CHARACTERIZING COPEPODS AND ITS FOOD VARIABILITY IN THE ARABIAN SEA Thesis Submitted to the Goa University For the Degree of Doctor of Philosophy In Marine Sciences By Ms. Analiza Maria D’souza Under the guidance of Dr. Mangesh Gauns Department of Marine Science Goa University, Goa 403206 April 2018 CHARACTERIZING COPEPODS AND ITS FOOD VARIABILITY IN THE ARABIAN SEA Thesis Submitted to the Goa University For the Degree of Doctor of Philosophy In Marine Sciences BY Analiza Maria D’souza Under the guidance of Dr. Mangesh U. Gauns SCHOOL OF EARTH, OCEAN AND ATMOSPHERIC SCIENCES Goa University, Goa 403206 APRIL 2018 Dedicated to Saint Francis and to my beloved family Dedicated to Saint Francis and to my beloved family Declaration As required under the university ordinance OB9.9, I state that the present thesis entitled “Characterizing copepods and its food variability in the Arabian Sea” is my original contribution and the same has not been submitted on any previous occasion. To the best of my knowledge the present study is the first comprehensive work of its kind from the area mentioned. The literature related to the problem of investigation has been cited. Due acknowledgements have been made wherever facilities and suggestions have been availed of. Place: Date: (Analiza Maria D’souza) STATEMENT I hereby state that all necessary corrections/modifications as advised by the examiners for my Ph.D thesis entitled “Characterizing copepods and its food variability in the Arabian Sea” are incorporated. Analiza D’souza ACKNOWLEDGEMENTS “Sing the praises of the LORD, you His faithful people, praise his holy name (Psalms 30:4).” I am completely obliged to the ALMIGHTY Lord for fulfilling my voyage of the doctoral degree. The boundless faith in the holy trinity gave me all the courage to face the ups and downs in this journey. The blessings of Mother Mary, angels and saints helped me to be patient along my path. Many helping hands of God’s messengers joined on the way of completion of this ENDEAVOR and I would like to convey my gratitude individually to all of them. First of all, I would like to thank my Ph. D. guide Dr. Mangesh Gauns, Principal Scientist, CSIR- National Institute of Oceanography, Goa. I am very GRATEFUL to you sir for introducing me to this subject, and for being my best critic throughout the research. Thank you for trusting in me and giving me the freedom to work on the objectives individually. I sincerely thank Dr. C. U. Rivonker, Head, Department of Marine Science, for all the support and guidance during my tenure. I take this opportunity to thank the members of the FRC panel for their critical and valuable reviews, especially Dr. M. K. Janarthanam (Dean, Faculty of Life Sciences) my VC's nominee Dr. Sameer Damare. I also acknowledge CSIR for an award of Junior Research fellowship followed by Senior Research Fellowship that has helped me financially to carry out this research work. I take this opportunity to thank former directors, Dr. S. Shetye and Dr. SWA Naqvi and present director Dr. Sunil Kumar Singh for giving me an opportunity to be associated with this institute and for the necessary laboratory facilities. A word of thank to Drs. Siby Kurian, Hema Naik and Damodar Shenoy the coordinators of SIBER programme for all the support ,rendered during sampling and assessments during the tenure. Special thank to Dr. S. G. P. Matondkar for his timely motivation and encouragement. I am greatly indebted to Dr. Gauri Mahadik for her patience in guiding me with copepod taxonomy. The basics and the important keys of copepod taxonomy were introduced to me by Gauri, who was trained by Dr. Grazia Mazzocchi, S.Z.N., Italy. I extend my gratitude to Dr. Mazzochi for her tips on taxonomy through emails conversation during my early phase of career. I thank Dr. Sunita Mochemadkar for helping me in copepod taxonomy. On this research journey, I was fortunate to meet Dr. Karen Wishner at IMBER conference (2012) and Dr. Sharon Smith at IIOE conference (2015). Their simplicity and eagerness to discuss on copepods and different aspects of it was very enlightening. I am grateful to both of them for their motivating words and important literature. Likewise, fortunately, I was introduced to Dr. Gadi Padmavati by my guide, who eagerly helped and confirmed the copepod identifications done by me. Thank you Dr. Gadi. I would like to thank Mrs. Supriya Karapurkar for the assistance rendered during stable isotope analyses. I would like to extend my gratitude to Dr. Pratirupa Bardhan for scientific as well as the technical support rendered during analyses of stable isotope on copepod samples. I am very grateful to you Prati for always understanding my need and for standing by my side in all the scientific discussions including preparation of manuscript on stable isotope work. I would like to thank Dr. Rasiq and Mr. Chandrasekhar Rao for their technical help in HPLC analyses for gut pigments of copepods. I am greatly indebted to Dr. Suraksha Pednekar for her generous attitude of instant suggestions on data plotting and statistics packages and above all for her kind gesture to help anytime. She was a great company in my research career. I extend my gratitude to Dr. Sanitha Sivadas for teaching me the basic statistics and for scientific discussion on my data. A BIG thank you to Ms. Prachi Naik and Ms. Lakshmipriya for all the moral support and appreciations. Both of them have a keen eye for drafting articles and both were always eager to help me with corrections of any form of write- up. Thank you for being my good companions in a critical time and for the insightful scientific discussions. I would like to thank Dr. Maya for her guidance on stable isotope work. Maya, you have been a blessing to me many a time and always helped me wholehearted, thank you for your warm and helpful nature. I express my sincere thanks to Librarian and his staff, at NIO and Goa University for their efficient support in procuring the literature and maintaining the best of oceanography journals. I also acknowledge the help rendered by HRDG group and ship cell at NIO during my tenure. I thank in a special way to Drs. Suhas Shetye, Geeta Nagvekar, Shrikant Patil, Dinesh Velip, Mahesh Nashnodkar, Prachi Tirodkar, Mahesh Majik, Dheeraj, Gayatree Narvekar, and Anil Pratihary for their spontaneous advice and concern that was of great help during various stages of my PhD journey. Special thanks to Dr. Sai Elangovan for all the help rendered, particularly during printing of thesis. I also thank my colleagues cum friends, Valeta, Shweta, Sandesh, Priyabrata, Adarsh, Arati, Amara, Jesley, Richita, Fatima and Ritu for being the best companions in this field of research and for all the learning experiences. I had spent awesome times with you all in laboratory and on field trips and cruises. I thank my other labmates, Prajyoti, Mrunmayi, Sumant, Wassim, Bhagyshri, Ankita, Meenal, Nandadeep, Pooja, Jonathan, Sujata and all the members of Biogeochemistry for being around with a team spirit, helpful, and understanding way. Also, thanks to Dr. Ayaz, Mr. Ankush, Mr. Anand and Mr. Dalvi, for helping me during sampling onboard. Thank you to my friends who were related to me in my PhD tenure particularly Dr. Sangeeta, Dr. Asha, Dr. Mandar, Genevieve and Vidya. Thank you all. I am deeply indebted to my family, my father Mr. Anthony, my mother Mrs. Angelina and my sister Ms. Anushka, without their co-operation, prayers and constant support, both moral and financial; it would have been an unattainable target for me. My parents always gave me the freedom to achieve great heights in my educational career. It is for their upbringing that I can stand on my own today. I would like to extend my gratitude to our priest, Fr. Richard who always shared his graceful motivation that helped me to withstand hard times of my PhD. Here, I would like to mention my sincere thanks to my uncles, aunties, cousins and well-wishers, who were always concerned about my work and me. Thank you, everyone, for all the support and love. ANALIZA MARIA D’SOUZA Contents Page No. List of Tables III-IV List of Figures V-IX List of Plates X Chapter 1 INTRODUCTION 1.1 GENERAL INTRODUCTION 1-7 1.2 REVIEW OF LITERATURE 7-30 1.3 ZONATION IN THE MARINE WATER SYSTEM 30-34 1.4 STUDY AREA 34-38 1.5 OBJECTIVES 39-40 SPATIAL AND TEMPORAL VARIATIONS OF Chapter 2 COPEPODS IN THE ARABIAN SEA 2.1 INTRODUCTION 49-51 2.2 METHODOLOGY 51-53 2.3 RESULTS 53-67 2.4 DISCUSSION 94-115 GENDER AND SPECIES-WISE INFLUENCE ON ẟ13C AND ẟ15N ISOTOPES OF COPEPODS IN Chapter 3 THE ARABIAN SEA 3.1 INTRODUCTION 116-118 3.2 METHODOLOGY 119-120 3.3 RESULTS 120-128 3.4 DISCUSSION 141-159 I SPATIAL DISCREPANCY INISOTOPIC COMPOSITION OF CALANOIDA AND POECILOSTOMATOIDA IN THE ARABIAN Chapter 4 SEA 4.1 INTRODUCTION 160-161 4.2 METHODOLOGY 161-162 4.3 RESULTS 162-167 4.4 DISCUSSION 173-180 FEEDING OF COPEPODS IN THE ARABIAN SEA, AS DETERMINED BY GUT Chapter 5 FLUORESCENCE TECHNIQUE 5.1 INTRODUCTION 181-183 5.2 METHODOLOGY 183-186 5.3 RESULTS 186-194 5.4 DISCUSSION 208-215 SUMMARY 216-222 REFERENCES 223-273 PUBLICATIONS II LISTS OF TABLES Chapter 1 Page No. Table 1.1 Summary of gut fluorescence studies on copepods from different part of world 41-42 Table 1.2 Summary of ẟ13C and ẟ15N values for copepods and bulk zooplankton from different part of the world 43-46 Chapter 2 Table 2.1 Mesozooplankton sampling locations from different cruises 68 Table 2.2 Result of ANOVA comparing inter-annual and seasonal variations in copepod abundances at continental shelf (G5) 68 Table 2.3