Thesis submitted to 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 RAJEESH KUMAR M. P. Centre for Marine Living Resources & Ecology Ministry of Earth Sciences Kochi- 682037 JUNE 2018 Deep-sea Anglerfishes (Pisces- Lophiiformes) of the Indian EEZ: Systematics, Distribution and Biology Ph. D. Thesis in Marine Biology Author Rajeesh Kumar M. P. Centre for Marine Living Resources & Ecology Ministry of Earth Sciences, Government of India Block C, 6th Floor, Kendriya Bhavan, Kakkanad Kochi682037, 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 Kochi682037, Kerala, India Email: [email protected] June 2018 Front cover New species of Himantolophus (Ceratioidei: Himantolophidae) collected onboard FORV Sagar Sampada from Andaman Sea. Cover Design- Shebin Jawahar This is to certify that the thesis entitled ―Deep-sea Anglerfishes (Pisces- Lophiiformes) of the Indian E.EZ: Systematics, Distribution and Biology‖ is an authentic record of the research work carried out by Mr. Rajeesh Kumar M. P. (Reg. No.: 4323), 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 Kochi Kendriya Bhavan, Kakkanad June 2018 Kochi 682037, Kerala, India I hereby declare that the thesis entitled ―Deep-sea Anglerfishes (Pisces- Lophiiformes) of the Indian EEZ: Systematics, Distribution and Biology‖ 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. Kochi-16 Rajeesh Kumar M. P. June, 2018 I am greatly indebted to my supervising guide Dr. V. N. Sanjeevan (Former Director, CMLRE) for providing inspiration, for his timely guidance and support throughout my research. I express my deepest gratitude to Dr. M. Sudhakar (Director, CMLRE) for providing facilities and support. I also express my heart full thanks to all teachers in my student life for their support and blessings. This work was carried out as part of the project “Deep-Sea and Distant Water Fishery Resources of the Indian EEZ” under Marine Living Resources (MLR) Programme of the Ministry of Earth Sciences, Government of India, implemented by the Centre for Marine Living Resources & Ecology, Kochi. I express my gratitude to Dr. G. V. M. Gupta (Scientist F & Member-RC) and Dr. Hashim Manjebrayakath (Scientist D & Project Coordinator) for their encouragement and support. I thank Dr. Smitha B. R. (Scientist D, CMLRE) for her personal advice and coordination of PhD programme in CMLRE. I thank, Dr. A. Shivaji (Scientist F), Mr. N. Saravanane (Scientist E), Dr. Sherine S. Cubelio (Scientist D), Dr. Anil Kumar Vijayan (Scientist D), Mrs. Ashadevi C. R. (Scientist D), Mr. Telson Noronha (Scientist C), Mr. C. Vasu (Scientist C), Mr. M. Subramanian (Scientist C), Mr. Abdul Basheer (PA to Director), Mr. B. Kishore Kumar (RTO) and Mr. Sasi Irumbanam of CMLRE for all their co-operation during the period of work. I would like express my sincere gratitude to T.W. Pietsch, Professor Emeritus, School of Aquatic and Fishery Science, College of the Environment, University of Washington, Seattle, United States; J. H. Caruso, Senior Professor of the Practice Emeritus, Department of Ecology and Evolutionary Biology, Tulane University, New Orleans, United States and Hsuan-Ching Ho, Associate Professor, Graduate Institute of Marine Biology, National Dong Hwa University, Taiwan, for their scientific guidance, help rendered during taxonomic ambiguity and friendship. The support and cooperation from the FORV Sagar Sampada Vessel Management team at CMLRE led by Dr. A. Shivaji (Scientist E) as well as the Captain, Crew, Chief Scientists and scientific team of FORVSS Cruises are greatly acknowledged, as is the cooperation extended by the Fishing Master Mr. Thomas Paul and Fishing Hands, Mr. Tapan Kumar Malo, Mr. S. B. Prakash, Mr. Pradeep, Mr. Binoy V., Mr. Rathinavel and Mr. Sunil Kumar. I express my love and thanks towards the causal fishing hands, Mr. Umesh, Mr. Sasi, Mr. Aby and Mr. Lijo for their untamed help during onboard sampling. It is my pleasure to acknowledge the Engineers from NORINCO Pvt. Ltd. for their friendship and support. I greatly acknowledge the untamed support and friendships of my teammates Dr. Vinu Jacob (Sub Inspector, Fisheries, Govt. of Kerala), Dr. Aneesh Kumar K. V, Mr. Midhun Shah Hussein, Mr. Sumod K. S and Ms. Deepa K. P. I thank my colleagues at CMLRE Dr. Usha Parameswaran (Bhuvan), Dr. Aiswarya Gopal, Mr. Muhammed Rafeeq M., Ms. Meera K. M., Mr. Thomy R., Ms. Sreerenjima G., Mrs. Chippy Khader for their cordial friendship. I thank my seniors Dr. Jayalakshmi K. J., Dr. Abdul Jaleel, Dr. Padmakumar K. B. and Dr. Sabu P. for all their support during the period of this work. I am grateful to Mr. Maneesh T.P., Mr. Jensen Rozario, Mr. sileesh, Mr. Siby T. Baby, Ms. Harishree, Mr. Jijith S. S., Mr. Jijomon, Mr. Arun C. N., Mr. Kevin, Mrs. Sini Salam, Ms. Mrunal Wanjale., Mr. Abhay Deshmukh, Nikki. R., Oxona K., Krishna Kumar, Dr. Kusum Komal, Dr. Rajool Shanis, Mr. Raviranjan, Mr. Ruffus thella, Mr. Rajeev and Mr. Sailesh Yadav for their help and support. I sincerely acknowledge, Mr. Vinod (Agri. Officer, Govt. of Kerala) and Ms. Neeraja Prakash for their immense help and friendship. Many thanks to Mrs. Mini M. K., Mrs. Bijimol, Mrs. Sajitha and Ms. Mariyakutty P. J. for their curatorial assistance in the laboratory. Help rendered for molecular work and sequence analysis from Dr. KK Bineesh, Scientist D, Andaman & Nicobar Regional Centre (ANRC), ZSI and KA Sajeela (CMFRI) greatly acknowledged. Special thanks extended to KKB for the help and friendship. I express my love towards my family— Achan, Amma, Brothers, Sister, Niblings, my in laws, for their blessings and support. Finally special thanks to my wife Shebin, for her photo editing work, untamed support, inspiration and humor. I thank god for sending a little Angel to our life, towards the completion of my work. Rajeesh Kumar M. P. Chapter 1 GENERAL INTRODUCTION ............................................................ 01 - 18 1.1 Introduction .............................................................................. 01 1.2 Order LOPHIIFORMES ........................................................... 03 1.2.1 General Characteristics ........................................................... 05 1.2.2 Phylogenetic Relationships ..................................................... 07 1.2.3 Food and Feeding ................................................................... 09 1.2.4 Reproduction and Early Life History ..................................... 10 1.2.5 Sexual Dimorphism ............................................................... 11 1.2.6 Sexual Parasitism ................................................................... 11 1.2.7 Bioluminescence .................................................................... 12 1.2.8 Economic Importance ............................................................. 13 1.2.9 Geographic Distribution ......................................................... 14 1.2.10 Fossil Lophiiformes ................................................................ 14 1.3 Indian Lophiiformes: Relevance of the Study ........................... 15 1.4 Objectives ................................................................................ 17 1.5 Outline of the Thesis ................................................................ 17 Chapter 2 STUDY AREA, SAMPLING METHODOLOGY AND ANALYSIS ........................................................................................... 19 - 34 2.1 Study Area ............................................................................... 19 2.2 Sampling ................................................................................. 21 2.3 Taxonomic Analysis ................................................................. 27 2.4 Biomass Estimation .................................................................. 28 2.5 Length Weight Relationship, Food and Feeding, and Reproductive Biology ............................................................... 30 2.6 Otolith Morphology and Morphometrics ................................. 31 2.7 Ecomorphological Studies ........................................................ 31 2.8 Molecular Identification of Lophiiformes ................................ 32 2.9 Data Analysis ........................................................................... 33 Chapter 3 LOPHIIFORMES OF THE INDIAN EEZ: SYSTEMATICS AND DIVERSITY ................................................