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Home , Papilla (fish anatomy), Reproductive system

Journal of the Indian Association 7 27,2000, 7-17

OBSERVATION ON THE URINOGENITAL PAPILLA AND IN SOME INDIAN GOBIIDS (: TELEOSTEI)

SAKTI HAR Department of Zoology, Ranwnanda College, Bishnupur, Bankura-722122. W. Bengal, India.

ABSTRACT

A thorough comparative study on the urinogenital papilla and sexual dimorphism has been made for the first time in both the of twelve Indian gobiids- Glassogobius giuris (Han1ilton); Acentrogobius cyanomos (Bleeker); Eleotriodes muralis (Valenciennes); Parapocryptes serperaster (Richardson); Apocryptes bato (Hamilton); Scartclaos viridis (Hamilton); Boleophthalmus boddarti (Pallas), Periophthalmus schlosseri (Pallas); P. koelreuteri (Pallas); Taenioides anguillaris (Linnaeus); T. buchanani (Day); Odontamblyopus rubicundus (Hamilton). The urinogenital papilla, originating as a free muscular from the ventral surface of the body-wall and shortly behind anus, is present in both the sexes. It is an important organ of primary recognition in all . In case of male the papilla is conical, broad at the base and in it is either flattened, distally truncated or bluntly rounded. The presence of permanent colour mark over the specific region of the body surface is another secondary sexual character in a few species. Besides, colouration may also be a nuptial secondary sex character developed in some during peak breeding season. The enlargement and colouration of the organ is subject to seasonal variations parallel to the seasonal gonadal cycle. The histological architecture of the papilla shows a high degree of cellular specialization and an interrelationship to the urinary and genital ducts. The functional efficacy and significance of the papilla in the breeding biology of these has been discussed. Keywords : Gobiidae, Sexual dimorphism, Urinogenital papilla Egami, 1960; Sinha, 1961; Kamalaveni, INTRODUCTION 1961 a; Sircar, 1970). Eggert (1931) in the The urinogenital papilla and sexual course of his description of the male dimorphism have been reported in fishes reproductive system of gobiiform and from time to time by different authors bleniiform fishes found errotile in (Eggert, 1931; Hora and Law, 1941; the urinogenital papilla of some of the Mookmjee, Mazumdar and Dasgupta 1941; gobies. Egami (1960) reported the presence Parkar, 1941; Weisel 1949; Braungart, of the urinogenital papillae and seasonal 1951; Lagler and chin, 1951; Clark, 1959; variations in their structure in both the 8 HAR

sexes of several species of Japanese gobies. Glossogobius giuris (Male) But these works have addressed only the Urinogenital papilla or "Papilla gross merphological aspects of the Urogenitalis" is a free muscular abruptly urinogenital papilla in gobiids. conical structure which arises from behind The nature ofthe urinogenital papilla, the anus and situated almost midventrally its seasonal variations parallel to those of and projected as an outgrowth infront of the gonadal activity, interrelationship the starting point of the anal . It is between the genital and urinary diets and visible from outside; if pressure is exerted sexual dimorphism have not been worked upon the ventral side ofthe anal opening in out on both the sexes in gobiids and hence, live specimen, the papilla will be erected. a comparative study on these aspects has The urinary and genital openings, though been undertaken for the first time on the very minute, lie separately at the distal Indian gobiids. bluntly pointed tip. The common and the common enters independently into the body of the MATERIAL AND METHODS urinogenital papilla and throughout the length of the papilla the two ducts remain Live specimens of different maturity distinctly separate. During breeding season stages of twelve gobiids were collected at the enlargement and the pigmentation of random from river, pond and muddy the papilla has been particularly observed estuaries of Bay of Bengal, Midnapur, West and in the non-breeding season the papilla Bengal during different seasons of the year. is reduced in size and devoid of Any sorts of variation either in structural pigmentation. Again from the preparatory details or in the seasonal or sexual colour period the papilla elongates gradually. The mark over the general body surface and anus and urinogenital papilla are distinct in these fishes was particularly from each other but at the base of the recorded. The papillae or both males and papilla it is confluent with the lower margin were dissected out and quickly fixed of the anus. The anal opening is circular in Bonin's fluid and Zanker's fluid. Serial and guarded by a sphincter (Text-fig.l). sections of the urinogenital papilla were cut at 6-Sp. thick and stained in Delafield's The histology (pl.l fig.l) of the haematoxylin, Iron Haematoxylin, and urinogenital papilla reveals peculiarities. counterstained with Eosin. Picroindigo The skin or the outer covering is made up carmine and Mallory's triple stain were of stratified squamous epithelium with also used to identify and study the many mucous glands and with a thin layer musculature and connective tissue elements of subcutaneous connective tissue. There is in the papilla. a thin basement membrane separating these two layers. Inner to the subcutaneous connective tissue layer, there lies the thick OBSERVATION longitudinal muscle bands with several blood sinuses. Inner to this there is an With relation to Urinogenital papilla OBSERVATION ON THE URINOGENITAL PAPILLA 9 equally thick circular muscle layer which visible to the naked eye. Unlike the male forms the inner core of the papilla. The tip of the flattened papilla is truncated. whole length ofthe papilla is traversed by The dorsal aspect ofthe papilla is medially the conduits ofthe accessory glands ofthe ridged but its ventral side is more or less testes, the spermatic duct and urinary duct. smooth. During the breeding season The two ducts are embedded at the centre enlargement ofthe papilla takes place and ofthe papilla. The space between the ducts i~ the off-season the papilla is indistinctly and the circular muscle layer is filled up by visible (Text-fig.2). the reticular connective tissue which is Histologically (Pl.fig.2) the urinogenital disposed at the lateral side only. At the papilla is to some extent different from that proximal end of the accessory glands of the of the male regarding disposition of different testes, some of the glandular follicles layers of muscles and structure of the become enlarged and form channels or urinary and genital ducts. The outer conduits, slowly converging to open into stratified squamous epithelium that forms the common spermatic duct. As the conduits the skin of the papilla is not provided with traverse the length of the papilla they mucous secreting glands and this layer is converge and open in to the main spermatic very thin. The subcutaneous connective duct either collectively or individually. The tissue is visibly distinct only at the lateral common duct, thus formed by the union of sides of the papilla. Unlike the male, the the conduits and the spermatic duct can be longitudinal muscles are absent or very named as spermigland duct. The inner few. The loose circular muscle layer forms wall of the spermatic duct is composed of the inner core of the papilla. The is short columner epithelial cells. The surrounded ventrally by reticular connective sperimigland duct runs below the urinary tissue. The urinary duct is oval and the duct which is composed of tall columner inner wall is covered with smooth cells. epithelial cells. The two ducts run, one Genital duct has infolded lumen and is above the other without union, up to the tip covered internally with short colmnner cells. of the urinogenital papilla. The lumen of the uninary duct is more The male urinogenital papilla of spacious. Like the male the two ducts Periophthabnus schlosseri, P. lwelreuteri, starting from the base to the tip of the Acentrogobius cyanomos shows the same papilla run separately without m1ion. morphological and histological pecularities, The u.g. papilla offemale P. lwelreuteri as that ofmale G. giuris. is exactly similar in structure to that of Glossogobius giuris (Female) female G. giuris.

The urinogenital papilla is a free Parapocryptes serperaster (Male) muscular, spically flattened structure which The urinogenital papilla or arises like that ofthe male from behind the pseudocopulatory organ is a conical, anus and situated almost mid ventrally muscular, elongated structure placed a11d projected as an out growth infront of behind that very close to the anus. The the starting point of the anal fin. It is also 10 HAR

sphincter muscles surrounding the anus anguillaris, T. buchanani, and the urinogenital papilla house a narrow Odontamblyopus rubicundus except in elliptical groove which may be called as having "integumentary " in them. "integumentary cloaca". During preparatmy Acentrogobius cyanomos (Female) period the papilla is comparatively short and white in colour. During the breeding The urinogenital papilla is an oval, season the pailla is exceedingly long, stout, brown, muscular structure placed inside a reddish in colour and hangs down from the rounded groove on the ventral body wall body wall. During the post-breeding season just behind the anus. The papilla begins to the size of the papilla is reduced to a mere enlarge and differentiate form the body projection which is white in colour but the wall and anus starting from the preparatary groove or depression surrounding the papilla period right up to the peak of the breeding and anus remains unchanged. The papilla, season. Due to the presence of a groove again, begins to enlarge at the onset of the encircling the proximal end of the genital breeding period. The urinary and genital papilla during the peak of the breeding passages remain separate at the tip of the season it may be well differentiated from papilla (Text-fig.5). the anus and its adjoining area. At the tip ofthe papilla two distinct apertures, urinary Histologically (Pl.l.fig.5) stratified and genital, open separately to the exterior squamous epithelium forms a thick outer (Text-fig.3). limit of the papilla. Subcutaneous connective tissue is very thin at the side. Histologically (Pl.l.fig.3) the The major portion ofthe papilla is occupied urinogenital papilla has much similarity by the circular muscle which lies beneath with that of male G. giuris in the disposition the subcutaneoues connective tissue. Inner of different muscle layers, inspite of the to this and adjoining the two ducts a bed of morphological differences between the two. loose connective tissue is prominent in the The stratified squamous epithelium, as mid ventral region. The longitudinal muscle usual forms the outer skin of the papilla are interposed between the circular muscle m1d this layer is provided with mucous and loose connective tissue layer. The secreting gland cells. Unlike G. giuris, the urinary duct is very spacious whereas the subcuta11eous conucetive tissue is very broad genital duct is rather narrow and is ventral and underneath this layer longitudinal to the urinary duct. The inner wall of both muscle layers are present. Inner to the the ducts are lined with tall and short longitudinal muscle layer, circular muscles columner epithelial cells. The two ducts form the inner core of the papilla. The traverse the urinogenital papilla and outer wall of the urinary and genital ducts ultimately open at the tip independently. are made up of a thin layer of reticular connective tissue. The inner wall ofthe two The morphology, histology and seasonal ducts is also lined with tall and short changes of the male urinogenital papilla columner epithelial cells. The lumen of the are more or less similar in Apoeryptes unirary duct situated dorsal to the oviduct bato, Boleophthalmus boddarti, Taenioides is more spacious and the two ducts run OBSERVATION ON THE URINOGENITAL PAPILLA I I separately through the papilla up to its tip of the papilla, longitudinal muscle bands to open to the exterior. being absent in this case. Instead of reticular connective tissue abounds in large The morphology, histology and seasonal number ofblood sinuses, a condition which changes of female urinogenital papilla are does not exist in any other reported found similar in Eleotriodes muralis, previously. The ducts, urinary and genital, Parapocryptes serperaster, Apocryptes are surrounded by well-developed circular bato, Boleop hthalmus boddarti, muscles. The two ducts remain separated Periophthalmus schlosseri, Taenioides althroughout loose connective tissue. anguillaris, T. buchanani, Odontamblyopus rubricundus. With relation to sexual dimorphism "Integumentary cloaca11 is found only in Sexual dimorphism is very well marked Parapocryptes serperaster. in all the twelve species of gobiids by the Eleotriodes muralis (Male) presence of two dissimilar types of urinogenital papilla in two sexes. In case of The urinogenital papilla is conical, male it is conical and in the female oval. muscular, elongated brown coloured Moreover, only in case of female structure and is situated behind the anal Acentrogobius cyanomos, a black spot is aperture. The proximal part ofthe papilla present at the base of the caudal peduncle is placed inside a round groove on the ventral and in the male it is wanting. The spot is body-wall and its dorsal tip is projected present throughout the year in the females. outside from the body-wall, during In the male Periophthalmus schlosseri, preparatory period. During breeding season the first has a black band, the urinogenital papilla is comparatively becoming cobalt externally and with a enlarged and looks like alonging appendage. scarlet edging whereas in the female the During the post breeding season the papilla first dorsal fin is moderatly developed unlike is reduced in length to an exceedingly male. The spot is seen in all the seasons of minute form. The urinary and genital the year. During the breeding season several apertures are not confluent but remain black patches have been found to be completely separated at the tip ofthe papilla developed on both the sides of truck and (Text-fig. 4). caudal in the female of Periophthalmus Histologically (Pl.l.fig.4) the outer schlosseri and in the male of P. koelreuteri. integument of the urinogenital papilla is Regarding other gobiids brighter colouration made up of stratified squamous epithelium is developed among the males during the which is separated from the subcutaneous peak ofthe breeding season. connective tissue by a thin basement · membrane. Inner to the subcutanous connective tissue, there is a circular DISCUSSION connective tissue lamina. Inner to this With relation to urinogenital papilla lamina, reticular connective tissue, givi~g spongy appearence, occupies a major portion Ofthe twelve species studied here, the 12 HAR

J-11"~~:..__-11' ----o.G.P,

TEK11'- fiG URi -t TEXT - f'IGURf • -4

TEXT~ nGURE • 5

~-11!4.;::---- c.o.d ·-=""'""-" o.. +-----U·1·P .------c.v.r 1;:""------C.ll.l" ,.,.-r---- b. m. ---~ ----:+<

URINO-GENITAL ORGANS

Text-figure 1 - Glossagabius, Text-Figures 2 G. giuris, Text-Figures 3- Acentrogobius cyanomos, Text-Figure 4 - Eleotriods muralis, Text-figure 5- Parapoeryptes serperaster k- ; s.d- duct; g- gut; S. V- Seminal vessicle; ug.p. Urinogenital papilla, u.bl. - ; c.s.d. -Common sperm duct; a. - Anua; Lcl- Integumentary cloaca; c.ur - C01nmon ureter; c.o.d. Common Oviduct; b.m - Body muscle; b.w. Body wall. p.sv Posterior Seminar vessicle; a.s.v. -Anterior seminal vessicle; OBSERVATION ON THE URINOGENITAL PAPILLA 13

s. Plate 1 Transverse section of urinogenital papillae of Glossogobius giuris, Acentrogobius cyanosmos; Eleolriodes mural is and Parapocryptes serperaster. Fig. I: Transverse section through the male urinogenital papilla ofGlossogobius giuris. Fig.2: Transverse section through the female urinogenital papilla ofGlossogobius giuris. Fig.3: Transverse section through the female urinogenital papilla ofAcentrogobius cyanosmos. Fig.4: Transverse section through the male urinogenital papilla ofEleothriodes muralis. Fig.5: Transverse section through the male urinogenital papilla of Parapocryptes serperaster. ABBREVIATIONS bs.mb. Bas.ement rnemb.er9-nce co.ce., Columber epithelial cells., cr.m. - Circular muscles; fe.t. - Fibrous connective tissue; l.m.- Longitudinal muscles; Sc.c.t.- Subcutaneous connective tissue; Sd­ Spermduct; st.sq.c- startified squamous epithelium; u.d.- uninany duct; mu.gl. mucous glands; ov. Oviduct. r.c. t. Reticular Connective tissue: Cr. l. - Circular lamina. 14 HAR urinogenital papilla has been found in both author. Weisel (1949) has mentioned the the sexes. In all the species the papilla is occurence of the urinogenital papilla in free, muscular organ which is visible to the Gillichthys mirabilis. He noticed variations naked eye and can easily be raised with in the size of the papilla in different seasons needle. In case of male the papilla is conical, of the year but he could not correlate it board at the base and tapering posteriorly. with the sexual cycle of the fish. Braungart Similarly in female the papilla is either (1951) made a comparative study of the flattened, distally truncated structure as reproductive system of several fishes. In Glossocgobius giuris and He observed the presence of a urinogenital Periophthalmus lwelreuteri or bluntly papilla in black crappie, Pomoxis rounded in the rest of the species, studied nigromaculatus. In this fish both the here. It arises from the ventral surface of urinary and genital ducts open to the the body-wall shortly behind the anus. The outside of the body separately through urinary and genital apertures open to the separate orifices, although both open to the exterior individually, in both the sexes, at outside on the same papilla. The present the tip ofthe papilla. The enlargement and study corroborates with Braungart's colouration of the organ is subject to findings so far as the opening of the two seasonal variations in parallel with the individual ducts to the outside independently seasonal gonadal cycle. at the tip of the papilla is concerned. Lagler and Chin (1951) have recorded lobate genital In course of his description ofthe male papilla in the harmaphroditic percid fish, reproductive system in a number of species Boleosoma nigrum. Clerk (1959) has Gobiiformes and Bleniiformes, Eggert (1913) recorded in a harmaphroditic serranid fish, found erectile tissue in the urinogenital Serranalus subuligarius, the of papilla of some gobies. Hora and Law (1914) the genital papilla when the sperm is have reported the presence of a prominent released from a small opening at the tip. urinogenital papilla or papilla urogenital is Egami (1960) reported the presence of the amongst certain species of Batasio urinogenital papilla in both the sexes of (Bagridae). Mookerjee, Mazumdar and several species of Japanese gobies. Dasgupta (1941) have recorded the presence According to him the papilla is different ofthe papilla or pseudocopulatory organ in both in size and shape between the two both the sexes of Heteropneustes fossilis sexes. He also observed that, at least in and Olarias batrachus. They say that there some species, the enlargement ofthe organ is a median cleft on the ventral surface of is subject to seasonal variations in parallel the female urinogenital papilla representing with the seasonal changes in gonadal the urinogenital opening, while in the male activity. The present study corroborates this oping lies at the tip of the papilla. The with Egami's findings form the formation of a temporary conspicuous morphological point of view but he did not papilla in femalies at the time of oviposition describe the histological nature of the as reported by Parker (1942) for Perea papills. Nair (1960) reported on the papilla flcwesgens has not been observed in any of in the male Heteropneustes fossilis and the species worked out by the present OBSERVATION ON THE URINOGENITAL PAPILLA IS

Rita rita. In Rita rita the papilla shows a Clarias batrachus and the two ducts join. large number of blood sinuses in its This colouration may be nuptial secondary substance, which probably help in the sex character, because this brilliant erection ofthe papilla. The present author colouration which developes at the peak of support Nair's observation on the basis of the breeding season soon starts disappearing his finding on some gobies. Kamalaveni after the is over. This eolouration (1961 a) has reported the presence of must have some recognition value or urinogenital papilla in the males of attracts to some extent the opposite sex Heteropeustes fossilis and Clarias during the peak season. batrachus; in both the sexes of Wallago Davidson (1935) described a very attu)· Eutropiichalus vacha; Notopterus peculiar secondary sex-character in the male notopterus; Notopterus chitala and pink , Onchorhycus gerbuscha. Mastoce1nbalus armatus. Though With the approach of the breeding season Mooke1jee, Mazumdar andDasgupta (1941) the snout elongates and a lnm1p is developed have recorded the presence ofthe papilla in on the dorsal side which starts from behind the females of Heterpneustes fossilis and the head and tapers caudally, the deepest Clarias batrachus, yet Kamalveni (1961d) portion being behind the dorsal fin. Parker has not observed it in the females of these and Brower (1935) discovered the two species. Kamalaveni did not furnish appearence of a melanomerphic mark on any histological data of the papilla nor did the ventral fin of the male of Fundulus make any comment on the organ in general heteroclitus. These spots appear only during dicussion. While describing the urinogenital the breeding season. They termed as organs of. Wallago attu, Sinha (1961) "nuptial secondary sex-character" by the recorded the presence of the papilla or authors.Fraser-Brunner(1940)recorded "pseudocopulatorypapilla" in both the sexes. that in the family Ostractiontidae sexual It is slender and pointed n the male but dimorphism is of common occurrence. It broad and truncated at the apex in the consists of colour difference, depth of the female. Sinha's (1961) findings are body and form of the snout in the two applicable in some of the gobiids under the sexes. In this studies of the morphology present investigation. and embryology of two species of marine Sircar (1970) has observed the papilla , Bagre nwrinus and Galeichthys in the males only ofHeteropheustes fosslis) felis. Merriman (1940) referred to the Clarias batrachus and Eutropiichthys morphological differences between the male rrmrius. Morphology and histology of the and female in both the species, Rosen and papilla have been studied and Gorden (1953) while working on the interrelationship of the urinary and genital evolution of nlale genitalia in helleri, ducts within the urinogenital papilla has observed sexual dimorphism in them. By a been investigated by serial sections. The complex series of morphogenetic processes, urinary and sperm ducts run side by side the anal fin of the male is transformed in to in the papilla for the major part of its a gonopodium. Egami (1960) reported length as in Heteropneustes fossilis and differencess both in shape and size of the 16 HAR genital papilla in several species of Japanese ACKNOWLEDGEMENT gobies. No sexual dimorphism in the shape I thankfully acknowledge my gratitude of the papilla was observed by him in to the Principal of Ramananda College, immature gobies. Such instances of the Bishnupur, Bankura for giving me absence of sexual dimorphism in immature permission and all facilities for this fishes, where dimorphism is pronounced in investigation. I specially indebted to Dr. the adult condition, has not been observed S.K. Chakarborty, Scientist, C.M.F.R.I. by the present investigator. Kamalaveni Mumbai for his criticism and co-operation. (1961 a) reported sexual dimorphism in Thanks are also due to my colleagues of Heteropneustes fossils and Clarias Zoology Department of this college for batrachus. She found no dimorphism in valuable suggestion and kind co-operation. Wallago attu and Mystus aor. All the species worked upon here in connection with the present investigation are sexually REFERENCES dimorphic. Norman (1963, P. 232) wrote; "Sexual differences in the dentition are also Braungart, D.C. 1951 A comparative known. In some Gobies and Blennies the study of the reproductive system of males have dentition is not found as sexually several teleost fishes. Copeia, : 203- dimorphic in any one of the twelve species 205. of gobiids worked out by the present Clark, E. 1959. Functional investigator. Sircar (1970) has reported hermaphroditism and self fertilization pronounced sexual dimorphism among in a serranid fish, Science, 129: 215- siluroid fishes, such as Heteropneoustes 216. follilis, Clarias batrachus, Eutropiichthys nutrias, in all the species named above the Davidson, A. 1935. The development of male have a muscular, tapering Secondary Sexual characters in pink minogenital papilla and the cloaca is absent, salmon (Oncorhynchus gorbuscha). J. the anal opening being distinct and separate. Morph., 57: 169-183. In the female of Eutropiichthys murias, a *Eggert, B. 1931. Die geschilechtsorgane membranous flap-like structure has been der Gobiifom1es and Bleniiformes. Wiss. found between the pelvic fins. The flap Zool., 139:249-558. extends posterior up to the anus and then again starts from behind the anus and Egami, N. 1960. Comparative morphology extends up to the base ofthe "genital " of the sex characters in several species or "accessory tube''. The presence of a soft of Japanese gobies, with reference to tubular "genital pouch" in the female and the effect of sex steroids on the muscular tapering urinogenital papilla in character. J.Fac.Sci. Tohyo. Univ:(Zool) the male is also a sexually dimorphic 9:67-100. character. Fraser- Brunner, A. 1940. Notes on the plectognath fishes. IV. sexual dimorphism 1n the family OBSERVATION ON THE URINOGENITAL PAPILLA 17

Ostraciontidae. Ann.Mag.nat.Hist., (Bloch); Part V, A comparative study of 11(6): 390-392. the of Mystus aor (Ham.) Proc. Indian Sci. Congr., Hora, S.L. and Law, N.C. 1941. Siluroid 47 (3): 471-473. fishes of India, Burma and Ceylon. 9, 10. Rec. Indian, Mus., 43: 9-42. Norman, T.R. 1963. A History of fishes. 2nd edition revised by P.H. Greenwood Kamalaveni, S. 1961 a. Morphology of London, Ernest Be1m. Ltd. the urinogenital system in some Indian teleostean fishes. Rec.Indian Mus., 59: Parker, J.B. 1942. Some observations on 83-118. the reproductive system of yellow perch (Perea flavescens) Copeia.: 223-226. Lagler, K.E. and Chin, M.B. 1951. Ovotestis in the percid fish, Boleosoma Parker, G.H. and Brower, H.P. 1935. A nigrwn (Refinesque). Copeia, 99-100. nuptial secondary sex character in Fundulus heteroclitus. Biol Bull Merriman, D. 1940. Morphological and Woods Hole. 68: 4-6. embrylogical studies on two species of marine fish, Bigre marinus and Rosen, D.E. and Gordon, M. 1953. Galeichthys felis.Zoologica, N.Y. 25: Functional and evolution of 221-248. Male genitalia in poecillid fishes. Zoologica.N. Y. 38: 1-47. Mookerjee, H.K. Mazumdar, S.R. and Dasgupta, B.N. 1941. On the nature Sircar, A.K. 1970. Morphology of the of the urinogenital papilla of Clarias urinogenital system of some siluroid batrachus (Linn), Heteropneustes fishes Proc.Zool.Soc. Calcutta. 23: 93- fossilis (Bloch.) Indian.J.vet.Sci. 11: 117. 334-336. Sinha, B.M. 1961. The urinogenital organs Nair P.V. 1960. Studies on the accessory of Wallago attu (Bloch). Proc. nat.Acad. glands and testes of some siluroid Sci.India 31(B); 296-304. fishes. Part I, Heteropneustes fossilis Weisel. C.F. 1949. The (Bloch); Part II, Clarias batrachus and testes of Gillichthys, a marine. (Bleeker); Part III, Rita rita teleost. Copeia.: 101-110. (Hamilton); Part IV, Mystus vittatus

* Not referred to in original