Gonadal development in males of the orange mud crab, Scylla olivacea (Herbst, 1796) (Decapoda, Brachyura, Portunidae)

in Crustaceana
Restricted Access
Get Access to Full Text
Rent on DeepDyve

Have an Access Token?

Enter your access token to activate and access content online.

Please login and go to your personal user account to enter your access token.


Have Institutional Access?

Access content through your institution. Any other coaching guidance?


This study describes the morphological and histological characteristics of the male reproductive system of the orange mud crab, Scylla olivacea (Herbst, 1796). Three maturation stages were determined on the basis of the vas deferens diameter, the gonad colour and the gonadosomatic index (GSI). Significant differences were observed in the GSI value as the crabs matured. All germ cells were present in the histological sections of the testes, and these decrease significantly in size as they progress from spermatogonia, spermatocytes, and spermatids to spermatozoa. Spermatophores are found in the anterior vas deferens (AVD) and median vas deferens (MVD) of all maturation stages but not in the posterior vas deferens (PVD), with a significantly smaller size in immature and maturing specimens. Thus, the classification of S. olivacea gonads into various maturation stages based solely on histological characteristics (i.e., the presence of spermatophores) is impossible. Therefore, the combination of both microscopic and macroscopic features is essential for determining the gonad maturation stages of male S. olivacea.


International Journal of Crustacean Research



BalasubramanianC. P.SuseelanC., 2000. Male reproductive system and spermatogenesis in the deep water crab Charybdis smithii McLeay (Brachyura: Portunidae). Indian J. Fish., 47: 275-282.

BarberB. J.BlakeN. J., 2006. Reproductive physiology. In: ShumwayS. E.ParsonsG. J. (eds.), Scallops: biology, ecology, and aquaculture (2nd ed.): 357-416. (Elsevier, Amsterdam).

CastilhoG. G.OstrenskyA.PieM. R.BoegerW. A., 2008. Morphology and histology of the male reproductive system of the mangrove land crab Ucides cordatus (L.) (Crustacea, Brachyura, Ocypodidae). Acta Zool., 89: 157-161.

GarciaT. M.SilvaJ. R. F., 2006. Testis and vas deferens morphology of the red-clawed mangrove tree crab (Goniopsis cruentata) (Latreille, 1803). Braz. Arch. Biol. Techn., 49: 339-345.

GardnerC. E.WilliamsH., 2002. Maturation in the giant crab, Pseudocarcinus gigas, and the potential for sperm limitation in the Tasmania fishery. Mar. Freshwater Res., 53: 661-667.

HanK.DaiY.ZhouZ.FuM.WangY.ZhangZ., 2012. Molecular characterization and expression profiles of cdc2 and cyclin B during oogenesis and spermatogenesis in green mud crab (Scylla paramamosain). Comp. Biochem. Phys. B, 163: 292-302.

HinschG. W., 1988. Ultrastructure of the sperm and spermatophores of the golden crab Geryon fenneri and a closely related species, the red crab G. quinquedens, from the eastern Gulf of Mexico. J. Crustacean Biol., 8: 340-345.

IkhwanuddinM.AzmieG.JuariahH. M.ZakariaM. Z.AmbakM. A., 2011. Biological information and population features of mud crab, genus Scylla from mangrove areas of Sarawak, Malaysia. Fish. Res., 108: 299-306.

IkhwanuddinM.AzmieG.Mohd IhwanZ.AmbakM. A., 2013. Mud crab: culture system and practice in Malaysia. (Penerbit Universiti Malaysia Terengganu, Terengganu).

IslamM. S.KurokuraH., 2012. Male reproductive biology of mud crab Scylla olivacea in a tropical mangrove swamps. J. Fish. Aquat. Sci., 7: 194-204.

IslamM. S.KurokuraH., 2013. Gonad development and size at maturity of the male mud crab Scylla paramamosain (Forsskål, 1755) in a tropical mangrove swamp. J. Fish., 1: 7-13.

KeenanC. P.DavieP. J. F.MannD. L., 1998. A revision of the genus Scylla De Haan, 1833 (Crustacea: Decapoda: Brachyura: Portunidae). Raffles Bull. Zool., 46: 217-245.

KrolR. M.HawkinsW. E.OverstreetR. M., 1992. Reproductive components. In: HarrisonF. W. (ed.), Microscopic anatomy of invertebrates, Decapoda, Crustacea, 10: 295-343. (Wiley-Liss, New York, NY).

López GrecoL. S.VazquezF.RodríguezE. M., 2007. Morphology of the male reproductive system and spermatophore formation in the freshwater ‘red claw’ crayfish Cherax quadricarinatus (Von Martens, 1898) (Decapoda, Parastacidae). Acta Zool., 88: 223-229.

Manjón-CabezaM. E.RasoJ. E. G., 2000. Morphological reproductive aspects of males of Diogenes pugilator (Roux, 1829) (Crustacea, Decapoda, Anomura) from southern Spain. Sarsia, 85: 195-202.

MedinaA.RodriguezA., 1992. Spermiogenesis and sperm structure in the crab Uca tangeri (Crustacea, Brachyura), with special reference to the acrosome differentiation. Zoomorphology, 111: 161-165.

Mota-AlvesM. I., 1975. Sobre a reprodução do caranguejo-uçá, Ucides cordatus (Linnaeus), em mangues do Estado do Ceará (Brasil). Arq. Cienc. Mar., 15: 85-91.

NascimentoF. A.ZaraF. J., 2013. Development of the male reproductive system in Callinectes ornatus Ordway, 1863 (Brachyura: Portunidae). Nauplius, 21: 161-177.

OshiroL. M. Y.SilvaR.SilveiraC. M., 1999. Rendimento de carne nos caranguejos Guaiá, Menippe nodifrons Stimpson, 1859 e Guaiamum, Cardisoma guanhumi Latreille, 1825 (Crustacea, Decapoda, Brachyura) da Baía de Sepetiba/RJ. Acta Biol. Leopold., 21: 83-88.

RaviR.ManisseriM. K.SanilN. K., 2014. Structure of the male reproductive system of the blue swimmer crab Portunus pelagicus (Decapoda: Portunidae). Acta Zool., 95: 176-185.

RyanE. P., 1967. Structure and function of the reproductive system of the crab Portunus sanguinolentus (Herbst, 1785) (Brachyura, Portunidae). Proceedings of the Symposium on Crustacea. Marine Biological Association of India, 2: 522-544.

Sal MoyanoM. P.GavioM. A.CuartasE. I., 2010. Morphology and function of the reproductive tract of the spider crab Libinia spinosa (Crustacea, Brachyura, Majoidea): pattern of sperm storage. Helgoland Mar. Res., 64: 312-321.

SantosC. M.LimaG. V.NascimentoA. A.SalesA.OshiroL. M. Y., 2009. Histological and histochemical analysis of the gonadal development of males and females of Armases rubripes (Rathbun 1897) (Crustacea, Brachyura, Sesarmidae). Braz. J. Biol., 69: 161-169.

SherkhaneU. D.PatilM. U.PandeG. S., 2010. Gross anatomy of male reproductive system and histology of testis and vas deferens in freshwater crab Barytelphusa cunicularis (Westwood 1836) (Decapoda: Crustacea). The Bioscan, 5: 599-603.

Shinozaki-MendesR. A.SilvaJ. R. F.HazinF. H. V., 2012. Development of male reproductive system of the blue land crab Cardisoma guanhumi Latreille, 1828 (Decapoda: Gecarcinidae). Acta Zool., 93: 390-399.

SilvaL. S.Martinelli-LemosJ. M.FerreiraM. A. P.RochaR. M., 2012. Gonadal development in the freshwater crab Sylviocarcinus pictus (H. Milne Edwards, 1853) (Brachyura: Trichodactylidae) from the Guamá River, state of Pará, Brazil. An. Acad. Bras. Ciênc., 84: 789-798.

SimeóC. G.RibesE.RotllantG., 2009. Internal anatomy and ultrastructure of the male reproductive system of the spider crab Maja brachydactyla (Decapoda: Brachyura). Tissue Cell, 41: 345-361.

SimeóC. G.KurtzK.ChivaM.RibesE.RotllantG., 2010. Spermatogenesis of the spider crab Maja brachydactyla (Decapoda: Brachyura). J. Morphol., 271: 394-406.

SokalR.RohlfE., 1995. Biometry — the principles and practice of statistics in biological research (3rd ed.). (Freeman, New York, NY).

SoundarapandianP.IlavarasanN.VaradharajanD., 2013a. Reproductive system of flower crab, Charybdis feriata (Linnaeus). Open Access Scientific Reports, 2: 701. DOI:10.4172/scientificreports.701.

SoundarapandianP.VaradharajanD.AnandT., 2013b. Male reproductive system of blue swimming crab, Portunus pelagicus (Linnaeus, 1758). J. Cytol. Histol., 5: 206.

StewartM. J.StewartP.SoonklangN.LinthongV.HannaP. J.DuanW.SobhonP., 2010. Spermatogenesis in the blue swimming crab, Portunus pelagicus, and evidence for histones in mature sperm nuclei. Tissue Cell, 42: 137-150.

SubramaniamT., 1995. Light and electron microscopic studies on the seminal secretions and the vas deferens of the penaeiodean shrimp, Sicyonia ingentis. J. Bioscience, 20: 691-706.

SuganthiA. S.AnilkumarG., 1999. Moult-related fluctuation in exdysteroid titre and spermatogenesis in the crab, Metopograpsus messor (Brachyura: Decapoda). Zool. Stud., 38: 314-321.

UmaK.SubramoniamT., 1979. Histochemical characteristics of spermatophore layers of Scylla serrata (Forskal) (Decapoda: Portunidae). Int. J. Inver. Rep., 1: 31-40.

VallinaM.Sal MoyanoM. P.CuartasE. I.GavioM. A., 2014. Reproductive system and size maturity of the paddle crab Ovalipes trimaculatus (Brachyura: Portunidae) along the Argentine coast. J. Crustacean Biol., 34: 357-366.

Van EngelW. A., 1990. Development of the reproductive functional form in the male blue crab, Callinectes sapidus. B. Mar. Sci., 46: 13-22.

WaihoK.FazhanH.IkhwanuddinM., 2016. Size distribution, length-weight relationship and size at the onset of sexual maturity of the orange mud crab, Scylla olivacea in Malaysian waters. Mar. Biol. Res., 12: 726-738.

WaihoK.MustaqimM.FazhanH.Wan NorfaizzaW. I.MegatF. H.IkhwanuddinM., 2015. Mating behaviour of the orange mud crab, Scylla olivacea: the effect of sex ratio and stocking density on mating success. Aquacult. Rep., 2: 50-57.


  • Male reproductive organs of Scylla olivacea (Herbst, 1796): A, internal male reproductive system of S. olivacea with the thin commissure (TC) located between the testes (T) and anterior vasa deferentia (AVD) and connecting the two sides of the gonad; B, extracted internal male reproductive system of S. olivacea without the linking commissure due to breakage during handling, with clear differentiation of testis (T), anterior vas deferens (AVD), median vas deferens (MVD) and posterior vas deferens (PVD); C, external male reproductive system of S. olivacea with the first (G1) and second (G2) gonopods clearly visible. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/15685403.

    View in gallery
  • Mean testis weight, mean anterior vas deferens (AVD) + median vas deferens (MVD) weight, mean posterior vas deferens (PVD) + ejaculatory duct (ED) weight and total gonad weight of male Scylla olivacea (Herbst, 1796) with increasing carapace width (CW).

    View in gallery
  • Male reproductive system of Scylla olivacea (Herbst, 1796) in histological cross-section (Mayer’s Hematoxylin-Eosin staining): A, testes containing spermatogonia (SPG), spermatocytes (SPC), spermatids (SPT) and spermatozoa (SPZ), magnification 20×; B, anterior vas deferens (AVD) containing closely packed spermatophores (SPH) with minimal spermatophoric matrix (SM), magnification 20×; C, cell wall of AVD made up of stratified cuboidal epithelium (SCE) and enveloped by a thin layer of elongated cells (EC), magnification 100×; D, median vas deferens (MVD) showing spermatophores (SPH) loosely distributed in spermatophoric matrix (SM) within the lumen cavity, magnification 4×; E, detailed zoom-in of a typical oval-shaped spermatophore (SPH) containing spermatozoa, surrounded by spermatophoric matrix (SM) with basophilic (black arrow) and acidophilic (white arrow) secretions, magnification 100×; F, cell wall of MVD consisting of a thin layer of elongated cells (EC) (outer layer) and simple squamous epithelium (SSE) (inner layer) with basophilic (black arrow) and acidophilic (white arrow) secretions, magnification 100×; G, posterior vas deferens (PVD) with only seminal fluid (SF) and no sign of spermatophores, magnification of 10×; H, cell wall of PVD composed of simple columnar epithelium (SLE), surrounded by a single layer of elongated cells (EC) with basophilic (black arrow) and acidophilic (white arrow) secretions, magnification of 100×. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/15685403.

    View in gallery
  • Different types of germ cells found in the histological cross-section of Scylla olivacea (Herbst, 1796) testis, magnification 100×: A, spermatogonia (SPG); B, spermatocytes (SPC); C, spermatids (SPT); D, spermatozoa (SPZ). This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/15685403.

    View in gallery


Content Metrics

Content Metrics

All Time Past Year Past 30 Days
Abstract Views 46 46 17
Full Text Views 6 6 6
PDF Downloads 0 0 0
EPUB Downloads 0 0 0