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Ecology of Calamus Sinicus

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Ecology of Calamus Sinicus ECOLOGY OF CALAMUS SINICUS (COPEPODA，CALANOIDA) IN OCEANS OF SOUTHERN CHINA by LEE KA LUN Thesis Submitted as Partial Fulfillment of the Requirements for the Degree of Master of Philosophy in Biology © The Chinese University of Hong Kong APRIL, 2003 系硬复® ！^ 0 6 丽 ” li Pv--— - ； UNIVERSITY /_舅 Contents Pages Abstract (in English) i Abstract (in Chinese) iv . Acknowledgements vi List of Figures vu List of Tables xv Chapter 1 General Introduction 1 Chapter 2 Seasonal population structure, life cycle and body allometry of the planktonic copepod Calanus sinicus 2.1 Literature Review ^ 2.1.1 Taxonomy of Calanidae 2.1.2 Geographical and seasonal distribution of Calanus sinicus 7 2.1.3 Body length and body weight of Calanus sinicus U 14 2.2 Introduction 2.3 Materials and Methods 17 17 2.3.1 Field sampling 2 3 2 Identification and enumeration of zooplankton 19 1 Q 2.3.3 Body length and weight of Calanus sinicus 上？ 2.4 Results 20 2.4.1 Temperature 2.4.2 Salinity 2.4.3 Ambient Chlorophyll a concentration 22 244 Seasonal occurrence and distribution of Calanus sinicus • • 〜 23 in northern Taiwan 2 4 5 Seasonal occurrence and distribution of Calanus sinicus •‘ 26 in Hong Kong 2.4.6 Life cycle of Calanus sinicus in northern Taiwan 27 2.4.7 Life cycle of Calanus sinicus in Hong Kong 28 2.4.8 Stage ratio index 2.4.9 Seasonal changes in biomass of Calarms sinicus 29 2.4.10 Seasonal changes in the abundances of other copepods in northern Taiwan 29 2.4.11 Seasonal changes in the abundances of other copepods in Hong Kong 30 2.4.12 Seasonal changes in biomass of other copepods 31 2.4.13 Seasonal variations in body size 31 2.4.14 Seasonal changes in sex composition in adults 32 2.5 Discussions 32 Chapter 3 Diel vertical migration and gut pigment rhythm of the planktonic copepod Calanus sinicus 3.1 Literature review ..87 3.1.1 Diel vertical migration of Calanus sinicus 87 3.1.2 Diel feeding rhythm of Calanus sinicus 91 3.1.3 Measurement of grazing rate 93 3.2 Introduction 96 3.3 Materials and Methods 98 3.3.1 Zooplankton sampling and physical parameters 98 3.3.2 Identification and enumeration 100 3.3.3 Gut pigment fluorescence 100 3.4 Results 101 3.4.1 Temperature and salinity 101 3.4.2 Ambient chlorophyll a concentration 102 3.4.3 Diel vertical migration 103 3.4.4 Gut pigment content 106 3.5 Discussion 107 Chapter 4 Use of molecular markers in population analysis of Calanus sinicus 4.1 Literature Review 134 4.2 Introduction 138 4.3 Materials and Methods 142 4.3.1 Collection, preservation, and identification of Calansii sinicus samples 142 4.3.2 DNA sequence determination for Calaniis sinicus 143 4.4 Results 1^4 4.5 Discussion 145 Chapter 5 Conclusion 150 Ecology of Calanus sinicus (Copepoda, Calanoida) in Coastal Oceans of Southern China by Lee Ka Lun Abstract Copepoda, with over 7,500 described species, are the largest class of small crustaceans. Calanus sinicus is a dominant copepod in the Yellow Sea, the East China Sea and the Sea of Japan. A major objective of this thesis is to examine the seasonal occurrence of C. sinicus in coastal oceans of southern China. Samples were collected at seven sampling stations in coastal water off northern Taiwan from November 2000 to March 2002. Outside Port Shelter, Hong Kong, samples were collected at five sampling stations from February 2001 to April 2002. Population density of C. sinicus was always higher in Taiwan than in Hong Kong. In Taiwan, numbers occurred throughout the year and were seasonally abundant. They were most common in winter months. In Hong Kong, a single peak in abundance occurred in March, 2001. Sex ratio among adults varied seasonally in Taiwan. There were usually more females than males. In general，the individuals collected i during the winter were larger than those collected during the summer. Diel vertical migration is a widespread phenomenon that is known to exist in almost all taxa of planktonic organisms. Because phytoplankton is most abundant near the surface, diel vertical migration is often associated with diel feeding rhythm. Diel vertical migration and diel feeding rhythm of Calarms sinicus were studied at two sites located near the northern tip of Taiwan in April and December 2001. The selective advantage of diel vertical migration in late copepodites and adult female of C. sinicus in April was the reduction of mortality by predation. In December, the copepods modified their migration behavior by remaining in the relatively food-rich surface waters when food availability is low. Several protocols based on molecular genetic characters have been successfully applied to marine zooplankton. This thesis also determined if Calarm sinicus populations in the oceans of southern China are derived from populations in the Yellow Sea and East China Sea, molecular approach will be used to examine intraspecific variations in the DNA sequences of C. sinicus specimens collected in coastal waters outside the Changjiang river mouth and off northern tip of Taiwan. The result showed that the Zhejiang-Fujian longshore currents carry C sinicus from the East China Sea into the South China Sea. ii 南中國沿岸海域大中華哲水蚤生態硏究 李嘉倫 摘要 橈足類是屬於甲穀門下的撓足類綱，已知曉的橈足類大約有七千五百種。 大中華哲水蚤是黃海，東中國海和日本海的優勢種類.本論文對南中國沿岸海 域的大中華哲水蚤的生態進行硏究。自2000年11月至2002年3月在台灣北 部沿岸海域及自2001年2月至2002年4月在香港牛尾海外邊的海域採樣站的結 果表明：大中華哲水蚤的在台灣北部的密度大于在香港的密度，在台灣北部是 全年性發生及具季節性豐度，尤在冬季常見，其在香港海域的紀錄只有2001年3 月。成年的大中華哲水蚤的性別比率同樣帶有季節性，但主要由雌性組成。 結果同時表明其在冬季的體長一般都大於夏季的體長。 曰周性垂直遷移現象常見於淡水及海水的浮游動物。由於浮游植物積 聚在水表層，所以，一般會進行日周性垂直遷移的浮游動物，通常是白天棲息在 較深的水層且不進食，晚上再游到水表層進行攝食行爲。本論文對大中華哲水 蛋在台灣北部沿岸海域的日周性垂直遷移現象和其攝食節律進行探討。在 2001年4月的結果表明，大中華哲水蚤的幼生期和成年雌性的垂直遷移是爲了 躲避以目視法捕食的敵人° 在2001年12月，浮游植物的供應量下降，大部份 的大中華哲水蚤停留在水表層尋找食物。 iv 由於不同的分子遺傳標記已成功應用在海洋的浮游動物，所以，本論文 同時利用分子標記以分析南中國沿岸的大中華哲水蚤是否從黃海和東中國海而 來。硏究利用在長江河口和台灣北部沿岸海域的大中華哲水蚤的標本進行 DNA序列分析。結果表明南中國沿岸的大中華哲水S是跟隨浙江和福建沿岸 流從黃海和東中國海而來。 X Acknowledgements I am most grateful to my advisor, Prof. Wong Chong Kim for his supervision and valuable comments during my two years of study. I am also indebted to my thesis committee members, Profs. Ang Put 0. Jr. and Chu Lee Man for their advice and help. It is also a pleasure to thank the helpful comments from Prof. Chu Ka Hou. Special regards are given to Prof. Hwang Jiang-Shiou of the National Taiwan Ocean University for his willingness to serve as the external examiner of this thesis and to provide the opportunity for me to work in Taiwan. Thanks are also given to Prof. Chen Qing Chao of the South China Sea Institute of Oceanology, Academia Science for his assistance in collecting zooplankton samples outside the Chang]iang River mouth. I greatly appreciate the assistance provided by the staff of Marine Science Laboratory and Department of Biology, the Chinese University of Hong Kong. I would like to thank Cheung Kwok Chu and Li Chi Pang for their technical advice during my laboratory work. I also thank Yung Yuk Hay for his help during the two years of field sampling in Port Shelter, Hong Kong. Thanks are extended to Wan Chak Lam and Wong Chun Kwan for their support and suggestions in the laboratory. Special thanks also given to all the staff and students at the National Taiwan Oceanic vi University who gave me assistance and helped me with the zooplankton samples collections in Taiwan. I must also thank all the members of my family here, especially my mother, for their support. Finally, a special debt of thanks is due my best friend, Kwan Ning Yan, because of her continuous encouragement and unconditional support during the past two years. vii List of Figures Fig. 1.1: (a) Sequential development of the apparent segmentation of Calarms sinicus throughout the copepodid stages, CI to CV, male and female, (b) The fifth pair of legs of swimming Calarms sinicus male and female. Fig. 2.1: Map of South China Sea and East China Sea showing location of 利 sampling stations outside Nuclear Power Plant I (NPPI) in coastal waters off the northern tip of Taiwan. Fig. 2.2: Map of Port Shelter, Hong Kong showing location of sampling stations. Fig. 2.3: Seasonal changes in (a) temperature (oC) and (b) salinity (O/QO) at 4j station B500 off northern Taiwan. Fig. 2.4: Seasonal changes in (a) temperature (oC) and (b) salinity (O/QO) at ^^ station B1000 off northern Taiwan. Fig. 2.5: Seasonal changes in (a) temperature (oC) and (b) salinity (O/QO) at ^^ station B2000 off northern Taiwan. Fig. 2.6: Seasonal changes in (a) temperature (OQ and (b) salinity (O/QO) at 46 station C500 off northern Taiwan. Fig. 2.7: Seasonal changes in (a) temperature (oC) and (b) salinity (O/QO) at ^^ station CI000 off northern Taiwan. Fig. 2.8: Seasonal changes in (a) temperature (oQ and (b) salinity (O/QO) at ^^ station C2000 off northern Taiwan. Fig. 2.9: Seasonal changes in (a) temperature (oC) and (b) salinity (O/QO) at ^^ station C5000 off northern Taiwan. Fig. 2.10: Seasonal changes in (a) temperature (oC) and (b) salinity (O/QO) at 冗 station I outside Port Shelter, Hong Kong. viii Fig. 2.11: Seasonal changes in (a) temperature (OQ and (b) salinity (O/QQ) at station II outside Port Shelter, Hong Kong. Fig. 2.12: Seasonal changes in (a) temperature (oC) and (b) salinity (o/oo) at station III outside Port Shelter, Hong Kong. Fig. 2.13: Seasonal changes in (a) temperature (oC) and (b) salinity (〇/oo) at 。 53 station IV outside Port Shelter, Hong Kong. Fig. 2.14: Seasonal changes in (a) temperature (oC) and (b) salinity (o/oo) at ^^ station V outside Port Shelter, Hong Kong. Fig. 2.15: Surface (white), middle (grey) and bottom (black) chlorophyll a ^^ concentration at different stations off northern Taiwan from November 2000 to March 2002. Fig. 2.16: Surface (white), middle (grey) and bottom (black) chlorophyll a 56 concentration at different stations outside Port Shelter, Hong Kong, from February 2001 to April 2002.
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