Thesis Submitted to the in Partial Fulfilment of the Requirements for The
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ECHINODERMS OF THE SOUTH EASTERN ARABIAN SEA: SYSTEMATICS & ECOLOGY Thesis submitted to the COCHIN UNIVERSITY OF SCIENCE AND TECHNOLOGY In partial fulfilment of the requirements for the award of the Degree of DOCTOR OF PHILOSOPHY in MARINE BIOLOGY under the FACULTY OF MARINE SCIENCES USHA V. PARAMESWARAN Centre for Marine Living Resources & Ecology Ministry of Earth Sciences Kochi 682037 SEPTEMBER 2016 Echinoderms of the south eastern Arabian Sea: Systematics & Ecology Ph. D. Thesis in Marine Biology Author Usha V. Parameswaran Centre for Marine Living Resources & Ecology Ministry of Earth Sciences, Government of India Block C, 6th Floor, Kendriya Bhavan, Kakkanad Kochi 682037, Kerala, India Email: [email protected] Supervising Guide Dr. V. N. Sanjeevan Former Director Centre for Marine Living Resources & Ecology Ministry of Earth Sciences, Government of India Block C, 6th Floor, Kendriya Bhavan, Kakkanad Kochi 682037, Kerala, India Email: [email protected] September 2016 Front Cover Brittle star Ophiothrix (Acanthophiothrix) purpurea, collected onboard FORV Sagar Sampada from the study area. CERTIFICATE This is to certify that the thesis entitled “Echinoderms of the south eastern Arabian Sea: Systematics & Ecology” is an authentic record of the research work carried out by Ms. Usha V. Parameswaran (Reg. No.: 4423), under my scientific supervision and guidance at the Centre for Marine Living Resources & Ecology (CMLRE), Kochi, in partial fulfilment of the requirements for award of the degree of Doctor of Philosophy of the Cochin University of Science & Technology and that no part thereof has been presented before for the award of any other degree, diploma or associateship in any University. Further certified that all relevant corrections and modifications suggested during the pre-synopsis seminar and recommended by the Doctoral Committee have been incorporated in the thesis. Dr. V. N. Sanjeevan Former Director Centre for Marine Living Resources & Ecology Ministry of Earth Sciences Kendriya Bhavan, Kakkanad Kochi 682037, Kerala, India Kochi September, 2016 DECLARATION I hereby declare that the thesis entitled “Echinoderms of the south eastern Arabian Sea: Systematics & Ecology” is an authentic record of research work conducted by me under the supervision of Dr. V. N. Sanjeevan, Former Director, Centre for Marine Living Resources & Ecology (CMLRE), Kochi and no part of it has been presented for any other degree or diploma in any University. Usha V. Parameswaran (Reg. No.: 4423) Kochi September, 2016 ACKNOWLEDGEMENTS My deepest gratitude goes first and foremost to Dr. V. N. Sanjeevan, my supervising guide and former Director, CMLRE for his scientific guidance and constant encouragement. The freedom of work accorded by him and his patient, timely, measured criticism has been invaluable to me in my research career. I also express my deepest gratitude to Dr. M. Sudhakar (Director, CMLRE) and Dr. P. Madeshwaran (Former Director, CMLRE) for providing facilities and for their encouragement. This work was carried out as part of the project ‘Marine Benthos of the Indian EEZ’ under the Marine Living Resources (MLR) Programme of the Ministry of Earth Sciences, Government of India, implemented at the Centre for Marine Living Resources & Ecology. I express my gratitude to Dr. G. V. M. Gupta (Scientist F), Dr. T. Shunmugaraj (Scientist F), Dr. A. Shivaji (Scientist E) and Dr. Anil Kumar Vijayan (Scientist D) who have coordinated the project during the period of this work (2008-2016). I also than Dr. Rosamma Philip (Head, Dept. of Marine Biology, Microbiology & Biochemistry, CUSAT) and Dr. R. Damodaran (Retd. Faculty of Marine Sciences, CUSAT) for granting access to samples collected through the MLR project ‘Marine benthos of the continental slope of the Arabian Sea & Bay of Bengal’ between 2003 and 2007. I acknowledge, with deepest gratitude, the scientific advice and unconditional support provided by Dr. Abdul Jaleel K. U. (Project Scientist, CMLRE) during the period of this work. I am also tremendously grateful to Dr. Smitha B. R. (Scientist C, CMLRE) for all the inspiration, encouragement, friendship and support. The scientific guidance and encouragement given by Dr. Sabine Stöhr (Swedish Museum of Natural History) is acknowledged with thanks. I thank Dr. Hashim M. (Scientist C), Mr. N. Saravanane (Scientist E), Dr. Sherine S. Cubelio (Scientist D), Mrs. Ashadevi C. R. (Scientist D), Mr. Telson Noronha (Scientist B), Mr. C. Vasu (Scientist B) and Mr. M. Subramanian (Scientist B) and Mr. B. Kishore Kumar (RTO) of CMLRE for their encouragement. I thank the current and former Dean and Director of the School of Marine Sciences, CUSAT and Registrar, CUSAT for their support. I also gratefully acknowledge the support and encouragement provided by my teachers, Prof. Rosamma Philip (HOD), Prof. A. V. Saramma, Prof. Babu Philip, Dr. A. A. Mohamed Hatha and Dr. Bijoy Nandan of the Department of Marine Biology, Microbiology & Biochemistry, CUSAT. The collection of samples was made possible by the continuous support and cooperation from the FORV Sagar Sampada Vessel Management team at CMLRE. I express sincere thanks to them as well as the Captain, Crew, Chief Scientists and scientific team of FORVSS Cruises. Most invaluable of all during sampling, was the unconditional cooperation extended by the Fishing Hands, Mr. Tapan Kumar Malo, Mr. S. B. Prakash, Mr. Pradeep, Mr. Binoy V., Mr. Rathinavel and Mr. Sunil Kumar. Analysis of sediment texture was carried out at the National Centre for Earth Science Studies (NCESS), Trivandrum, and I express sincere thanks to the Director (NCESS), as well as Dr. T. N. Prakash (Scientist), Dr. Reji Sreenivas (Scientist), Dr. Tiju Varghese, Ms. Praseetha and supporting staff of the Sedimentology Lab, NCESS for their help. I have been most fortunate to work in close association with Dr. Abdul Jaleel K. U., Ms. Aiswarya Gopal, Mr. Vinu Jacob, Ms. Jini Jacob and Ms. Shruthi Venugopal, who all share my passion for science and comedy. I thank them for the long-winding scientific discussions (or arguments), the unending laughter, which has made lab work a pure joy. Most of all, I thank them the help accorded to me in preparing this thesis and also for the constancy of their friendship. I am equally fortunate and grateful for the kindness and timely support extended to me by Ms. Nikitha S. Linda, Mr. Muhammed Rafeeq, Ms. Sreerenjima G., Dr. Priyaja P. and Mr. Kevin P. V. during the preparation of this thesis. I thank colleagues Ms. Meera K. M., Mr. Sumod K. S., Mr. Rajeesh Kumar M. P., Ms. Chippy Khader, Mr. Thomy R., Mr. Maneesh T. P., Dr. Lathika Cicily Thomas, Ms. Anu Shaji, Mr. Sudheesh Keloth, Ms. Deepa K. P., Mrs. Mini M. K. and all project staff at CMLRE for the help extended to me during the period of this work. Encouragement provided by Mr. Niyas M. A., Mr. Anil Kumar P. R., Dr. T. Ganesh, Dr. K. J. Jayalakshmi, Dr. Rajool Shanis and Dr. Sabu P. is gratefully acknowledged. During a critical juncture in my life, I benefitted from timely advice from Dr. M. Bhasi (SMS, CUSAT), who with a few kind words, inadvertently set me on the path to writing this thesis. I express my deepest gratitude to him. I sincerely thank my friends Ms. Gayathri B. Menon, Ms. Trupti Dabade, Ms. Annie John, Ms. Dhanya N., Dr. Kavya Nambiar, Mr. Arun Jacob, Ms. Racheal Chacko, Lt. Cdr. Vivek Kuriakose, Ms. Kamala Suryanarayanan and Ms. Megha Mangalam for all their encouragement and support. I also thank Ms. Supriya Rangarajan, Ms. Vaidehi Powani, Mr. Mohammed Azil and little Ms. Fathima for being sources joy. At the junction between ‘friends’ and ‘family’, I have a small but significant set of benevolent and enthusiastic supporters. Whether I refer to them ‘friends of the family’ or ‘families of friends’ is matter of mere technicality. I thank Dr. B. Rajendran, Mrs. Saraswati Rajendran and Ms. Meera Rajendran; Mr. Vaidyanathan P., Mrs. Lalitha Vaidyanathan, Dr. Ranjini Vaidyanathan and Mr. Raghav Vaidyanathan; Mr. K. K. Shenoy, Mrs. Purnima Shenoy, Ms. Radhika Shenoy and Ms. Renuka Shenoy; Dr. Abdul Jaleel K. U and Mrs. Suleikha; Mr. Abhilash K. S. and Dr. Vrinda Sukumaran for all the love and for being pillars of support to me and my family. In this regard, I also thank the Mr. Abraham Mathew, Mrs. Bindu Abraham, and last but not least Mr. Mahesh Gupta, Mr. Biju Neelambran, Mr. Anil Unnikrishnan, and all staff of RBG Commodities Ltd., Kochi. I am fortunate to have been raised by a group of people comprising of parents, grandparents, aunts and uncles. They have made me understand the value of hard work, honesty and commitment and have variously instilled in me a sense of insatiable inquisitiveness, of wonder and of humour. I sincerely thank them and my brother for making me who I am and for their ardent encouragement for all my pursuits. Usha For my father INDEX Acknowledgements List of tables List of figures List of acronyms & abbreviations 1 Introduction 1 1.1 General Introduction 1 1.2 The Echinoderms 4 1.2.1 Phylum Echinodermata: definition 4 1.2.2 General characteristics 4 1.2.3 Evolutionary position 7 1.2.4 Systematics & diversity 8 1.2.5 Ecological roles 19 1.2.6 Global diversity & distribution 21 1.3 Echinoderms of Indian waters: status of knowledge 23 1.4 Echinoderms of the south eastern Arabian Sea (SEAS): relevance of present study 27 1.5 Objectives 28 1.6 Outline of the thesis 29 2 Study area, sampling design & analysis 33 2.1 The south eastern Arabian Sea 33 2.1.1 Geomorphology & sediment nature 33 2.1.2 Hydrography