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Zoological Survey of India MEMOIRS OF THE ZOOLOGICAL SURVEY OF INDIA Vol. 17 No. 2 MONOGRAPH ON NICHOLLSIA KASHIENSIS CHOPRA & TIWARI1950 (CRUSTACEA, ISOPODA, PHREATOICOIDEA, NICHOLLSIDAE) By LAKSHMAN PRASAD GUPTA Zoological Survey of India, Q.P.R.S., Patna-800016 Edited by the Director, Zoological Survey of India 1989 Copyright 1989, Government of India Published % June, 1989 PRICE: Inland : Rs. 100.00 Foreign j £ 10*00 or % 14*00 PRODUCTION t PUBLICATION UNIT, ZOOLOGICAL SURVEY OF INDIA, CALCUTTA. PRINTED AT THE BANl PRESS, 16, HEMENDRA SEN STREET, CALCUTTA-700 006, PRODUCED BY THE PUBLICATION DIVISION AND PUBLISHED BY THE DIRECTOR, ZOOLOGICAL SURVEY OF INDIA, CALCUTTA 700 020 MEMOIRS OF THE ZOOLOGICAL SURVEY OF INDIA Vol. 17 (2) 1989 Pages 1—160 CONTENTS PAGES INTRODUCTION ... i DISTRIBUTION ... 3 MATERIAL AND METHODS ... 3 GENERAL ECOLOGY ... 9 Biological community ... 10 Vertical migration ... 10 Parasites ... U Observations in the Aquarium ... 11 CLASSIFICATION ... 13 EXTERNAL MORPHOLOGY ... 15 BODY WALL AND MOULTING ... 34 THE MUSCULAR SYSTEM ... 38 FOOD, FEEDING AND DIGESTIVE SYSTEM 44 Food and feeding habit ... 45 Different modes of feeding ... 50 Alimentary canal ... 52 Foregut ... 53 Histology ... 54 Musculature of the foregut ... 55 Functions of Proventriculus (Stomach) 58 Digestive gland or hepatopancreas 60 Histology of the digestive gland ... 63 [ii] PAGES Intestine ... 66 Histology of Intestine ... 67 Musculature ... 68 Movement of food in alimentary canal 69 pH of the gut ... 70 EXCRETORY SYSTEM ... 71 Segmental glands of the head ... 72 The maxillary gland ... 73 Segmental glarids or coxal glands of the thorax 75 Histology of the maxillary gland ••• 75 Histology of other glands of head and thorax 76 Athrocytes or Branchio-pericardial organs 76 Tegumental glands ... 77 CIRCULATORY SYSTEM ... 77 Heart ••• 78 The Arterial System ••• 81 The Venous System ••• 83 The Blood - 85 Flow of blood - 86 Heartbeat ••• 87 Effect of body size ... 87 Effect of temperature ... 89 RESPIRATORY SYSTEM ••• 89 Morphology and histology of the pleopods 89 Movement of blood in pleopods ... 91 Pleopod beat ••• "1 Effect of temperature and size ... 91 Intestinal respiration ... 93 NERVOUS SYSTEM ••• 93 1. The Supra-oesophageal ganglia or the Brain 93 t iii i 2. Circum-oesophagea! connective 3. The suboesophageal ganglia The Ventral nerve cord Histology of the nervous system REPRODUCTIVE SYSTEM Male Reproductive system Histology of the testicular lobes Histology of the vasdeferens Histology of the penis The spermatozoa Orientation of spermatophores Androgenic gland Female Reproductive system Histology of the ovary Histology of the oviduct Brood pouch or brood chamber Pairing, Mating and egg deposition Copulation Transfer of eggs to the brood pouch Size of the Brood Incubation and Release of brood ... Reproductive cycle Sex Ratio Chromosomes DISCUSSION SUMMARY REFERENCES ACKNOWLEDGEMENTS LIST OF ABBREVIATIONS PLATES INTRODUCTION The discovery of a phreatoicid isopod in wells at Varanasi (Chopra, 1947) was hailed as an incident of great zoogeographical importance because, barring a couple of species of phreatoicids from the Cape Province of South Africa, the living species of the suborder phreatoi- coidea of isopods are now confined to the Australian continent, Tasmania and New Zealand. Described by Chopra and Tiwari (1950) as a new species under a new genus, Nichollsia kashiense was found to be common in deep wells at Varanasi, and some specimen were also available from a well at Lohagara near Naini in Allahabad District of Uttar Pradesh. A few year later, Tiwari (1955) described another species, Nichollsia menoni, from an abandoned well in Monghyr in Bihar State, extending the distribution of the genus eastward in the Gangetic Plain. Subsequent surveys have revealed that Nichollsia kashiensis occurs in several other localities in U. P. and Bihar, i.e., Ramnagar (opposite Varanasi on the right bank of Ganga), Chapra and Patna (Gupta, 1980). In a personal communication to Dr. K. K. Tiwari, Prof. P. J. Sanjeeva Raj of Madras Christian College, Tambaram (Madras), informs that he has material of phreatoicid isopods from Andhra Pradesh (locality not revealed) in South India obtained during deep drilling operations for boring tube wells. The phreatoicoidea is a primitive suborder of isopods which seems to have evolved during the palaeozoic in the old landmass known as Gondwanaland. Apart from Australia, Newzealand and Tasmania where several living species belonging to many genera still survive in surface as well as subterranean waters, recent representatives of this group have a discontinuous distribution in India, in the Gangetic Plain and Andhra Pradesh (Peninsular India), and in the Cape Province of South Africa. Outside the Australian continent, the only fossil representatives of this group have been reported from Siberia by Birs- tein (1962) who recorded Paleophreatoicus soyanensis from Permian beds. Judging from the present and past distribution, it appears that phreatoicoidea must have evolved during palaeozoic and had wide distribution in the old landmass of Gondwanaland, and has now disappeared from most of the Afro-Euro-Asian landmass, leaving living relicts in South Africa and India, and sizeable population inf Australasia. 2 Mem. zool, Surv, India Inspite of its zoogeographical interest and biological antiquity, prac­ tically no work has been done on the anatomy, physiology, histology etc. of the phreatoicids, though the group seems to have received con­ siderable attention from taxonomists. Nicholls (1943) in a monograph (in two parts) has thoroughly revised the group, giving detailed descrip­ tion of all the known species till then. Except for a brief account of the anatomy of Mesamphisopus capensis by Barnard (1927) and Golu- hotelson thomsoni by Engemann (1964), we have practically no informa­ tion about the internal morphology and other aspects of phreatoicoidea. The group shows considerable ecological diversity inhabiting several aquatic and semi-aquatic niches on surface and in subterranean habitats and its morphological characters show an interesting assem­ blage of primitive and specialised features. It was, therefore, consi­ dered desirable to have detailed and in depth investigation on at least one species of phreatoicid isopod to findout the similarities that the group shares with other isopods and differences that separate them from each other. Because of its easy availability and hardiness under captivity (individuals can survive for several months in water from ponds or well at room temperature and with very little care), Nicholhia kashiensis was selected for the purpose. The work is divided into a number of chapters, each dealing with one topic. The account begins with description of the external mor­ phology, ecology and observations on feeding habits, locomotion, respiration, etc. of specimens in captivity. In subsequent chapters each organ system is dealt with separately, and the material includes mor­ phology, gross structure and histology of each system. Where possible, experiments on live specimens to elucidate the process of ingestion of food, digestion, defecation etc., and of circulation of blood, palpitation of heart etc., were done. The result is a compre­ hensive account of the Indian species of phreatoicoidea, Nicholhia kashiensis. As such, it is the first work of its kind done on a represen­ tative of Isopoda, phreatoicoidea. Separated as they are by time span and geographic space from their relatives in Australisia and South Africa, the Indian representatives of suborder are naturally, widely, divergent from the main evolutionary branch of the group extant in Australia, New Zealand, etc. Comparison with the Australian and S. African kins brings out certain interesting morphological and anatomical similarities and differences, which show the common affinities and evolutionary divergence of the different GUPTA : Monograph on N, kashiensis 3 stems spatially separated from each other by discontinuous stretches of land and oceans. The details of similarities and divergences are discussed separately in each. While it is not possible to generalise on the basis of single species, it is hoped that similar studies will be undertaken on selected species of Phreatoicoid isopods from Australia and South Africa so as to provide comparative material that could throw light on the evolutionary history of the group and its relationship with other crustaceans. DISTRIBUTION So far, species of the genus Nichollsia have been recorded from a number of isolated localities in Uttar Pradesh and Bihar in the Gangetic Plains (Fig. 1). From Uttar Pradesh. N. kashiensis has been recorded from deep wells at Lohagara (near Naini in Allahabad District), Varanasi and Ramnagar (oppostive Varanasi on the right bank of Ganga). In Bihar, two species of this genus occur. N. kashiensis has been collected by the author from Chapra and Patna. Another species, Nichollsia menoni, has been described from Monghyr. It appears that diligent surveys in the Gangetic Plains are likely to reveal that this genus may be more widespread in the subterranean waters. Though no published report is now available, it appear that sub­ terranean Phreatoicoids are also available in Andhra Pradesh in South India (pers. commun., P. J. Sanjeeva Raj). MATERIAL AND METHODS Animals were collected from the wells with the help of plankton and ordinary nets which were dropped and left overnight and then taken out. For collection of bottom detritus an iron bucket full of water was dropped from
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