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Enhydrina Schistosa (Daudin)*

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Enhydrina Schistosa (Daudin)* PLACENTATION IN THE SEA-SNAKE, ENHYDRINA SCHISTOSA (DAUDIN)* BY L. R. KASTURIRANGAN.~ Re~ived October 16, 1950 (Communicated by Dr. N. Kesava Panikkar, ~.A., D.SC., V.A.SC.) CONTENTS PAGE 1. INTRODUCTION ..... .. 1 2. ACKNOWLEDGMENTS .... 2 3. HISTORICAL RESUME .... 2 4. MATERIAL AND METHODS .. 3 5. THE REPRODUCTIVE SYSTEM .. 4 6. THE UTERINE CHAMBERS .. 7 7. THE ANATOMY OF THE BLASVOCYST 8 8. THE HISTOLOGY OF THE PLACENTA-- (a) The Euryplacental Region .... 12 (b) The Stenoplacental Region .: .. 17 9. DISCUSSION ...... .... 21 10. SUMMARY ...... .... 29 ll. REFERENCES ...... .... 30 12. EXPLANATION OF PLATES .. .... 31 INTRODUCTION Enhydrina schistosa is a sea-snake belonging to the family Hydrophida~ the members of which are predominantly oceanic in habit. As the open sea is not a habitat suited for eggs being laid out to hatch as on dry land, the sea-snakes have perforce to retain the eggs within the body till the young ones complete their development and are ready to be born. "Viviparity," in the broadest sense of the word, seems to be nearly universal in the family. As direct observation of the birth of young sea-snakes is possible only in marine aquaria, the phenomenon has been recorded only very rarely (Raj, 1926). It appears likely that Laticauda eolubrina and one or two other species of this genus which occur close to the shore have not progressed beyond the oviparous stage (Smedley, 1930, 1931; Smith, 1930), but the great majority of the species included in the family is either ovo-viviparous * Part of a thesis approved for the Degree of Master of Science of the University of Madras; published wah the permission of the Syndicate of the Umverslty. :~ Present address of the author: Central Marine Fisheries Reaearcb Station, West Hill, Calicut. m 1 2 L.R. Kasturirangan or truly viviparous. There is scope for wide variation among the species in the degree of adaptation in response to the new mode of reproduction, but the phenomenon appears to remain largely unexplored. The subject has been invested with interest from the fact that true allantoic placenta- tion has been described in several species of viviparous reptiles in the past sixty years, and these studies of reptilian placentation throw light on current theories on the mode of origin of the placenta in mammals. It seemed desirable, therefore, to investigate the adaptive condition in the commonest species of sea-snakes to begin with. Two readily availa- ble species at Madras, e.g., Enhydrina schistosa (Daudin) and Hydrophis cyanocinctus Daudin have been investigated. E. schistosa has not only true viviparity but also a distinct type of allantoplacenta, the characteristics of which are quite unlike those recorded by previous workers. The present account deals only with E. schistosa, the placenta of the other species being reserved for description in a separate communication. ACKNOWLEDGMENTS The work was carr:ed out in the University Zoological Research Laboratory, Madras, during the tenure of a Research Studentship in 1939- 1941. Grateful acknowledgment is made to the Syndicate of the Madras University for the award of the Studentship. I am indebted to Professor R. Gopala Aiyar, the then Director of the University Laboratory, for suggest- ing the problem, guiding the investigation at every stage of its progress, and for constant help and encouragement. It gives me pleasure to record here my sincere thanks to him. I am grateful to Dr. H. S. Rao and Dr. N. K. Panikkar for kindly going through the typescript and making valuable suggestions towards its improvement. HISTORICAL RESUME Since the publication in 1891 of the classical work of Giacomini on the development of Chalcides tridactylus (=-Seps chalcides) it has been known that a true allantoplacenta occurs in some of the Reptilia, doubtless in the course of the normal evolutionary process independently of the Mammalia. Giacomini described also the placenta of Chalcides ocellatus in 1906. Though Giacomini's original descriptions were not available to the present author the handsomely illustrated paper on the placenta of Chalcides tridactylus by Cate-H~edemaker (1933) was helpful in studying the placenta of E. schis- tosa. Later research in this field has been carried out mostly on species from Australia and New Zealand. The work of Flynn (1933) on Tiliqua Placenlalio~z #t the Sea-S~take, Enhydrina schistosa (Daudi~z) 3 scincoides, of Harrison and Weekes (1925) on Lygosoma entrecasteauxi, of Weekes (1927-I935) on several species of Lacertilia and two species of Ophidia and of Boyd (1942) on Hoplodactylus maeulatus may be mentioned here. Weekes's investigations have covered a wide range of species. Her final paper (Weekes, t935) reviews and discusses the results of ten years' work and provides a comprehensive summary, which should be of very great help to every investigator of reptilian placentation. Two terrestrial snakes, Denisonia superba and D. suta, also figure in Weekes's list. The present work is, however, the first record of allantoplacentation in the Hydrophidce. MATERIAL AND METHODS The material for this study, consisting of 10 embryos of Enhydrina schistosa,* was obtained from pregnant females during November and December 1939. A total of 55 specimens was dissected between August 1939 and September 1941 of which 20 were males, 26 were non-pregnant females and 9 were gravid females. The 2 earliest obtained embryos were 194ram. long and the remaining 8 ranged from 264 ram. to 286 mm. in length. The stages of development reached in the latter are practically identical and form the basis for the description of placentation given below. Bouin's fluid (aqueous) gave very good fixation. Zenker's fluid (with acetic acid) and Formol bichromate mixture were the other fixatives used. Dis- sections of the uterine chamber and the fcetal membranes were performed under a Zeiss Binocular microscope. Portions of the uterine wall and the allantoic membranes were separated from one another, stained in alum- carmine and mounted whole. It was not found necessary to cut microtome sections of the complete uterine chamber which was quite large in size (35 ram. across). The uterine wall, the chorio-allantoic membrane and the inner allantoic membrane remained adherent to one another in the form of a single sheet which was rolled up and sectioned. Other parts of the f~etal membranes and the yolksac were studied by sectioning suitable pieces. The pre-pregnant and post-pregnant oviducts of the females and the testes of the males were also sectioned to study the seasonal changes in the histo- logical characters. Sections cut varied from 5 to 10 microns in thickness. Heidenhain's Iron Hzematoxylin followed by Orange G gave very satis- factory results. Delafield's H~ematoxylin and Eosin, Mann's Methylblue Eosin and Mallory's triple stain were also occasionally used. * The genus Enhydrina includes only a single species for which three specific names have been proposed at different times. According to Smith (1926, 1943) E. schistosa has priority over E. valakadyn and E. bengalensis, which should be treated as synonyms. 4 L.R. Kasturirangan THE REPRODUCTIVE SYSTEM The ovaries are elongated bodies 50 to 100 mm. in length. They are ridge-like and hang from the dorsal wall of the body cavity immediately mesial to the oviducts (Text-Fig. 1). The cranial end of the ovary is at the same level as the tip of the oviducal funnel. The right ovary which is a little longer than the lcft, is placed a little farther forwards. The ovarian eggs are of all sizes, the largest observed being 15 to 20 mm. in length. They are not grouped together at the two ends of the ovary but are scattered irregu- larly. The full-grown egg about to be shed from the ovary was observed only once. It was 70 to 75 mm. long and extremely soft and flabby. The oviducal funnel on the same side appeared to be very large, the funnel mouth TExT-FIG. 1. Enhydrina schistosa, pregnant female with a single uterine chamber. a.e. alimentary canal, cloacal reg,on, o.f. oviducal funnel. d.a. dorsal aorta, p.c.v, postcaval veto. k. kidney, r.v. efferent renal vein. o. ovary, ut. a. uterine artery. o cd. oviduct, caudal tubular region, ut. c,h. uterine chamber. o, cr. oviduct, cranial tubular region, ut. v. uterine veins. Plcicentalion in lhe Sea-Snake, Enhydrina schistosa (Daudin) 5 being very close to the ripe egg and stretched wide open. Whether active movements of the funnel wall assist in the entry of the egg and its passage into the oviduct is difficult to determine. The unripe egg 20 ram. 1Ohg and the ripe egg 70 mm. long greatly differed in colour and consistency. The former was quite firm and pale ivory in coloar and the latter very flaccid and distinctly yellow in colour. The primary egg-membrane does not increase in thickness with the growth in size of the egg. While it gives a firm outline to the small volume of yolky material in the unripe egg, it is not strong enough to hold firmly the vastly greater amount of yolky material in the ripe egg. The change in colour from ivory to yellow might be due to the difference in composition of the yolk deposited in the later stages or perhaps to the greater percentage of water present. Corpora lutea were observed in the ovaries of all the pregnant snakes and of some of the non-pregnant ones. The yellow luteal bodies were usually large, oval in outline, 10 mm. long and 4 to 6 mm. broad, and had a median longitudinal furrow where the follicle had ruptured and healed. The oviducts are two tubular bodies placed alongside of the alimentary canal (Text-Fig.
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