MARINE BIOLOGICAL ASSOCIATION OF INDIA MEMOIR I THE RIBBON-FISHES OF THE FAMILY TRICHIURIDAE OF INDIA BY P. S. B. R. JAMES Centre/ Marine Fisheries Research Institute Mandapam Camp 1967 MARINE BIOLOGICAL ASSOCIATION OF INDIA MARINE FISHERIES P. 0., MANDAPAM CAMP INDIA © Copyright 1967 MARINE BIOLOGICAL ASSOCIATION OF INDIA MANDAPAM CAMP Printed by A. J. Deshpande, at Western Printers & Printers, Bombay-13 and published by G. Luther, Managing Editor, Marine Biological Association of India, Mandapam Camp, Madras State, India. CONTENTS Foreword 111 Preface Introduction Part One SYSTEMATICS TAXONOMY OF THE INDIAN SPECIES OF TRICHIURIDAE 5 1. Key to Species of Indian Ribbon-Fishes 6 2. Genus Eupleurogrammus Gill 8 3. Genus Trichiurus Linnaeus 15 4. Genus Lepturacanthus Fowler 23 Part Two ANATOMY II. COMPARATIVE OSTEOLOGY WITH REMARKS ON THE PHYLOGENY OF RIBBON-FISHES 29 1. General Features 30 2. Neurocranium 32 3. Individual Bones of the Skull 34 4. Branchiocranium 40 5. Pectoral Girdle and Fin 48 6. Pelvic Girdle and Fin 50 7. Dorsal Fin 51 8. Anal Fin 52 9. Vertebral Column 53 10. Relationship between Subfamilies Trichiurinae and Lepidopodinae 57 11. Osteological key 57 12. Relationships of the Genera of the Family frichiuridae 58 lU. HYPEROSTOSIS 61 1. Observations 61 2. Discussion and Conclusions 63 IV. ALIMENTARY CANAL 06 1. Oral Armature 66 2. Gross Anatomy 71 3. Histology V. VASCULAR SYSTEM 77 1. Heart 77 2. Arterial System 77 3. Venous System 79 VI. NERVOUS SYSTEM 81 1. Brain 81 2. Cranial Nerves 81 VII. OTHER INTERNAL ORGANS 84 1. Auditory Organ 84 2. Air-bladder 84 3. Renal Organs 84 4. Gonads .. 85 VIII. PARASITES 86 (with special reference to Eupleurogrammus intermedius) IX. FOOD AND FEEDING HABITS .. 87 1. Qualitative and Quantitative Analysis 90 2. Variations in the Food 93 3. Food in Relation to Age 97 4. Selectivity of Feeding 103 5. Condition of Feed .. .. 109 6. Feeding Habits 113 7. Food of Species 116 119 8. Discussion 123 X. LENGTH-WEIGHT RELATIONSHIP 128 XI. RELATIVE CONDITION FACTOR 132 Xn. REPRODUCTION 1. Maturity 132 2. Spawning 137 3. Comparison of Spawning Habits of the four species of Ribbon-fishes from Indian waters 147 4. Relation between Size of Gonads and Size of Fish .. 151 5. Gonado-somatic Index 154 6. Size at First Maturity 157 7. Fecundity 163 8. Sex-ratio 168 XIII. LIFE-HISTORY 170 1. Eggs and Larvae 170 2. Description of the Young Stages .. 172 XIV. AGE AND GROWTH 176 1. Length Frequency Analysis 176 2. Growth Rate 181 3. Use of Skeletal Structures 182 4. Summary of all Evidence 188 5. Comparison of Species .. 188 6. Growth of Young Fish 189 XV. RACIAL INVESTIGATIONS 191 1. Relation between Standard Length and Snout-vent Length 193 2. Morphometric Characters 194 . 3. Comparison of Sexes 196 4. Comparison of Samples .. 201 XVI. DISTRIBUTION AND FISHERY ALONG THE INDIAN COAST 211 1. Distribution of Species along the Indian Coast 211 2. Fishing Methods and Sdasons along Palk Bay and the Gulf of Mannar 214 3. Particulars of Catch 217 218 4. Economic Importance 220 REFERENCES 227 ADDENDUM FOREWORD The Marine Biological Association of India founded in 1958 for the promotion of the cause of marine sciences in the Indian Region has been pub­ lishing regularly from 1959 its Journal which has already established itself as one of the leading scientific periodicals. In addition to the above, the Association has been holding symposia, the first one in January 1962 on Scombroid Fishes and the second in January 1965 on Crustac^. The Proceedings of these symposia issued already have become indispensable works of reference wherever researches on the concerned subjects are carried out. One of the objectives of the Association has been to bring out monographs and other comprehensive accounts on the fauna and flora of the seas around India and the present Memoir entitled "The Ribbon-fishes of the family Tri- chiuridae of India" by Dr. P.S.B.R. James is the first of this series. It is based on his painstaking study for over three years on this economically important family of fishes about which the knowledge we had previously was only of a desultory nature. In addition to his own observations, all available information on the subject has been brought together to form a comprehensive account—the first of its kind in this country on any family of fishes. Dr. James has endeavoured to make the account succinct and lucid. The biologists as well as the fishery scientists are expected to be benefited by this publication which is bound to serve as a valuable work of reference for a long time to come. Dr. James deserves to be congratulated for the preparation of the same and it is hoped that this pioneer attempt will be emulated by other fishery workers and research scholars and more accounts of this nature will be forthcoming in furtherance of our knowledge on our fish and fisheries. The issue of this Memoir is certainly another milestone in the activities of the Marine Biological Association of India. The fact that the Association could do this in addition to the publication of the Journal and the Proceedings of the Symposia without any aid from any organisation in this country is a most unique achievement and shows the sustained patronage it is receiving from right thinking scientists all over the world. The Association is releasing the Memoir on the Ribbon-fishes with the earnest hope that it will be well received and found useful by all fishery biologists. Marine Biological Association of India, Mandapam Camp, S. JONES May 8, 1967. President PREFACE Ribbon-fishes of the family Trichiuridae constitute one of the important commercial fisheries of India. There have been some earlier reports about their distribution, abundance and biological aspects but no attempt has been made towards a comprehensive study of this group of fishes. Four species of ribbon-fishes are now known to occur in the Indian Seas and while knowledge about their biology would be useful from the fishery point of view, it was also thought that a study of their systematic position and some aspects of anatomy would lead to a better understanding of the group as a whole. This account, which is mainly based on my thesis approved for the Doctor of Philosophy Degree in Zoology of the Banaras Hindu University, deals with the systematic position, some aspects of anatomy, biology and fishery of the ribbon-fishes from the Indian Seas. The analyses, interpretation of data and illustrations are ray own. I am deeply grateful to Dr. S. Jones under whose supervision and constant encouragement this work was carried out at the Central Marine Fisheries Research Institute, Mandapam Camp. 1 am also grateful to Dr. S. P. Ray- Chaudhuri for all the help received during the course of this work. For con­ structive criticism and valuable suggestions, I am thankful to Dr. Carl L. Hubbs. I am also thankful to Dr. C. V. Kulkarni for many valuable suggestions. I wish to record my sincere thanks to Dr. R. Raghu Prasad, Dr. R. V. Nair, Mr. K. Virabhadra Rao, Mr. S. K. Banerji and Dr. E. G. Silas who have greatly helped me in "various directions in this study. I am also thankful to my wife, Mrs. Indira V. James for all the assistance. Some of the photographs presented in this work were kindly taken by Dr. R. P. Varma and Mr. K. G. Nambiar to whom my thanks are due. To the crew of M.L. 'Sagitta' and M.L. 'Mathi', 1 am thankful for the help during my collection trips. For kindly making available some specimens of related fishes for my study, I am thankful to the authorities of the British Museum, U.S. National Museum and the Leiden Museum. This work was done partly during the tenure of a Senior Research Training Scholarship of the Ministry of Scientific Research and Cultural Affairs, Govern­ ment of India and partly under a Junior Fellowship of the Council of Scientific and Industrial Research, New Delhi. I am thankful to the authorities of these two organisations for the awards. Thanks arc due to Western Printers & Publishers, Bombay, for executing the printing of the memoir in such an excellent manner and in special to Mr. G. S. Pohekar, for all the interest taken in this connection. Central Marine Fisheries P.S.B.R. JAMES Research Institute, Mandapam Camp, July 24, 1965. INTRODUCTION Ribbon-fishes of the family Trichiuridae are widely distributed in the Indo-Pacific and the Atlantic and four of the species constitute an important commercial fishery at several places along the Indian coast. Various species of this family have been recorded by many earlier authors from different local­ ities with brief remarks on their commercial importance but the present knowledge about their biology in general is very meagre. To mention some earlier works from India, on one or more of the species, Venkataraman (1944) dealt with the feeding habits and Chidambaram and Venkataraman (1946) their natural history. Devanesan and Chidambaram (1948) briefly reported on their fishery in Madras State. Jacob (1949) gave some details of their bionomics and Chacko (1950) reported the occurrence of eggs and larvae of Trichiurus savala i= Lepturacanthus savala) from waters around Krusadai Island, in the Gulf of Mannar. Mahadevan (1950) described the alimentary canal of T. haumela (= T. lepiurus) and Prabhu (1950 and 1955) gave an account of its breeding habits and some aspects of the biology. Vijayaraghavan (1951) dealt with a qualitative and quantitative analysis of the stomach contents of T. haumela and T. savala. From outside India there have been several reports on these or related species of ribbon-fishes but in this connection the works of Delsman (1927) on the eggs and larvae of Trichiurus spp.