Sulawesi, with its systematic revision(インドネシア・ Title スラウェシ産デオキノコムシ亜科(コウチュウ目、ハネカクシ科) の系 統地理学と分類学的検討) 氏名 Ogawa, Ryo Author 専攻分野 博士(農学) Degree 学位授与の日付 2015-03-25 Date of Degree 公開日 2020-03-25 Date of Publication 資源タイプ Thesis or Dissertation / 学位論文 Resource Type 報告番号 甲第6345号 Report Number 権利 Rights JaLCDOI URL http://www.lib.kobe-u.ac.jp/handle_kernel/D1006345 ※当コンテンツは神戸大学の学術成果です。無断複製・不正使用等を禁じます。著作権法で認められている範囲内で、適切にご利用ください。 PDF issue: 2020-05-01 Doctoral Dissertation Phylogeography of the subfamily Scaphidiinae (Coleoptera, Staphylinidae) in Sulawesi, with its systematic revision Ryo OGAWA Laboratory of Insect Biodiversity and Ecosystem Science, Graduate School of Agricultural Science, Kobe University February, 2015 Doctoral Dissertation Phylogeography of the subfamily Scaphidiinae (Coleoptera, Staphylinidae) in Sulawesi, with its systematic revision インドネシア・スラウェシ産デオキノコムシ亜科(コウチュウ目, ハネカクシ科) の系統地理学と分類学的検討 Ryo OGAWA Laboratory of Insect Biodiversity and Ecosystem Science, Graduate School of Agricultural Science, Kobe University February, 2015 Table of Contents Abstract ........................................................................................................................... 1 Declaration ...................................................................................................................... 2 Chapter 1 ― Introduction ............................................................................................... 3 1. 1 Family Staphylinidae .......................................................................................... 4 1. 1. 1 Phylogeny of the Staphylinidae ................................................................... 4 1. 1. 2 Mycophagy, myxomycophagy, and termitophily ........................................ 5 1. 2 Subfamily Scaphidiinae ...................................................................................... 6 1. 2. 1 Phylogeny of the Scaphidiinae .................................................................... 7 1. 2. 2 Sulawesian scaphidiines .............................................................................. 7 1. 3 Sulawesi Island ................................................................................................... 8 1. 3. 1 Wallacea ...................................................................................................... 8 1. 3. 2 Geological history of Sulawesi .................................................................. 10 1. 3. 3 Invertebrate biogeography of Sulawesi ..................................................... 11 1. 4 Aims of this study ............................................................................................. 11 References ................................................................................................................... 12 Chapter 2 ― Materials and Methods ............................................................................ 17 2. 1 Sampling methods ............................................................................................ 18 2. 1. 1 Sampling localities .................................................................................... 19 2. 1. 2 Deposition of voucher specimens .............................................................. 19 2. 1. 3 Taxon sampling for molecular phylogenetics ........................................... 20 2. 2 Dissection and morphological observation....................................................... 20 2. 3 Morphological terminology .............................................................................. 20 2. 4 DNA extraction, amplification, and sequencing .............................................. 21 2. 5 Sequence alignment, model selection, and phylogenetic analyses ................... 22 i References ................................................................................................................... 30 Chapter 3 ― Phylogeny of the Subfamily Scaphidiinae .............................................. 33 3. 1 Phylogenetic analyses based on 28S sequences ............................................... 35 3. 2 Ancestral state reconstruction for feeding habits ............................................. 38 3. 3 Estimation of divergence time .......................................................................... 40 References ................................................................................................................... 42 Appendix ..................................................................................................................... 44 Chapter 4 ― Systematics of the Subfamily Scaphidiinae from Sulawesi .................... 47 4. 1 Genus Scaphidium ............................................................................................ 49 4. 2 Genus Cyparium ............................................................................................... 73 4. 3 Scaphisoma group ............................................................................................ 75 4. 3. 1 Termitophilous scaphidiines ...................................................................... 75 4. 3. 1. 1 Genus Baeoceridium .......................................................................... 76 4. 3. 1. 2 Genus Termitoscaphium ..................................................................... 77 4. 3. 1. 3 Genus Vituratella ................................................................................ 78 4. 3. 2 Genus Scaphisoma ..................................................................................... 80 4. 4 Genus Birocera ............................................................................................... 100 4. 5 Genus Bironium .............................................................................................. 101 4. 6 Baeocera group .............................................................................................. 103 4. 6. 1 Genus Baeocera ....................................................................................... 104 4. 6. 2 Genus Scaphobaeocera ........................................................................... 108 4. 6. 3 Genus Xotidium ....................................................................................... 112 4. 7 Genus Scaphoxium ......................................................................................... 114 4. 8 Genus Scaphicoma ......................................................................................... 116 References ................................................................................................................. 126 Figures ...................................................................................................................... 131 ii Chapter 5 ― Phylogeography of the Subfamily Scaphidiinae from Sulawesi ........... 157 5. 1 Phylogenetic relationships of Sulawesian species .......................................... 158 5. 2 Genus Scaphidium .......................................................................................... 158 5. 3 Termitophilous scaphidiines ........................................................................... 160 5. 4 Genus Scaphisoma .......................................................................................... 161 5. 5 Genus Scaphicoma ......................................................................................... 163 References ................................................................................................................. 164 Chapter 6 ― General Discussion ................................................................................ 165 6. 1 Fauna of Sulawesian scaphidiines .................................................................. 166 6. 2 “Vicariance” versus “dispersal” ..................................................................... 166 6. 3 Biogeographical origins of Sulawesian scaphidiines ..................................... 167 6. 4 Conclusion ...................................................................................................... 168 References ................................................................................................................. 169 Acknowledgements ..................................................................................................... 170 Appendix ...................................................................................................................... 173 iii iv Abstract Sulawesi, which is the largest island in the biodiversity hotspot of Wallacea as a transitional ecozone between the Oriental region and the Australasian region, has been formed through the collisions of three continental plates (Sundaland, Philippines, and Sahul). Thus this island’s endemic fauna has multiple elements in phylogeography. The origins of Sulawesi's fauna have been debated on the variety of organisms by many authors, but those of invertebrate fauna are still controversial with regard to the role of “vicariance” versus “dispersal”. As mycophagous beetles, the Scaphidiinae (Coleoptera, Staphylinidae) depend on the fungi grown up in humid forests and thus likely migrate with the forests on the continental plates in displacement. Therefore, phylogeographical studies of them are expected to support the “vicariance” hypothesis. This study, for the consideration of the role of "vicariance” versus “dispersal" on the scaphidiine fauna
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