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The Ecology of Chaetognatha in the Coastal Waters of Eastern Hong

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The Ecology of Chaetognatha in the Coastal Waters of Eastern Hong The Ecology of Chaetognatha in the Coastal Waters of Eastern Hong Kong TSE，Pan Thesis submitted as partial fulfillment of the requirctnents for the degree of Master .of Philosophy Biology ©The Chinese University of Hong Kong February 2007 The Chinese University of Hong Kong holds the copyright of this thesis. Any person(s) intending to use part or whole of the materials in the thesis in a proposed publication must seek copyright release from the Dean of the Graduate School. 統系‘書圓 |f 0 3 M 18 )i| university~ SYSTEMy^ Thesis Committee Professor Chu Ka Hou (Chair) Professor Wong Chong Kim (Thesis Supervisor) Professor Ang Put O Jr (Committee Member) Professor Cheung Siu Gin (External Examiner) Abstract Chaetognaths occur in almost all marine habitats, including estuaries, open oceans, tide pools, polar waters, marine caves, coastal lagoons, and the deep sea. Their biomass is estimated to be 10-30% of that of copepods in the world oceans. Chaetognaths are voracious predators and have a great significance in transferring energy from small zooplankton to higher trophic levels, including juvenile fish and squid. However, their functional role in various oceanic and coastal ecosystems is hardly studied. In this study, species composition, species diversity and seasonal abundance of chaetognaths were investigated in two hydrographically different areas in the eastern coast of Hong Kong. Tolo Harbour, located in the northeastern comer of Hong Kong, is semi- enclosed and poorly flushed bay with a long history of eutrophication. It opens into the Mirs Bay, which is exposed to water currents from the South China Sea. Zooplankton samples were collected monthly from July 2003 to July 2005 at six sampling stations. A total of 20 species of chaetognaths were identified. They included 6 species of the genus Aidanosagitta, 4 species of the genus Zonosagitta, 3 species of the genus Ferosagitta and 1 species each from the genera Serratosagitta, Decipisagitta, Flaccisagitta, Krohnitta, Mesosagitta, Pterosagitta and Sagitta. The three most abundant species, Flaccisagitta enflata, Aidanosagitta neglecta and Aidanosagitta delicata, comprised 39.7 %, 28.2 % and 22.0 %, respectively of the total chaetognath populations in all the samples collected throughout the study period. Tolo Harbour supported larger populations, but fewer species of chaetognaths than the surrounding open waters. These observations suggest that some species of chaetognaths were carried into the coastal waters of Hong Kong by water currents from the South China Sea and established large populations in the productive and poorly flushed waters of Tolo Harbour. I Diel vertical migration of Flaccisagitta enflata was studied in Tolo Harbour (8-9 July 2005 and 15-16 December 2005) and in Mirs Bay (8-9 September 2005 and 12-13 January 2006). Adult F. enflata carried out more active vertical migration than the juveniles in both Tolo Harbour and Mirs Bay. Food availability, pycnocline and hypoxia do not affect the vertical distribution of F. enflata. Both adult and juvenile F. enflata fed more actively at night. The predation impact of Flaccisagitta enflata on the copepods, cladocerans and larvaceans were compared between sampling stations inside Tolo Harbour and in the surrounding waters outside Tolo Harbour. Though copepods were the most common prey of Flaccisagitta enflata, cladocerans and larvaceans were more preferred prey. The predation impact of F, enflata on copepods (0-1.14 %) was negligible in the subtropical coastal waters of Hong Kong. In contrast, the predation impact on cladocerans (0-7.71 %) and larvaceans (0-6.19 %) and was more significant. II 摘要 毛顎類出現在幾乎所有的海洋生態之中，包括河口、開洋、潮塘、極地海域、海 洋石窟、沿海瀉湖和深海。其生物量預計相等於世界大洋中10-30 %的橈足類。毛顎 類是一種袍猛的捕獵者，它們在食物鏈中擔當著重要的角色’負責把能量從細小的浮 游動物之中傳送到轉高等的動物裹，例如魚和魷魚。不過現時對毛顎類在各大洋和沿 海生態所擔當角色，並沒有深入的硏究。 我對香港東面兩個擁有不同地理特性的沿岸水域進行了有關毛顎類品種組成，品 種多樣性及季節分佈的硏究，這兩個水域包括吐露港和大鵬灣。吐露港位於香港的東 北角，它是一個半封閉，水流不通暢和具有悠久營養化歷史的海灣。而大鵬灣是一個 暴露在南中國海水流中的海灣，它的西面連接著吐露港東面的出水口。從2003年7月 至2005年7月之間，我們每個月在六個採樣站裹分別收集一次浮游動物的樣品。綜合 所有的樣品，一共發現了 20種毛顎類，它們包括了六種的屬，四種 Zonosagitta ’ 三種 的屬禾口一種分另IJ來自 Serratosagitta ’ Decipisagitta， Flaccisagitta，Krohnitta，Mesosagitta，P/ero^sagZ/r^jt 和 的屬。其中三種最豐富的 品種包括了 Flaccisagitta enflata，Aidanosagitta neglecta 禾口 Aidanosagitta delicata�這三 種毛顎類分別佔據了所有毛顎類樣品中的39.7 % ’ 28.2 %和22.0 %。毛顎類在吐露港 的密度是高於周邊的水域，但其品種在吐露港是相對少於周邊的水域。這說明了在香 港的沿岸水域，有一部分毛顎類的品種是由南海水流所帶進來的，毛顎類在水流不通 暢和高度營養化的吐露港形成了一個高密度的群體。 我們對Flacc—tta enflata的晝夜垂直遷移在吐露港(2005年7月8日至9曰和 2005年12月15日至16曰）和大鵬灣(2005年9月8日至9日和2006年1月12日至13 日）各自進行了兩次硏究。成熟的F. 有明顯的晝夜垂直遷移，相反，幼年F. III 的晝夜垂直遷移並不明顯。無論是食物的充足度，由溫差和鹽度差別所形成的 密度躍層或者是水中的缺氧，都無法証明對F. �〃的晝夜垂直遷移有任何影響。在 夜間，成熟和幼年厂的進食量有明顯的增加。 這個硏究同時比較了 Flaccisagitta enflata在吐露港及附近水域之間對中型浮游生 物捕食的影響。雖然橈足類在的食物中佔了最大的比重，但厂更加喜 好水蛋類和被囊類作爲食物。F. 的捕食對橈足類(0-1，14 %)在亞熱帶沿岸水域 所造成的影響十分微小，相反，其捕食對水蛋類(0-7.71 %)和被囊類(0~6.19%)所造成 的影響比較顯著。 IV Acknowledgements I would like to express my deep gratitude to my supervisor, Prof. C. K. Wong, for his advice, encouragement and guidance during the past two years. I also would like to express my gratitude to Prof. K. H. Chu and Prof. Put. O. Jr. Ang, for their valuable suggestions. My gratitude is also extended to the external examiner Prof. S. G. Cheung of the City University of Hong Kong for reading and improving my thesis. I am deeply grateful to Prof. Q. C. Chen of the South China Sea Institute of Oceanology for helping with the identification of chaetognaths. I am indebted to the staff of the Simon F. S. Li Marine Science Laboratory for their help and support during the entire course of this study. Particular thanks to Mr. K. C. Cheung for providing technical assistance and Mr. Y. H. Yung for helping with the collection of samples. I would like to thank Ms. Eva Y. W. Yau, Ms. Alice S. Y. Hui, Ms. Alice A. Y. Lie, Ms. Vivian C. Y. Li, and Mr. Y. K. Kwok for their help in sample collection, sample analysis and carrying out experiments. Last but not least, I need to specially thank my family members for their love and support. V Table of contents Page Abstract (in English) I Abstract (in Chinese) III Acknowledgements V Table of contents VI List of tables XII List of figures XVI VI Chapter 1. General introduction 1 1.1. Chaetognatha 1 1.2. Morphology and anatomy 1 1.3. Food and feeding behaviour 3 1.4. Locomotion 3 1.5. Study location 3 1.6. Significance of study 4 1.61. Species composition and seasonal abundance 4 1.62. Diel vertical migration and diel feeding activity 5 1.63. Predation impact 5 1.7. Objectives 6 Chapter 2. Species composition and seasonal abundance of 7 Chaetognatha 2.1. Introduction 7 2.11. Species composition 7 2.12. Study sites 9 2.13. Objective 9 2.2. Materials and methods 11 2.21. Field sampling 11 2.22. Laboratory analysis 11 2.23. Data analysis 12 VII 2.3. Results 12 2.31. Temperature and salinity 12 2.32. Species composition 14 2.33. Dominant species 18 2.34. Species richness and species diversity 20 2.4. Discussion 23 2.41. Species composition 23 2.42. Flaccisagitta enflata 24 2.43. Aidanosagitta neglecta 25 2.44. Aidanosagitta delicata 26 2.45. Species richness and species diversity 26 2.46. Prey abundance 27 2.47. Spatial influences 27 2.5. Conclusion 28 Chapter 3. Diel vertical migration and feeding activity of Flaccisagitta 29 enflata 3.1. Introduction 29 3.11. Abundance 29 3.12. Diel vertical migration 29 3.12. Diel feeding activity 30 3.13. Study sites 30 3.2. Objectives 31 VIII 3.3. Materials and methods 31 3.31. Sample collection 31 3.32. Laboratory analysis 32 3.33. Data analysis 32 3.4. Results 33 3.41. Hydrography 33 3.42. Flaccisagitta enflata abundance 33 3.43. Diel vertical migration 38 3.44. Dietary composition 42 3.45. Prey abundance 47 3.46. Food containing ratio 51 3.5. Discussion 51 3.51. Flaccisagitta enflata abundance 51 3.52. Diel vertical migration 55 3.53. Diel feeding activity 57 3.6. Conclusion 58 Chapter 4. Predation impact of Flaccisagitta enflata on mesozooplankton 59 4.1. Introduction 59 4.11. Flaccisagitta enflata 59 4.12. Prey 59 4.13. Predation impact 60 4.14. Feeding rate and digestion time 60 IX 4.2. Objective 61 4.3. Materials and methods 61 4.31. Sample collection 61 4.32. Sample analysis 61 4.33. Measurement of digestion time 62 4.34. Data analysis 62 4.4. Results 64 4.41. Water temperature 64 4.42. Flaccisagitta enflata distribution 64 4.43. Prey distribution 66 4.44. Food containing ratio 66 4.45. Gut content 66 4.46. Prey selectivity 72 4.47. Predation impact 72 4.5. Discussion 77 4.51. Flaccisagitta enflata distribution 77 4.52. Food containing ratio 79 4.53. Gut content 80 4.54. Cannibalism 81 4.55. Prey selectivity 82 4.56. Digestion time 82 4.57. Predation impact 83 4.6. Conclusion 84 X Chapter 5. Conclusion 85 References 86 ‘ XI List of tables Tables Page Table 1. Occurrence (+) of chaetognath species in China seas. ++ 8 represents dominant chaetognath species. The results in this table based on Du et al (2003), Chen (1982) and the present study. Table 2. Average density and relative abundance of chaetognath species 15 at sampling stations inside (SI, S2 and S3) and outside (S4, S5 and S6) Tolo Harbour. The value at each sampling station represents the average over the entire study period from July 2003 to July 2005. Table 3. Correlation (rs) between density of dominant chaetognath 21 species and physical parameters (Spearman Rank Order Correlation), n is the number of samples collected from the entire study period from July 2003 to July 2005. Table 4. Species richness ⑶，species diversity (Shannon's diversity 22 index，H) and species evenness (Shannon's equitability, E) of Chaetognatha at sampling stations inside (SI, S2 and S3) and outside (S4, S5 and S6) Tolo Harbour over the entire study period from July 2003 to July 2005. Table 5. Density (mean 土 SD) of juvenile and adult Flaccisagitta enflata 37 in Tolo Harbour (S2) and Mirs Bay (S4).
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