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Cyclophorus and Cyclotus Taivanus Ssp

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Cyclophorus and Cyclotus Taivanus Ssp 國立臺灣師範大學生命科學系 博士論文 大山蝸牛屬及台灣山蝸牛屬之種化事件 與山蝸牛科之系統發育學研究 Cyclophoridae phylogeny and the speciation events of Cyclophorus and Cyclotus taivanus ssp. 研究生：李彥錚 Yen-Chen Lee 指導教授：呂光洋 Kuang-Yang Lue 巫文隆 Wen-Lung Wu 中華民國九十七年六月 96.11. 版學位論文授權書 國 立 臺 灣 師 範 大 學 學 位 論 文 授 權 書 本授權書所授權之論文為授權人在國立臺灣師範大學 生命科學 研究所 96 學年度第 2 學期取得 博 士學位之論文。 論文題目：大山蝸牛屬及台灣山蝸牛屬之種化事件與山蝸牛科之系統發育學 研究 指導教授：呂光洋‧巫文隆 授權事項： □同意 一、 授權人 非專屬無償授權本校及國家圖書館將上列論文資 □不同意 料以微縮、數位化或其他方式進行重製，並可上載網路收錄於本校博 碩士論文系統、國家圖書館全國博碩士論文資訊網及臺灣師範校院聯 合博碩士論文系統，提供讀者基於個人非營利性質之線上檢索、瀏覽、 下載、傳輸、列印或複印等利用。 二、 論文全文電子檔上載網路公開時間：【第一項勾選同意者，以下須擇一勾選】 □ 即時公開 □ 自＿ 2009 年＿＿8 月＿＿31 日始公開。 授權人姓名： （請親筆正楷簽名） 學 號：891430048 註：1. 本授權書須列印並簽署兩份，一份裝訂於紙本論文書名頁，一份繳至圖書館辦理離校手續。 2. 授權事項未勾選者，分別視同「同意」與「即時公開」。 中 華 民 國 97 年 6 月 27 日 Content Abstract………………………………………………………………………....………..III Chapter 1 General introduction ……………………………………...…………..………..1 Chapter 2 Molecular phylogeny of the family Cyclophoridae (Gastropoda: Architaenioglossa) in East Asia…………………………………..……………3 Chapter 3 Ring speciation and morphological adaptations of genus Cyclophorus in Taiwan………………………………………………………………………...37 Chapter 4 The phylogenetic evolution and morphological adaptations in Cyclotus taivanus ssp.………………………………………...…………..…………….63 Chapter 5 The cyclophorids fauna of Taiwan……………..……..………………………91 Chapter 6 Summary and conclusion……………………………………………………151 References………………………………………………………………………………155 Acknowledgment…………………………………………………………...…………..163 I II Abstract Cyclophoridae consists of four subfamilies and about 300 species currently arranged in 38 genera, occuping varies habitats, with great morphological diversity allover the world. However, the relationship among Cyclophoridae is thus far not clear. In order to investigate this, I sampled cyclophorid snails around Taiwan and its adjacent areas, and then sequenced part of the mitochondrial COI (cytochrome oxidase subunit I) and the 16S rRNA gene from 32 species of 10 cyclophorid genera to establish a phylogenetic tree of Cyclophoridae. Phylogenetic relationships based on mtDNA sequences suggest that Cyclophorus, Cyclotus, Leptopoma, and Platyrhaphe are monophyletic while the traditional genus Japonia is polyphyletic, and the previous J. zebra should be placed into a new genus Pilosphaera. In addition, Pilosphaera yentoensis n. sp. and Japonia boonkioensis n. sp. will also be described as new species. Members of Cyathopoma are tiny white cyclophorid snails occurring in East Asia, Madagascar and the Seychelles. Phylogenetic relationships of Cyathopoma are uncertain. Combined with COI and radular data, I conclude that Cyathopoma and Cyclotus are only distantly related. Cyathopoma iota has been considered to be a controversial member of this group. Through molecular and radular data, I found C. iota to be closer to C. taiwanicum than to C. micron, and concluded that C. micron, C. ogaitoi, C. iota and C. taiwanicum all belong to Cyathopoma. There are 10 genera and 29 cyclophorid species in Taiwan. Among them, the most interesting taxa are Cyclophorus and Cyclotus, both sharing similar ecological niches and representing by a north and south form in morphology. In order to clarify their relationship, I have to find out their sister group as out groups to compare with the members among Cyclophorus and Cyclotus. The gene trees of Cyclophoridae indicate that Japonia and Pterocyclus are sister group of Cyclophorus and Cyclotus, respectively. The former two will be used as the out groups of Cyclophorus and Cyclotus in their phylogenetic studies. Both COI and 16S rRNA gene trees of Taiwan Cyclophorus show prominent geographic structure. The Mantel test showed significant positive correlation between fixation index (FST) and cumulative geographic anti-clockwise distance (origin in the region around Tainan, anti-clockwise pass through south cape, Taidung, Hualian, Iran, Taipei, Taichung and meet the original populations in Jia-yi). There are finite gene flew between adjacent populations. And there are series of clines around the Central Range. Cyclophorus of Taiwan is a proposed “ring species”. In the morphology and environmental variables correlation study, I found the currently shell morphology may III be caused by the adaptations of recent long term climate. In traditional classification, Cyclotus taivanus consists of five subspecies, with clear morphological diversity. The molecular phylogenetic relationships of this group have never been discussed before. I sequenced part of the mitochondrial COI (cytochrome oxidase subunit I) and the 16S rRNA gene from 27 sampling sites. I also measured 9 shell traits to investigate the relationships between C. taivanus ssp. Even though the morphology PCA revealed a more or less continuous distribution of individuals in morph-space, the two highly divergent haplotype clades in COI and 16S rRNA analysis indicated the presence of two independently evolving lineages. The sequence divergence between two clades was almost as high as between other Cyclophoridae species. Therefore C. adamsi should be a considered valid species. For the environmental analysis, temperature may be a limiting element to the distribution of C. adamsi and C. taivanus group. The ecological divergence probably is the ruling force of speciation in my case. The PLS analysis results indicate, that phenotypic plasticity may be a key element of variable shell in C. taivanus group. The ecological divergence probably appears rule of speciation in C. taivanus ssp. case. The speciation process may be incomplete among C. t. dilatus, C. t. diminutus, C. t. peraffinis, and C. t. taivanus, and the adaptation of climatic pressure continue being a rule of speciation process. This study provides an opportunity to understand that no matter how similar two taxonomic groups are, occupying similar niche, undergoing the same geology history, with morphological adaptation to the same long term climate, they may have different speciation model. IV Chapter 1 General introduction Kobelt (1902) had been introduced cyclophorids in 1902, there were only 7 genera and 13 species in Taiwan. After Kobelt, Pilsbry and Hirase (1906) reported 7 genera and 21 cyclophorid species of Taiwan. There were 9 genera and 19 Taiwan cyclophorid species in Kuroda’s report of 1941 (Kuroda 1941). After these pioneers, Lee and Wu reported 9 genera and 29 species of Cyclophoridae in Taiwan (Lee & Wu, 2001). However, the phylogeny of cyclophorid is not clear. Besides, some new species and new records were found recently. Cyclophorids are very common snails in Taiwan, the genus Cyclophorus and Cyclotus are the most mysterious groups. The Cyclophorus moellendorffi found in southern Taiwan was considered a subspecies of C. formosaensis found in northern Taiwan by some authors (Kuroda, 1941; Chang, 1984; Lai, 1990; Higo & Goto, 1993). Besides, in the eastern side of Central Mountain Range, C. friesianus and C. latus were also believed to share a subspecies relationship by some authors (Kuroda, 1941; Chang, 1984; Lai, 1990; Higo & Goto, 1993; Hsieh et al., 2006). It is interesting that all the southern Cyclophorus species possess keeled shells, but the northern species possess round shells. Cyclotus taivanus ssp. are another interesting group. Pilsbry and Hirase (1905, 1906) reported C. t. peraffinis and C. t. adamsi as a subspecies of C. taivanus by their glossy surface and tall spire, respectively. Lee & Wu (2001) reported C. t. dilatus and Cyclotus t. diminutus as subspecies of C. taivanus by their wide out lip and very small shell, respectively. Like Cyclophorus, Cyclotus taivanus ssp. could divide north form (C. t. adamsi) and south from (C. t. dilatus, C. t. diminutus and C. t. peraffinis), roughly. The north form possess tall spire, the south form possess low spire. It is interesting that both Cyclophorus and Cyclotus taivanus ssp. occupy similar habitat, have two morphologic forms. Do Cyclophorus and Cyclotus taivanus ssp. undergo similar speciation process? The objectives of this study were (1) to establish the Molecular phylogeny of the family Cyclophoridae, (2) to understand the speciation model morphological adaptations of genus Cyclophorus, (3) to understand the speciation model morphological adaptations of genus Cyclotus, (4) to discuss the biogeography and morphology of Taiwan Cyclophoridae. 1 2 Chapter 2 Molecular phylogeny of the family Cyclophoridae (Gastropoda: Architaenioglossa) in East Asia Abstract Cyclophoridae consists of four subfamilies and about 300 species currently arranged in 38 genera, occupying varies habitats, with great morphological diversity. The molecular phylogenetic relationships of this group have never been discussed before. In order to investigate the relationships between cyclophorids, I sequenced part of the mitochondrial COI (cytochrome oxidase subunit I) and the 16S rRNA gene from 32 species of 10 genera of cyclophorid (including Cyclophoridae of China and Japan). I constructed phylogenetic trees using neighbor joining, Bayesian and maximum likelihood analyses on part of COI and 16S rRNA gene data set comprising. Phylogenetic relationships based on mtDNA sequences suggest that Cyclophorus, Cyclotus, Leptopoma, and Platyrhaphe are monophyletic while the traditional genus Japonia is polyphyletic and the previous J. zebra should be placed into a new genus Pilosphaera. In addition, Pilosphaera yentoensis n. sp. and Japonia boonkioensis n. sp. will also be described as new species. Besides, members of Cyathopoma are tiny white cyclophorid snails occurring in East Asia, Madagascar and the Seychelles. Phylogenetic relationships of Cyathopoma
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