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研究生: 陳淑芳
Chen, Shu-Fang
論文名稱: 東亞特有八角金盤屬(五加科)之親緣地理學研究
Phylogeography of Fatsia (Araliaceae): an endemic genus in East Asia
指導教授: 蔣鎮宇
Chiang, Tzen-Yuh
學位類別: 碩士
Master
系所名稱: 生物科學與科技學院 - 生物學系
Department of Biology
論文出版年: 2004
畢業學年度: 92
語文別: 中文
論文頁數: 85
中文關鍵詞: 種化時間八角金盤親緣地理親緣網狀圖核醣體核酸ITS序列葉綠體DNA
外文關鍵詞: trnL-F, ITS, nested clade analysis, phylogeography, Araliaceae, Fatsia, divergence time
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  •   八角金盤屬(Fatsia)為五加科(Araliaceae)植物,特產於東亞。此屬植物全世界共三種,其中F. polycarpa 產於台灣全省海拔1,800~2,500 公尺之山地;F. japonica 產於日本本島及琉球島,通常生長在海岸附近之暖林帶;F. oligocarpella 為日本小笠原群島之特有種,生長於低海拔之闊葉林中,此種數量稀少,只侷限分佈於小笠原群島中的父島及母島,日本學者將其列為瀕臨絕種物。由於八角金盤屬為東亞特有植物,相當適合東亞親緣地理之研究。

      本研究選取八角金盤屬三種植物共36 個族群,114 個樣本,利用PCR 擴增八角金盤屬植物之葉綠體DNA 介於trnL 及trnF 基因間的noncoding spacer,及細胞核ITS片段進行定序,藉由其核酸變異來重建八角金盤屬之親緣地理,並估算族群內及族群間的分化情形。由葉綠體trnL-trnF 遺傳變異的偵測結果,在顯示八角金盤屬種間均擁有較低的遺傳歧異度,其主要原因為葉綠體基因片段仍處於lineage sorting 階段,各種間還擁有許多祖先多型(ancestral polymorphism),故未能提供足夠的訊息以進行親緣分析。細胞核ITS 的分析結果則呈現較多的親緣訊息,其種間Tajima’D 為顯著負值,顯示八角金盤屬植物為近代族群擴張之結果。Neighbor joining 樹狀圖分析顯現小笠原八角金盤與與台灣八角金盤各形成一monophyly,進一步以親緣網狀圖分析,得八角金盤屬之族群由日本本島向南方及東南方進行族群擴張。利用分子時鐘估算小笠原八角金盤與與台灣八角金盤之種化時間,分別是距今0.96-1.87 百萬年前及0.33-0.64 百萬年前所種化。由八角金盤之親緣地理分析結果,推論八角金盤的族群擴張歷史受限於冰河時期的環境氣候與地理障壁,形成現今的分佈模式,而成為各島與特有種。

      Fatsia, a genus of the Araliaceae, is endemic to the Archipelago of west Pacific Rim and Bonin Islands. These woody, perennial plants, with terminal inflorescence, are insect-pollinated. Drupaceous fruits attract birds for seed dispersal. Phylogeography of F. japonica (Japan), F. oligocarpella(Bonin), and F. polycarpa (Taiwan) was reconstructed based on nucleotide variation of cpDNA trnL-trnF intergenic spacer and nuclear ribosomal ITS region. ITS genealogy reveals a close phylogenic relationship between F. polycarpa and a haplotype of Okinawa, as well as between F. oligocarpella and a clade of Honshu and Ryukyus. Paraphyly of F. japonica indicates its ancestry. According to a molecular clock, the splitting of F. oligocarpella from F. japonica was dated to 0.96-1.87 million years before present, coinciding the divergence time for most endemic plants on the oceanic islands. Nevertheless, coalescence has not been attained at the cpDNA spacer within either species due to possession of shared ancestral polymorphisms. A very low level of genetic variation of cpDNA was detected in F. polycarpa , π = 0.00071, while genetic diversity was estimated at π = 0.00285 and π = 0.00282 in F. japonica and F. oligocarpella , respectively. Evidence of a close affinity to Oplopanax , a genus of North America and Japan, and natural habitats in mountainous areas in Taiwan suggests a likely temperate origin of Fatsia. Accordingly, F. polycarpa may have been founded in Taiwan via migrants from F. japonica , which possesses numerous genetic polymorphisms in both chloroplast and nuclear DNAs. This unusual, southward colonization was dated to 0.33-0.64 million years before present. In contrast to the low genetic variation of cpDNA, postglacial demographic expansion, as indicated by significantly negative Tajima’s D statistics, may have recovered the genetic diversity of the nrITS in F. polycarpa , which was estimated at π = 0.00327, a level comparable to that of F. japonica and F. oligocarpella. Very low levels of genetic differentiation were detected between populations within all taxa, although current, across-island gene flow is likely to be constrained. Phylogeographical analysis suggests that such low geographical structuring in Fatsia, like many plants of the archipelago, has been highly mediated by periodical glaciation.

    誌謝................................................................................................................... Ⅰ 中文摘要........................................................................................................... Ⅱ 英文摘要........................................................................................................... Ⅲ 目錄................................................................................................................... Ⅳ 表目錄............................................................................................................... Ⅶ 圖目錄............................................................................................................... Ⅷ 第壹章 緒言....................................................................................................... 1 ㄧ、研究背景理論.................................................................................... 1 (一) 島嶼生物地理學................................................................. 1 (二) 親緣地理學......................................................................... 4 (三) 分子標記之理論發展......................................................... 7 二、八角金盤之生物學研究.................................................................. 10 (一) 分類地位與形態特徵........................................................ 10 (二) 地理分布與生育地環境.................................................... 10 (三) 相關文獻整理.................................................................... 11 三、研究文獻回顧.................................................................................. 14 (一) 日本、台灣與小笠原群島之自然環境與地質歷史....... 14 (二) 東亞島弧地質歷史........................................................... 17 四、動機與目的...................................................................................... 19 第貳章 材料與方法......................................................................................... 20 一、研究材料.......................................................................................... 20 二、資料分析.......................................................................................... 24 第參章 結果..................................................................................................... 26 一、八角金盤屬之核苷酸鹼基組成分析.............................................. 26 (一) 葉綠體DNA trnL-trnF 之noncoding spacer 片段.......... 26 (二) 細胞核DNA ITS 之noncoding spacer 片段.................... 26 二、八角金盤屬之遺傳變異分析.......................................................... 28 (一) 葉綠體DNA 之遺傳變異................................................ 28 (二) 細胞核DNA 之遺傳變異................................................ 28 (三) 中性假說測試................................................................... 29 (四) 遺傳分化指數(FST)分析................................................... 29 三、親緣關係樹之構築.......................................................................... 31 (一) 傳統樹狀圖的建立........................................................... 31 (二) 網狀親緣樹狀圖的建立................................................... 31 第肆章 討論..................................................................................................... 33 ㄧ、Fatsia 之遺傳變異分析................................................................... 33 (一) 種間之遺傳變異探討....................................................... 33 (二) 種內之遺傳變異探討....................................................... 35 二、中性假說測驗.................................................................................. 36 三、族群分化之探討.............................................................................. 37 四、葉綠體 DNA 與細胞核DNA 之親緣樹狀圖結果比較............... 39 五、八角金盤之親緣地理...................................................................... 41 (一) 種化時間之推估............................................................... 41 (二) 物種傳播路徑之假說....................................................... 41 (三) 八角金盤屬的生物地理與種化假說............................... 44 第伍章 結論..................................................................................................... 46 第陸章 參考文獻............................................................................................. 47 附錄一............................................................................................................... 80

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