臺灣博碩士論文加值系統

臺灣過去對於海膜藻科與那國藻屬的研究多為海藻相調查。劉（2006）雖曾探討過臺灣產海膜藻科與那國藻屬的分類問題，但其偏重分子序列分析。因此，在此研究中，我以臺灣產海膜藻科與那國藻屬為研究對象，透過分子序列分析與形態觀察，來探討與那國藻屬和蜈蚣藻屬內藻種的親源關係並釐清輔助細胞基團絲的發育模式，以期增進對臺灣產與那國藻屬的藻種多樣性和分佈的了解。
RbcL序列分析結果顯示臺灣產與那國藻屬有八個不同的分化群（＝藻種），其中有六種是世界新種（盾葉與那國藻、東方與那國藻、羽葉與那國藻、北方與那國藻、細葉與那國藻以及大葉與那國藻）。由於藻體外觀的相似，這六種在野外易被誤認為是已知的藻種，但若輔以內部營養結構特徵以及rbcL序列分析來進行比較，則可成功的區分這些藻種。
與那國藻屬的八個藻種依據外觀形態及分子親源關係探討發現：1）薄葉與那國藻和盾葉與那國藻於分子親源關係上互為姐妹種，但其外觀形態有明顯的差異。薄葉與那國藻的葉片為不規則二叉分枝且其葉片上半部會扭曲呈大花穗狀，而盾葉與那國藻的葉片偏羽狀分枝且呈盾狀。2）雖然東方與那國藻、臺灣與那國藻和大葉與那國藻彼此間的分子親源關係並不明顯，但此三種的藻體葉片皆具有不規則二叉分枝。東方與那國藻具匍匐生長的藻體且葉片分枝數最多，其藻體具濃密的葉片分枝。另一方面，臺灣與那國藻的葉片長且厚，分枝的間距較寬且葉片邊緣平整，其藻體具疏鬆的葉片分枝。大葉與那國藻可和其他二種區分開的特徵為具有較長的葉柄，葉片分枝數少、分枝寬度差異大且葉片邊緣不平整。3）羽葉與那國藻、北方與那國藻和細葉與那國藻於分子親源關係上互為姊妹種，而且具有相似的藻體外觀─羽狀分枝和長而明顯的葉片中肋。其中，北方與那國藻的藻體呈花穗狀，葉片最長且分枝寬而稀疏。另一方面，細葉與那國藻的葉柄最長，葉片末端細而尖銳，而羽葉與那國藻的葉片分枝雖密卻整齊規律。
另外在與那國藻屬囊果胚胎發育方面的觀察發現薄葉與那國藻、盾葉與那國藻以及北方與那國藻的輔助細胞基團只具有二條基團絲，且其形成方式與Lin et al.（2008）所描述有關東方蜈蚣藻的輔助細胞基團發育模式較為相像，但與Kawaguchi et al.（2004）所認為的與那國藻屬具有三至四條基團絲的輔助細胞基團有許多相異之處。Kawaguchi et al. 提出的與那國藻屬分類特徵也需要稍微修正，即與那國藻屬的輔助細胞接受由連接絲傳送過來的合子細胞核時，多數的基團絲細胞立即開始向兩側進行一至四回縱向細胞分裂產生絲狀或圓形細胞，但不一定會形成多串葡萄狀細胞群。
本研究在與那國藻屬姐妹屬─蜈蚣藻屬複合群的研究上，亦加入LSU來進一步探討“鋸齒藻屬”分化群和蜈蚣藻屬分化群的關係。RbcL與LSU序列合併的分析結果顯示“鋸齒藻屬”分化群與蜈蚣藻屬分化群的支持度皆有提高，而蜈蚣藻屬形成的單系群也得到較高支持度。因此，本研究的結果得出rbcL序列分析適用於屬內種間的親源關係分析，而以rbcL與LSU序列合併的分析，可較完滿解決蜈蚣藻屬複合群的系統分類問題。

The previous studies on the taxonomy of Yonagunia from Taiwan were mainly focused on the marine flora investigation. Liu (2006) had discussed the taxonomic problem of Yonagunia from Taiwan, but her study results were largely relying on rbcL sequence analyses. In this study I focused the genus Yonagunia from Taiwan to infer the phylogenetic relationships between Yonagunia and Grateloupia and to clarify the development of auxiliary cell ampullar filaments by molecular analyses and morphological observation to improve the species diversity and distribution of Yonagunia from Taiwan.
In this study, rbcL sequence analysis showed that there are eight species of Yonagunia present in Taiwan and its neighboring islands. Among the eight species, six are new species (Y. peltata sp. nov., Y. orientalis sp. nov., Y. pinnata sp. nov., Y. borealis sp. nov., Y. minima sp. nov. and Y. maxima sp. nov.). The new six species were normally placed under the species of Y. formosana or “Carpopeltis maillardii” due to their similar gross morphologies.
Among the eight species of Yonagunia identified in this study, they can be separated based on morphology of habit and molecular phylogeny. Y. tenuifolia and Y. peltata are sister clades (= species). Y. tenuifolia has irregularly dichotomous branched blades and the upper parts of blades are often strongly twisted, whereas Y. peltata has pinnate branched blades presenting as a shield. Although Y. orientalis, Y. formosana and Y. maxima have irregularly dichotomous branched blades, the thalli of Y. orientalis are creeping and have dense branched blades. The thalli of Y. formosana possess long and loose branched, thick blades, whereas those of Y. maxima are with the longest stipe and their branches have the great width. Y. pinnata, Y. borealis and Y. minima are closely related sister groups and share similar habits, i.e. pinnate branched blades with thickened rib. The thalli of Y. borealis possess long and wide loose branches, whereas those of Y. minima have the longest rib and the distal ends of branched blades are thin and tapering. On the other hand, the thalli of Y. pinnata possess dense but regular branched bladelets.
In this study, I found the auxiliary cell ampullae of Y. tenuifolia, Y. peltata and Y. borealis are composed of only two orders of unbranched ampullar filaments. Their development pattern is similar with Grateloupia orientalis what Lin et al. (2008) described, but is different with what Kawaguchi et al. (2004) described that auxiliary cell ampullae contain three to four orders of ampullar filaments. I also found that, after diploidization, most cells of ampullar filament divided laterally one to four times to produce filamentous or globular cells, but the newly formed filaments did not form grape-like, cellular clusters.
In this study, I also conducted a combination of rbcL and LSU sequence analysis for resolving the taxonomic problem of the Yonagunia/Grateloupia/Prionitis complex. The rbcL + LSU sequence analysis showed that “Prionitis” clade and Grateloupia clade received stronger supports than those derived from either rbcL sequence analysis or LSU sequence analysis along. The genus Grateloupia is shown to be monophyletic and received a strong support. In conclusion, rbcL sequence analysis is suitable for delineating the interspecific phylogenetic analysis within a genus, whereas a combination of rbcL and LSU sequence analysis resolved the generic relationships of the Yonagunia/Grateloupia/Prionitis complex better and is suitable for inferring the intergeneric relationships.

謝辭.......................................................i
中文摘要..................................................ii
英文摘要...................................................v
表目錄...................................................xii
圖目錄..................................................xiii
第一章、前言...............................................1
一、海膜藻科分類研究史及分類特徵...........................1
1.1海膜藻科分類研究史......................................1
1.2海膜藻科生活史及相關形態發育............................9
1.2.1海膜藻科生活史........................................9
1.2.2海膜藻科相關形態發育.................................12
1.3海膜藻科分類特徵.......................................13
二、臺灣海膜藻科分類研究史及分類問題......................14
1.1臺灣海膜藻科分類研究...................................14
1.2臺灣海膜藻科待解決之分類問題...........................17
三、研究動機及目的........................................18
第二章、材料及方法........................................22
2.1 藻種採集與保存........................................22
2.1.1 蠟葉標本製作流程....................................25
2.2 分子序列分析方法......................................25
2.2.1 去氧核醣核酸萃取（DNA Extraction）..................25
2.2.2 基因擴增（Gene Amplification）：聚合酶連鎖反應（Polymerase Chain Reaction；PCR）........................27
2.2.3定序反應及基因定序（Sequencing Reaction and Gene Sequencing）..............................................28
2.2.4 親源關係分析（Phylogenetic Analysis）...............29
2.2.5 海膜藻科rbcL和LSU序列間變異分析.....................30
2.3 形態觀察..............................................31
2.3.1 染劑配製............................................31
2.3.2 切片染色方法........................................32
第三章、結果..............................................34
3.1海膜藻科與那國藻屬與相關藻屬之分子序列分析.............34
3.1.1 RbcL序列分析........................................34
3.1.1.1親源關係分析.......................................34
3.1.1.2 海膜藻科與那國藻屬與相關藻屬之rbcL序列間變異分析..39
3.1.2 LSU序列分析.........................................40
3.1.2.1親源關係分析.......................................40
3.1.2.2海膜藻科與那國藻屬與相關藻屬之LSU序列間變異分析....43
3.1.3 RbcL + LSU序列之親源關係分析........................44
3.2形態觀察...............................................53
3.2.1 Yonagunia tenuifolia Kawaguchi et Masuda（薄葉與那國藻）......................................................55
3.2.2 Yonagunia peltata Y.-C. Chuang et S.-M. Lin sp. nov.（盾葉與那國藻）..........................................64
3.2.3 Yonagunia orientalis Y.-C. Chuang et S.-M. Lin sp. nov.（東方與那國藻）......................................73
3.2.4 Yonagunia formosana (Okamura) Kawaguchi et Masuda（臺灣與那國藻）..............................................82
3.2.5 Yonagunia pinnata Y.-C. Chuang et S.-M. Lin sp. nov.（羽葉與那國藻）..........................................92
3.2.6 Yonagunia borealis Y.-C. Chuang et S.-M. Lin sp. nov.（北方與那國藻）.........................................101
3.2.7 Yonagunia minima Y.-C. Chuang et S.-M. Lin sp. nov.（細葉與那國藻）.........................................110
3.2.8 Yonagunia maxima Y.-C. Chuang et S.-M. Lin sp. nov.（大葉與那國藻）.........................................119
第四章、討論與結論.......................................128
第五章、參考文獻.........................................135

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