本研究分析2007年6月至2021年11月間，以研究船「海研三號」和「新海研三號」的改良桁桿式蝦拖網採集於臺灣西部沿岸（苗栗至枋寮）的頭足類樣品，並同步蒐集水質資料。研究期間共捕獲頭足類6科13屬34分類物種（taxa）、1,545尾個體，前六優勢種分別為虎斑烏賊（Sepia pharaonis）、唇瓣烏賊（S. lycidas）、金烏賊（S. esculenta）、貝瑞氏四盤耳烏賊（Euprymna berryi）、日本暗耳烏賊（Inioteuthis japonica）和火槍魷（Loliolus beka），合計佔73.4%。臺灣西部沿海的頭足類群聚組成有顯著的地理分布差異（p＜0.05），可區分為以七股–茄萣為界的「南群」和「北群」，和水深約100 m以深的「深水群」。這些頭足類分布的空間變化可能與溫鹽適應、海流和棲地偏好有關。雖然頭足類的種類組成並無明顯的年間變化，但優勢種的時序排序有差異。過去「北群」的最優勢種由火槍魷轉變為唇瓣烏賊，過去「南群」前四優勢種中的越南烏賊（S. vietnamica）已大量減少，不再名列本研究中南群的前四優勢種。此外，優勢種除了唇瓣烏賊外，其他種類在後期（2016–2021年）的豐度皆較前期（2007–2010年）為低。頭足類豐度的時間序列變化可能與海溫上升及漁業資源變動有關。優勢的虎斑烏賊、唇瓣烏賊和金烏賊有季節遷移的特性，亦顯示臺灣西部沿岸水深75 m以淺為烏賊適合的產卵場和育幼場。

The R/V Ocean Researcher III and the R/V New Ocean Researcher III were used to simultaneously investigate the cephalopod resources and measure hydrological and water quality data along the coastal waters off western Taiwan. During the study period, a total of 1,545 individuals were captured, representing 34 taxa from 13 genera and 6 families of cephalopods. The top six dominant species were Sepia pharaonis, S. lycidas, S. esculenta, Euprymna berryi, Inioteuthis japonica, and Loliolus beka, accounting for 73.4% of the total catch. There were significant geographic differences in the cephalopod assemblages along the western Taiwan (p < 0.05), including the "N group" and "S group" delineated by the Chiku–Jiading area, as well as the "deep-water group " divided by the water depth of 100 m. The spatial variation of cephalopods may be influenced by environmental adaptation, ocean currents, and habitat preferences. There were no significant annual variations in species composition; however, the ranking of dominant species changed over time. The most dominant species in the "N group" had transitioned from L. beka to S. lycidas. Among the top four dominant species in the "S group," the S. vietnamica had experienced a significant reduction and was no longer included in the list of the top four dominant species in the present study. Except for S. lycidas, the abundance of other dominant species was lower in the later period (2016–2021) compared to the earlier period (2007–2010). The temporal variations in cephalopod abundance may be related to global warming and changes in fishery resources. The seasonal migration characteristics of the dominant species, such as S. pharaonis, S. lycidas, and S. esculenta, indicate that the coastal waters off western Taiwan with the water depth shallower than 75 m serve as suitable spawning and nursery grounds for cephalopods.

目　錄
論文審定書………………………………………………………………………………i
誌謝…………………………………………………………………...…………………ii
摘要………………………………………………………………………….....…….....iii
Abstract………………...…………………………………….....……..……...….…...…iv
目錄………………………………………………………………..……………….……v
圖次…………………………………………………………………………...…....….viii
表次……………………………………………………….………………….....……….x
附錄…………………………………………………………..……...………….………xi
第一章 前言………………………………………………..………………..………….1
1.1頭足類的多樣性…...……………………………………………………………1
1.2臺灣西部海域的水文環境………...……………………………………………1
1.3頭足類種類組成的時空變化………………………...……………...……….…2
1.3.1頭足類種類組成的空間差異………………………………………………2
1.3.1.1水深差異……………………………………………………………….2
1.3.1.2南北緯度差異………………………………………………………….3
1.3.2頭足類種類組成的十年差異………………………………………………3
1.4頭足類的分布與環境因子之關聯性………………………………...…………4
1.4.1鄰近海域研究………………………………………………………………4
1.4.2國內研究……………………………………………………………………5
1.5頭足類的遷移與棲地利用……...………………………………………………5
1.6研究目的………………………………………………………………………...6
第二章 材料與方法………………………………..………………………………...…7
2.1底拖調查與水質資料蒐集………………………...……………………………7
2.2樣本鑑定與生物學參數的量測…………………………...………..…………..7
2.3數據分析………………………………………….……...…………………..….8
2.3.1 α多樣性指數分析…………………………..……...………………………8
2.3.2群聚組成分析……………………………………....….………..………….8
2.3.2.1選擇群聚組成的固定因子………………………………...…………..8
2.3.2.2多變量分析………………………..………………………….......……9
2.3.3豐度的時間變化分析………………………………………...……...……10
2.3.4環境因子分析……………………………………………...…...…………10
2.3.5環境與生物因子分析……………………………………...…...…………10
2.3.6優勢種之生物學分析………………………………………...…..……….11
2.3.7數據分析軟體…………………………………………….…...…………..11
第三章 結果....…………………..………………………………....……….…………12
3.1臺灣西部沿岸之環境…………………………….………………...………….12
3.1.1底質環境……………………………………………….……...…………..12
3.1.2環境因子……………………..………………….…………...……………12
3.1.2.1水溫……………………………………...……………..….………….12
3.1.2.2鹽度……………………………………………………...……………13
3.1.2.3溶氧………………………………..……………………...…………..13
3.1.2.4葉綠素甲……………………………..…………………...…………..14
3.1.3環境因子之主成分分析…………………………………………………..14
3.2臺灣西部沿岸頭足類的物種多樣性……………………………...…………..14
3.3臺灣西部沿岸頭足類的生物量……………………………………………….15
3.4臺灣西部沿岸頭足類分布的水深變化……………………..……...…………15
3.5臺灣西部沿岸100 m以淺頭足類的空間分布……………………………….16
3.6臺灣西部沿岸頭足類豐度的年間變化…………………………...…………..17
3.7環境因子與頭足類分布之關聯性………………………………...…..………17
3.8優勢種與生活史策略……………………………….…………...…………….18
3.8.1虎斑烏賊（Sepia pharaonis）……………………….………...…………18
3.8.2唇瓣烏賊（Sepia lycidas）…………………………………...…………..19
3.8.3金烏賊（Sepia esculenta）………………………….....…………………19
第四章 討論………………………………..………………………………………….21
4.1臺灣西部沿岸底拖頭足類的多樣性……………………………...…………..21
4.2臺灣西部沿岸底拖頭足類分布的水深差異………………………………….22
4.3臺灣西部沿岸底拖頭足類分布的南北緯度差異…………………………….23
4.4臺灣西部沿岸底拖頭足類豐度的年間變化……………………...…………..24
4.5優勢種的生活史策略與棲地利用…………………………………………….25
4.5.1 虎斑烏賊（Sepia pharaonis）……………………….………………..25
4.5.2唇瓣烏賊（Sepia lycidas）………………………………...…………..26
4.5.3金烏賊（Sepia esculenta）……………………….…………...……….27
第五章 結論與建議………………………………………………….....……………..28
第六章 參考文獻…………………….…….…………..………….…...….…………..30
 
圖　次
圖1. 本研究於2007–2021年間，在臺灣西部沿岸海域的14個拖網測站地理位置圖。…………………………………………………………………………………….37
圖2. 2007–2021年間，本研究於各測站之底拖水深範圍盒形圖。……...…...…….38
圖3. 頭足類之不同類群的背外套膜長（Dorsal Mantle Length）量測方法示意圖。…………………………………………………………………………………….39
圖4. 2007–2021年間，臺灣西部沿岸各測站四季的底層（Bottom）與表層（Surface）之環境因子。…………………………………...………………..……40
圖5. 2016–2021年間，臺灣西部沿岸各測站四季之葉綠素甲濃度。…...…......…..41
圖6. 2007–2021年間，本研究於臺灣西部沿岸14個測站採樣之環境因子主成份分析（Principal component analysis, PCA）結果。………………………………42
圖7. 本研究於2007–2021年間，臺灣西部沿岸海域各測站之底拖網數與物種數累積曲線圖。……………………………………….………………………..……..43
圖8. 本研究各測站之物種多樣性指數。…………………………………….……...44
圖9. 2007–2021年間，本研究於七股（CG）與澎湖水道（PC）之頭足類物種組成集群分析結果。…………………………………………………….…….……...46
圖10. 2007–2021年間，本研究在七股（CG）與澎湖水道（PC）捕獲之頭足類於不同採樣水深的豐度分布圖。…………………………………….…….……...47
圖11. 本研究之100 m以淺頭足類前九優勢種與剩餘非優勢種（Other species）於各測站平均豐度圖。………………………………………………………..…...48
圖12. 2007–2021年間，臺灣西部沿岸水深100 m以淺頭足類物種組成之非度量多維度空間分析（nMDS）結果。……………………………………….………….49
圖13. 2007–2021年間，臺灣西部沿岸水深100 m以淺頭足類物種組成於各年份之集群分析結果。…………………………………………………………………….50
圖14. 2007–2021年間，臺灣西部沿岸水深100 m以淺頭足類物種組成於各月份之集群分析結果。………………………………………………………………….…51
圖15. 2007–2021年間，本研究捕獲之頭足類優勢種的平均豐度在前期（2007–2010年）與後期（2016–2021年）分布圖。….…………………..……....……52
圖16. 2007–2021年間，本研究於臺灣西部沿岸捕獲的前六優勢頭足類之豐度與底層水溫散佈圖。…………………………………………………..…….………..53
圖17. 2007–2021年間，本研究於臺灣西部沿岸捕獲的前六優勢頭足類之豐度與底層溶氧散佈圖。…………………………………….……………...…..….……..54
圖18. 2007–2021年間，本研究於臺灣西部沿岸捕獲的前六優勢頭足類之豐度與底層鹽度散佈圖。…………………………………………………..……………...55
圖19. 本研究捕獲的虎斑烏賊（Sepia pharaonis）（A）外套膜長-體重關係曲線，（B）不同性成熟階段之體長頻度圖。…………………….………………...…..56
圖20. 本研究捕獲的虎斑烏賊（Sepia pharaonis）各月份體長頻度圖。……...….57
圖21. 本研究捕獲的頭足類前六優勢種之北群（N）和南群（S）於不同月份之平均豐度堆疊圖。……………………………………………………………..…...58
圖22. 本研究捕獲的虎斑烏賊（Sepia pharaonis）於不同發育成熟度（Maturity）的月份深度分布圖。…………………………………………………………….59
圖23. 本研究捕獲的唇瓣烏賊（Sepia lycidas）（A）外套膜長-體重關係曲線，（B）不同性成熟階段之體長頻度圖。………………………………………….....60
圖24. 本研究捕獲的唇瓣烏賊（Sepia lycidas）各月份體長頻度分布圖。…..…...61
圖25. 本研究捕獲的唇瓣烏賊（Sepia lycidas）於不同發育成熟度（Maturity）的月份深度分布圖。……………………………………………………………….62
圖26. 本研究捕獲的金烏賊（Sepia esculenta）（A）外套膜長-體重關係曲線，（B）不同性成熟階段之體長頻度圖。………………………………..………..….63
圖27. 本研究捕獲的金烏賊（Sepia esculenta）各月份體長頻度分布圖。…...…..64
圖28. 本研究捕獲的金烏賊（Sepia esculenta）於不同發育成熟度（Maturity）的月份深度分布圖。…………………………………………………………….…65
圖29. 2007–2021年間，本研究採樣的四季底層水溫於前期（2007–2010年）和後期（2016–2021年）分布圖。………………...…………………………………66
 
表　次
表1. 2007–2021年間，本研究於各測站之網次數（n）、底拖水深（Trawl Depth, m）的最大值（Max）、最小值（Min）、中位數（Median）和平均值±標準差（Mean±SD）。…………………………………..……………………...….......…...67
表2. 2007–2021年間，本研究於臺灣西部沿岸14個測站四季的環境因子Kruskal-Wallis test結果。……………………………………...………….…….…….…...68
表3. 2007–2021年間，本研究於臺灣西部沿岸14個測站採樣之環境因子主成份分析（Principal component analysis, PCA）各軸排序結果。…………………….69
表4. 本研究各測站的頭足類平均豐度（ind./104 m2）與相對百分比（Relative abundance, RA）。…………………………………………..……….……………70
表5. 本研究各測站的頭足類平均生物量（g/104 m2）與相對百分比（Relative abundance, RA）。……………………………………………………….…….….72
表6. 2007–2021年間，臺灣西部沿岸水深100 m以淺頭足類之種類組成於南北群（Groups N&S）的相似度貢獻百分比（Similarity percentage, SIMPER）結果。…………………………………………………………………………………….74
表7. 2007–2021年間，本研究於臺灣西部沿岸海域捕獲的頭足類於「北群」（N）和「南群」（S）之物種平均豐度（ind./104 m2）與排序表。……………...…75
表8. 2007–2021年間，本研究頭足類前九優勢種於各年份之平均豐度（ind./104 m2）與Mann-Whitney U test分析結果。………….………………….….…..…….76
表9. 本研究捕獲的前六優勢種與環境因子之斯皮爾曼等級相關係數分析（Spearman’s rank correlation coefficient）結果。……………………………….77
表10. 2007–2021年間，臺灣西部沿岸苗栗至枋寮捕獲的虎斑烏賊（Sepia pharaonis）形質測量資料。……………………………...………………………….……78
表11. 2007–2021年間，臺灣西部沿岸苗栗至枋寮捕獲的唇瓣烏賊（Sepia lycidas）形質測量資料。…………………………………………………....….………...78
表12. 2007–2021年間，臺灣西部沿岸苗栗至枋寮捕獲的金烏賊（Sepia esculenta）形質測量資料。…………………………………………………….…………...78
表13. 本研究於臺灣西部沿岸海域捕獲的頭足類物種名錄與過去文獻之比較。.79
表14. 本研究與過去文獻的集群分析各分群物種貢獻度百分比之比較。…..…...82
附　錄
附圖1. 2007–2021年間，臺灣西部沿岸水深100 m以淺頭足類物種組成之集群分析結果。…………………………………………………………………..….……..83
附表1. 2007–2021年間，本研究於臺灣西部沿海之各航次採樣日期、採樣測站、測站水深範圍與底拖網數表。……………………………………..…………...84
附表2. 2007–2021年間，本研究於臺灣西部沿海14個測站各年四季採樣的底拖網數表。……………………………………………………………………….……86
附表3. 2007–2021年間，本研究的底層水溫（Temperature, °C）事後檢定Dunn’s test分析結果。……………………………...………….........……………..……..88
附表4. 2007–2021年間，本研究的底層水鹽度（Salinity, psu）事後檢定Dunn’s test分析結果。…………………………………………..…………………..……….91
附表5. 2007–2021年間，本研究的底層水溶氧（Dissolved oxygen, mg/l）事後檢定Dunn’s test分析結果。….…………………..…………………….……..……….94
附表6. 2007–2021年間，本研究的底層葉綠素甲（Chlorophyll-a, μg/L）事後檢定Dunn’s test分析結果。……………………………………….……..……..……..97
附表7. 2007–2021年間，本研究於臺灣西部沿海捕獲的頭足類物種名錄。….......99
附表8. 2007–2021年間，本研究於臺灣西部沿岸各測站捕獲的頭足類個體數表。…………………………………………………………………………………...101
圖版. 2007–2021年間，本研究捕獲的前六優勢物種樣本照。…………………...103

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