臺灣博碩士論文加值系統

淡水生態系因人類活動影響而易受外來種入侵。外來種若對引入地區的生態或經濟造成影響則稱為入侵種。原生於亞洲大陸的淡水雙殼貝河殼菜蛤 Limnoperna fortunei (Dunker, 1856) 已先後入侵至香港、台灣、日本、阿根廷、烏拉圭、巴拉圭，巴西和玻利維亞。河殼菜蛤在台灣的確定族群，於1986、1990、2000年分別發現於台北縣新店溪直潭壩、南投縣日月潭和桃園縣石門水庫。為釐清台灣3個水系間河殼菜蛤的族群關係，本研究應用粒線體COI片段的501個核苷酸序列，分析台灣、日本和阿根廷6個水系共326隻個體，其A + T高達65.1 %，21個變異點均為同義突變，區分出17種單基因型。台灣 (Hd = 0.233, π = 0.000275) 和阿根廷 (Hd = 0.378, π = 0.000253) 的河殼菜蛤族群之遺傳多樣性偏低，且mismatch analysis呈現單一高峰，顯示該族群為單次引入，且有明顯奠基者效應；而日本的遺傳多樣性很高 (Hd = 0.895, π = 0.00097) 和2 ~ 3個高峰的mismatch distribution，推測是多次引入所造成。H1為3國家河殼菜蛤共有單基因型，推測H1可能為原生地廣泛分布的單基因型；三個國家各有其獨特的單基因型：台灣 (H2 ~ H4)、日本 (H5 ~ H15) 和阿根廷 (H16和H 17)，顯示3國家的河殼菜蛤均為不同來源。除了台灣和日本的河殼菜蛤族群有明顯分化的單基因型之外，大部分單基因型間僅相差一個鹼基變異，使NJ、MP、ML和MJ network的分析結果，無法釐清多數單基因型的親緣關係。此外，日本的河殼菜蛤入侵族群呈現高度遺傳多樣性，推測原生族群應有眾多高度分化單基因型共存。河殼菜蛤在台灣發現的3水系—新店溪、大漢溪和日月潭水系的族群遺傳多樣性均低；3個水系族群之間，至少共有2種單基因型 (H1>80 %和H2)，應為奠基者效應所造成。3水系間的平均遺傳距離 (0.259 % ~ 0.387 %) 和FST值 (0.032 ~ 0.0490)，顯示台灣3水系族群關係相近，推測可能是單次引入至某一水系再被引入至另外2水系，而河殼菜蛤於台灣水系間的擴散，可能透過工程船或休閒船引入至另一水域。

Freshwater ecosystems are prone to be invaded by alien species because of human activity. Invasive species are alien species that caused ecological and economical impact in introduced area. Limnoperna fortunei (Dunker, 1856) is a freshwater mussel that native to Asian continent, has invaded Hong Kong, Taiwan, Japan, Argentina, Uruguay, Paraguay, Brazil and Bolivia. L. fortunei in Taiwan were found in 1986, 1990 and 2000 at Chih-Tan Dam (Sindian River, Taipei County), Sun Moon Lake (Nantou County) and Shihmen Reservoir (Taoyuan County), respectively. 21 polymorphic sites (all synonymous mutation) revealed 17 haplotypes based on 501-bp of mitochondrial cytochrome c oxidase subunit I partial sequences from 326 individuals were applied to clarify the population relationships of L. fortunei among 3 drainages in Taiwan and other three sampling sites in Japan and Argentina. Population genetic diversity is extremely low in Taiwan (Hd = 0.233, π = 0.000275) and Argentina (Hd = 0.378, π = 0.000253) and with only one peak in mismatch distribution suggested strong founder effect, but were high and 2-3 peaks in Japan (Hd = 0.895, π = 0.00097). H1 (>80 % in Taiwan and Argentina) were the only shared haplotype among 3 countries, suggested that H1 might be widely distributed in native area. There were high ΦST = 0.382 and specific haplotypes in 3 countries (H2-H4 in Taiwan, H5-H15 in Japan, H16 & H17 in Argentina) suggested different population origin of introduced L. fortunei. Haplotype phylogeny trees were reconstructed by Neighbor-Joining, Maximum Parsimony, Maximum Likelihood and Median-Joining network methods were all showed low resolution because of most haplotypes were differed by one mutation between them. L. fortunei in Japan revealed high genetic diversity suggested numerous divergent haplotypes were distributed in native population. Genetic diversity of L. fortunei in Taiwan among Sindian River, Dahan River and Sun Moon Lake were low. 2 haplotypes, H1 and H2, were shared among 3 populations with frequency over 80% of highest haplotype H1, which probably caused by founder effect. Mean genetic distance (0.259 % ~ 0.387 %) and FST valvues (0.032 ~ 0.049) between 3 populations suggested closer population relationships, indicating the same origin of 3 populations that single introduction event happened in one drainage and then introduced into other two. The dispersal vectors of L. fortunei between drainages might be engineering ships or recreational boats in Taiwan.

誌謝 ………………………………………………………………………I
摘要 ………………………………………………………………………II
目錄 ………………………………………………………………………V
表目錄 ……………………………………………………………………VI
圖目錄 ……………………………………………………………………VII
壹、前言 …………………………………………………………………1
貳、文獻回顧 ……………………………………………………………5
參、材料與方法 …………………………………………………………11
肆、結果 …………………………………………………………………17
伍、討論 …………………………………………………………………23
陸、參考文獻 ……………………………………………………………32
表 …………………………………………………………………………42
圖 …………………………………………………………………………52
附錄 ………………………………………………………………………60

Table 1. Synonym and recorded locality of Limnoperna fortunei in Southeastern
Asia……………………………………………………………………... 42
Table 2. Sampling sites and numbers of sequencing individuals of Limnoperna
fortunei in Taiwan…………………………………………………. 43
Table 3. Sampling sites and numbers of sequencing individuals of Limnoperna
fortunei in China, Japan and Argentina…………………………………. 44
Table 4. Characteristics of mtDNA COI based on 501-bp sequences of
Limnoperna fortunei in 6 populations among Taiwan, Japan and
Argentina………………………………………………………………... 45
Table 5. Polymorphic sites and designated haplotypes based on mtDNA COI
501-bp sequences in Limnoperna fortunei…………………………….... 46
Table 6. Genetic diversity of mtDNA COI based on 501-bp sequences of
Limnoperna fortunei in 6 populations among Taiwan, Japan and
Argentina………………………………………………………………... 47
Table 7. Haplotypes distribution and frequency of Limnoperna fortunei in
Taiwan, Japan and Argentina…………………………………………… 48
Table 8. Pairwise genetic distance and FST of Limnoperna fortunei between
Taiwan, Japan and Argentina…………………………………………… 49
Table 9. Pairwise genetic distance and FST of Limnoperna fortunei between 6
drainage areas in Taiwan, Japan and Argentina………………………… 49
Table 10. Analysis of molecular variance among 6 drainage areas of Limnoperna
fortunei in Taiwan, Japan and Argentina………………………………...50
Table 11. Analysis of molecular variance among Sindian River, Dahan River and
Sun Moon Lake drainage areas of Limnoperna fortunei in Taiwan……..50
Table 12. Pairwise nucleotide differences and genetic distance of haplotypes of
Limnoperna fortunei……………………………………………………. 51

Figure 1. Sampling sites of Limnoperna fortunei in Sindian River, Dahan River
and Sun Moon Lake drainage area in Taiwan………………………….. 52
Figure 2. Sampling sites of Limnoperna fortunei in Taiwan, Japan and Argentina 53
Figure 3. Planktonic larvae of Limnoperna fortunei under microscope…………. 54
Figure 4. Relationships between genetic distance and ratio of transition and
transversion based on mtDNA COI 501-bp sequences of Limnoperna
fortunei…………………………………………………………………. 55
Figure 5. Haplotype frequency and distribution of Limnoperna fortunei in
Argentina, Japan and Taiwan…………………………………………... 56
Figure 6. Phylogenetic tree with outgroups based on mtDNA COI haplotypes of
Limnoperna fortunei using Neighbor-Joining, Maximum Parsimony,
Maximum Likelihood and Baysian method …………………………… 57
Figure 7. Meadian-Joining network of haplotypes of Limnoperna fortunei in
Taiwan, Japan and Argentina…………………………………………... 58
Figure 8. Mismatch distribution of Limnoperna fortunei among 6 drainage areas
in Taiwan, Japan and Argentina based on mtDNA COI 501-bp
sequences……………………………………………………………......59

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