臺灣博碩士論文加值系統

台灣產蜈蚣藻屬外形多變，由單一到頻繁茂盛的枝狀形，滑溜到皮質的葉片形，而且有些具有不規則的增生小枝或葉片。前人對於台灣海膜藻科的分類研究大多只有在外觀形態上的探討，容易造成同種異名或異種同名的誤判，因此本研究以囊果胚胎發育比較，配合rbcL序列分析來探討台灣產海膜藻科蜈蚣藻屬及相關藻屬種類的系統分類及親緣關係。RbcL序列分析結果指出台灣產蜈蚣藻屬具有七個種類，其中，有2種為前人鑑定有誤藻種，應為世界新種【Prionitis huangii sp. nov. （過去誤植為“G. okamurae”和G. “filicina”），P. taiwanensis sp. nov. （過去誤判為G. “carnosa”、G. “livida”和G. “sparsa”）】，一為G. ramossissima，有兩種為本研究首先發現的世界新種（G. gelidioides sp. nov. 和G. lewisii sp. nov.），另有兩種疑為新種（Grateloupia sp. 1和Prionitis sp. 1），但因採集的標本缺乏雌配子體，需行進ㄧ步研究確定。另外，台灣產蜈蚣藻屬加上來自世界各地的相近屬種由rbcL親緣分析結果顯示，可區分為兩個不同的次分化枝。ㄧ為以Grateloupia屬模式種（G. filicina）為主的次分化枝，其中包含的台灣種類有G. gelidioides sp. nov.、G. lewisii sp. nov.、G. ramossissima和Grateloupia sp. 1。另一為包含Prionitis屬（鋸齒藻屬）模式種（P. lanceolata）為主的次分化枝，包含的台灣種類有P. huangii sp. nov.、P. taiwanensis sp. nov.和Prionitis sp. 1。由台灣產蜈蚣藻屬囊果胚胎發育過程的研究結果，筆者認為果胞受精後輔助細胞基團發育成為果胞子體周圍的周生體形成模式可分為兩種：1）為受精作用後產孢絲開始分裂時，輔助細胞基團絲未有任何細胞分裂或只有少數幾個細胞行1次的細胞分裂，且周生體是由髓層細胞和內皮層細胞所形成，此種模式的種類有G. gelidioides sp. nov.、G. lewisii sp. nov.和G. ramossissima；2）為受精作用後產孢絲開始分裂時，輔助細胞基團絲的大部分細胞會進行4-6次細胞分裂，並配合髓層細胞和內皮層細胞形成果孢子體周圍的周生體，包含此種模式的種類有P. huangii sp.nov. 和P. taiwanensis sp.nov.。筆者認為此兩種不同的周生體發育模式可用來區別Grateloupia 屬（周生體是由髓層細胞和內皮層細胞所形成）和Priontis 屬（周生體是由輔助細胞基團絲行5-6次細胞分裂組成的）。此研究結果解決了單以藻種外形或序列分析無法解釋的Grateloupia與Prionitis的系統分類重大問題。

The external morphology of the species in the genus Grateloupia is very variable. The thalli are composed by single or few, soft blades to bush, rigid, terete branches, or by luxuriant to cartilaginous blades, or by from regular to irregular pinnate proliferations. The previous studies on the taxonomy of Grateloupia from Taiwan were mainly focused on the differences in the vegetative structures. As the results, many species were wrongly named synonymously or homonymously. In this study, I used two approaches (the cystocarp development and rbcL sequence analysis) to clarify the taxonomic problems occurred in the species of Grateloupia from Taiwan and to infer their phylogenetic relationships between the species from Taiwan and the other species of Grateloupia from around the world. In this study, the molecular analyses showed that there are seven species of Grateloupia present in the waters of Taiwan. Among the seven species I found, four are new species (G. gelidioides sp. nov., G. lewisii sp. nov., P. huangii sp. nov. and P. taiwanensis sp. nov.). Grateloupia gelidioides and G. lewisii are reported as new to science as the first time, whereas P. huangii was mis-identified as “G. okamurae” and G. “filicina” and P. taiwanensis was misapplied to G. “carnosa”、G. “livida” and G. “sparsa” before. The record of G. ramossissima is confirmed and another two species treated here as Grateloupia sp. 1 and Prionitis sp. 1 are also found for the first time. Because of lacking female specimens for a detailed study, the two unnamed species call for a further study. In addition, as shown in the phylogenetic tree inferred from the rbcL sequences of the species of Grateloupia from Taiwan and worldwide, they can be divided into two subclades. One is called as Grateloupia sensu stricto subclade, consisting of the generic type of Grateloupia, G. filicina from Italy, G. gelidioides, G. lewisii., G. ramossissima and Grateloupia sp. 1 from Taiwan, whereas the other is treated as Prionitis sensu stricto subclade, which includes the generic type of Prionitis, P. lanceolata from California, USA, and P. huangii, P. taiwanensis and Priontis sp. 1 from Taiwan. Furthermore, based on the observations of the cystocarp development, there are two types of the formation of the pericarp after fertilization: 1) the pericarp is not formed by the ampular filaments, which are undivided or little differentiated, but is generated by growth of filamentous, medullar and transformed inner cortical cells. The species possessing the first type of pericarp are G. gelidioides, G. lewisii and G. ramossissima; 2) the pericarp is formed by the divided, differentiated ampullar filaments after fertilization along with filamentous medullar and transformed inner cortical cells. This species having the second type of pericarp include P. huangii and P. taiwanensis. Therefore, I conclude that the behavior of the ampular filaments after fertilization can be used as important features for separating the species currently placed in the genus Grateloupia into two different two genera, Grateloupia and Prionitis, which is currently sunken under the genus Grateloupia, in addition to the architecture of the ampullar filaments. In the true species of Grateloupia, the pericarp is mainly formed by transformed inner cortical and filamentous medulla cells. Nevertheless, the pericarp in the true species of Prionitis is consisting of the differentiated ampullar filaments, transformed inner cortical cells and medulla filaments. As the results shown above, this research successfully resolved the long standing, taxonomic problem of the genera Grateloupia and Prionitis, which can not be fully explained by rbcL sequence analysis or vegetative, morphological studies, by carefully examining the cystocarp development.

第1章 前言……………………………………………………………………………1
1.1海膜藻目海膜藻科蜈蚣藻屬系統分類研究史…………………………………1
1.1.1海膜藻目的研究歷史…………………………….…………………………1
1.1.2海膜藻科的研究歷史……………………………….………………………5
1.1.3目前海膜藻科各屬的分類情形…………………….………………………6
1.1.4蜈蚣藻屬（Grateloupia）目前分類情形……………….……………………9
1.2 海膜藻科蜈蚣藻屬特徵及生活史………………………….…………………31
1.2.1 藻體形態…………………………..………………………………………31
1.2.2 生活史……………..………………………………………………………33
1.3 台灣海膜藻科蜈蚣藻屬系統分類學及其相關研究………………………….38
1.4 目前分子生物技術在海膜藻科蜈蚣藻屬研究上的應用…………………….40
第2章 本篇研究目的…………………………………………..……………………43
第3章 材料及方法………………………………………..…………………………44
3.1 藻種採集與保存………………………………………….……………………44
3.2 DNA分析………………………………………………………………………44
3.2.1 DNA萃取（DNA Extraction）……………………………………………44
3.2.2 基因擴增（Gene amplification）…………………………………………49
3.2.3 定序反應及基因定序（Sequencing Reaction and Sequencing）……..….50
3.2.4 親源關係分析（Phylogenetic analysis）…………………………………50
3.3 形態觀察. ……………………………………………………………………...52
3.3.1 染劑材料的準備…………………………………………………………..53
3.3.2 染色步驟…………………………………………………………………..54
第4章 結果…………………………………………………………………………..56
4.1 RbcL序列分析…………………………………………………………………56
4.2 形態觀察……………………………………………………………………….73
4.2.1 Grateloupia gelidioides sp. nov…………………………….….….……….75
4.2.2 Grateloupia lewisii sp. nov………………………………….…..…………85
4.2.3 Grateloupia ramossissima Okamura....…………………………………....95
4.2.4 Grateloupia sp. 1……………………………………………....................105
4.2.5 Prionitis huangii sp. nov.………………….............…………….….….....112
4.2.6 Prionitis taiwanensis sp. nov.…………………………………….…….…127
4.2.7 Prionitis sp. 1………………………………………………......…………142
第5章 討論…………………………………………………………………………149
第6章 結論……………………………………..…………………………………..160
第7章 引用文獻……………………………………………………………………161

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