Population Biology, Reproductive Biology, Fisheries, and Future Perspective to Develop Three-Spot Swimming Crab Portunus sanguinolentus as New Aquaculture Candidate: A Review

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Khor Waiho
Rusydi Rozaimi
Supawadee Poompuang
Suriyan Tunkijjanukij
Alexander Chong Shu-Chien
Youji Wang
Mhd Ikhwanuddin
Sila Sukhavachana
Hanafiah Fazhan


The three-spot swimming crab Portunus sanguinolentus is an easily recognisable portunid species owing to the three red spots on its carapace. Like other portunid swimming crabs, P. sanguinolentus is an edible species found across the Indo-Pacific region, and it is gaining attention as one of the important edible crabs in the coastal crab fishery sector of many countries. In addition to being one of the main crab species (along with P. pelagicus and P. trituberculatus) that support the livelihood of coastal communities, P. sanguinolentus also exhibits high aquaculture potential. This species is often associated with coastal intertidal and subtidal zones, with juveniles and males preferably inhabiting depths up to 30 m, whereas females may be found as deep as 80 m. As P. sanguinolentus is a highly exploited portunid species with great aquaculture potential, this review summarises the available literature to provide a holistic compilation of the current knowledge, ranging from taxonomy and phylogeny, growth and life cycle, reproductive maturity and breeding, to feeding behaviour and food preference; we also highlight the knowledge gaps that require future emphasis. This is the first comprehensive review of the life history and other biological information of P. sanguinolentus. This review serves as an essential baseline for sustainable fishery management and aquaculture development of this species.


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Abdullah-Zawawi, M.R., N. Afiqah-Aleng, M. Ikhwanuddin, Y.Y. Sung, S. Tola, H. Fazhan and K. Waiho. 2021. Recent development in ecdysone receptor of crustaceans: Current knowledge and future applications in crustacean aquaculture. Reviews in Aquaculture 12: 1938-1957. DOI: 10.1111/raq.12552.

Al-Aidaroos, A.M., A.A.J. Kumar, A.E. Al-Haj, A. Al-Sofyani, M.P. Crosby and M.M El-Sherbiny. 2019. Morphology of the complete larval stages of Portunus segnis (Forskål, 1775) (Crustacea: Brachyura: Portunidae) from the Gulf of Aqaba, Saudi Arabia. Zootaxa 4638: 199-218. DOI: 10.11646/zootaxa.4638.2.2.

Alberts-Hubatsch, H.A., S.Y. Lee, J.O. Meynecke, K. Diele, I. Nordhaus and M. Wolff. 2016. Life-history, movement, and habitat use of Scylla serrata (Decapoda, Portunidae): Current knowledge and future challenges. Hydrobiologia 763: 5-21. DOI: 10.1007/s10750-015-2393-z.

Ali, A., S.E. Sherif, J.A. Alla, S. Maulu, A.A Tantawy, M.F.K. Soliman, M.M. Hassan, M.M. Hassan and H.S. Khalil. 2021. Morphometric, histochemical, and ultrastructural analysis of the reproductive system and spermatogenic stages of male blue crab (Callinectes sapidus Rathbun, 1896). Journal of Marine Science and Engineering 9: 1105. DOI: 10.3390/jmse9101105.

AquaMaps. 2019. Computer generated distribution maps for Portunus sanguinolentus (threespot swimming crab), with modelled year 2050 native range map based on IPCC RCP8.5 emissions scenario. https://www.aquamaps.org. Cited 6 Jan 2022.

Ariyama, H. 1996. Life history of the red-spotted swimming crab Portunus (Portunus) sanguinolentus in Osaka Bay. Benthos Research 51: 1-8. DOI: 10.5179/benthos1996.51.1_1.

Ates, M.C.D., G.F. Quinitio, E.T. Quinitio and R.C. Sanares. 2012. Comparative study on the embryonic development of three mud crabs Scylla spp. Aquaculture Research 43: 215-225. DOI: 10.1111/j.1365-2109.2011.02818.x.

Azra, M.N. and M. Ikhwanuddin. 2015. Larval culture and rearing techniques of commercially important crab, Portunus pelagicus (Linnaeus, 1758): Present status and future prospects. Songklanakarin Journal of Science and Technology 37: 135-145.

Babu, B.T., K. Shyamasundari and K.H. Rao. 1989. Studies of the seminal receptacle of the crab Portunus sanguinolentus (Portunidae: Brachyura). Folia Morphologica 37: 4-8.

Beninger, P.G., R.W. Elner, T.P. Foyle and P.H. Odense. 1988. Functional anatomy of the male reproductive system and the female spermatheca in the snow crab Chionoecetes opilio (O. Fabricius) (Decapoda: Majidae) and a hypothesis for fertilization. Journal of Crustacean Biology 8: 322-332. DOI: 10.2307/1548271.

Campbell, G.R. and D.R. Fielder. 1986. Size at sexual maturity and occurrence of ovigerous females in three species of commercially exploited portunid crabs in S.E. Queensland. Proceedings of the Royal Society Queensland 97: 79-87.

Danielsen, H.E.H., A.M. Hjelset, B.A. Bluhm, C. Hvingel and A.L. Agnalt. 2019. A first fecundity study of the female snow crab Chionoecetes opilio Fabricius, 1788 (Decapoda: Brachyura: Oregoniidae) of the newly established population in the Barents Sea. Journal of Crustacean Biology 39: 485-492. DOI: 10.1093/jcbiol/ruz039.

Dash, G., S. Sen, K.M. Koya, K.R. Sreenath, S.K. Mojjada, M.S. Zala and S. Pradeep. 2013. Fishery and stock assessment of the three-spot swimming crab Portunus sanguinolentus (Herbst, 1783) off Veraval, Gujarat. Indian Journal of Fisheries 60: 17-25.

Dineshbabu, A.P., B. Sreedhara and Y. Muniyappa. 2007. Fishery and stock assessment of Portunus sanguinolentus (Herbst) from south Karnataka coast, India. Journal of the Marine Biological Association of India 49: 134-140.

Duan, B., S. Mu, Y. Guan, S. Li, Y. Yu, W. Liu, Z. Li, X. Ji and X. Kang. 2022. Genetic diversity and population structure of the swimming crab (Portunus trituberculatus) in China seas determined by genotyping-by-sequencing (GBS). Aquaculture 555: 738233. DOI: 10.1016/j.aquaculture.2022.738233.

Food and Agriculture Organisation of the United Nations (FAO). 2021. Fisheries and Aquaculture-Global Production by Production Source (1950-2019). https://www.fao.org/fishery/statistics query/en/global_production/global_ production_quantity. Cited 12 Jan 2022.

Fazhan, H., K. Waiho, Y. Fujaya, N. Rukminasari, H. Ma and M. Ikhwanuddin. 2021. Sexual dimorphism in mud crabs: a tale of three sympatric Scylla species. PeerJ 9: e10936. DOI: 10.7717/peerj.10936.

Fazhan, H., K. Waiho, M. Ikhwanuddin, A.C. Shu-Chien, Y. Fujaya, Y. Wang, H.J. Liew, C. Chen, M.H. Abualreesh, A. Jaya-Ram, and A.R. Ahmad-Ideris. 2022. Limb loss and feeding ability in the juvenile mud crab Scylla olivacea: Implications of limb autotomy for aquaculture practice. Applied Animal Behaviour Science 247: 105553. DOI: 10.1016/j.applanim.2022.105553.

Fisher, M.R. 1999. Effect of temperature and salinity on size at maturity of female blue crabs. Transactions of the American Fisheries Society 128: 499-506. DOI: 10.1577/1548-8659(1999)128<0499:EOTASO>2.0.CO;2.

Froese, R. and C. Binohlan. 2000. Empirical relationships to estimate asymptotic length, length at first maturity and length at maximum yield per recruit in fishes, with a simple method to evaluate length frequency data. Journal of Fish Biology 56: 758-773. DOI: 10.1111/j.1095-8649.2000.tb00870.x.

Gao, B., D. Sun, J. Lv, X. Ren, P. Liu and J. Li. 2019. Transcriptomic analysis provides insight into the mechanism of salinity adjustment in swimming crab Portunus trituberculatus. Genes and Genomics 41: 961-971. DOI: 10.1007/s13258-019-00828-4.

Gardner, C. 1997. Effect of size on reproductive output of giant crabs Pseudocarcinus gigas (Lamarck): Oziidae. Marine and Freshwater Research 48: 581-587. DOI: 10.1071/MF97023.

Greenwood, J.G. and D.R. Fielder. 1979. The zoeal stages and megalopa of Portunus rubromarginatus (Lanchester) (Decapoda: Portnnidae), reared in the laboratory. Journal of Plankton Research 1: 191-205. DOI: 10.1093/plankt/1.2.191.

Gunderson, D.R. 1980. Using r-K selection theory to predict natural mortality. Canadian Journal of Fisheries and Aquatic Sciences 37: 2266-2271.DOI: 10.1139/f80-272.

Hamasaki, K., K. Fukunaga and K. Maruyama. 1992. Effects of temperature on egg incubation period and synchronization of hatching rate in the swimming crab Portunus trituberculatus. Saibai Giken 20: 73-85.

Hamasaki, K., K. Fukunaga and K. Maruyama. 2003. Egg development and incubation period of the swimming crab Portunus trituberculatus (Decapoda: Portunidae) reared in the laboratory. Crustacean Research 32: 45-54. DOI: 10.18353/crustacea.32.0_45.

Hamasaki, K., N. Matsui and M. Nogami. 2011. Size at sexual maturity and body size composition of mud crab Scylla spp. caught in Don Sak, Bandon Bay, Gulf of Thailand. Fisheries Science 77: 49-57. DOI: 10.1007/s12562-010-0307-6.

Hartnoll, R.G. 1985. Growth in Crustacea - twenty years on. Hydrobiologia 449: 111-122.

Haukioja, E. and T. Hakala. 1979. Asymptotic equations in growth studies-an analysis based on Anodonta piscinalis (Mollusca, Unionidae). Annales Zoologici Fennici 16: 115-122.

Hill, A.E., J. Brown and L. Fernand. 1997. The summer gyre in the western Irish Sea: shelf sea paradigms and management implications. Estuarine, Coastal and Shelf Science 44: 83-95. DOI: 10.1016/S0272-7714(97)80010-8.

Hines, A.H. 1989. Geographical variation in size at maturity in brachyuran crabs. Bulletin of Marine Science 45: 356-368.

Ikhwanuddin, M., M.N. Azra, N.F. Noorulhudha, A. Siti-Aishah and A.B. Abol-Munafi. 2016. Embryonic development and hatching rate of blue swimming crab, Portunus pelagicus (Linnaeus, 1758) under different water salinities. Turkish Journal of Fisheries and Aquatic Sciences 16: 669-677. DOI: 10.4194/1303-2712-v16_3_21.

Islam, M.S., S. Shokita and T. Higa. 2000. Larval development of the swimming crab Charybdis natator (Crustacea: Brachyura: Portunidae) reared in the laboratory. Species Diversity 5: 329-349. DOI: 10.12782/specdiv.5.329.

Jacob, R., P.N. Prasad and M.S. Kusuma. 1990. Maturity and dimensional studies in female crabs of Portunus sanguinolentus and P. pelagicus (Decapoda: Portunidae). Indian Journal of Marine Sciences 19: 221-223.

Jantrarotai, P., K. Sirisintruwanich, S. Pripanapong and C. Chayarat. 2006. Morphlogical study in zoeal stages of mud crabs: Scylla olivacea (Herbst, 1796). Kasetsart Journal: Natural Science 40: 507-516.

Josileen, J. and N.G. Menon. 2004. Larval stages of the blue swimmer crab, Portunus pelagicus (Linnaeus, 1758) (Decapoda, Brachyura). Crustaceana 77: 785-803. DOI: 10.1163/156854004774248681.

Kepel, A.A. and L.A. Tsareva. 2005. First record of tropical crabs Portunus sanguinolentus (Herbst, 1783) and Plagusia depressa tuberculata Lamarck, 1818 in Peter the Great Bay, Sea of Japan. Russian Journal of Marine Biology 31: 124-125. DOI: 10.1007/s11179-005-0054-1.

King, M. 1995. Fisheries Biology, Assessment and Management. Fishing News Books, London, UK. 341 pp.

Koolkalya, S., T. Thapanand, S. Tunkijjanujij, V. Havanont and T. Jutagate. 2006. Aspects in spawning biology and migration of the mud crab Scylla olivacea in the Andaman Sea, Thailand. Fisheries Management and Ecology 13: 391-397. DOI: 10.1111/j.1365-2400.2006.00518.x.

Kornienko, E.S. 2020. The reproductive strategy of hermit crabs in temperate waters. Russian Journal of Marine Biology 46: 319-329. DOI: 10.1134/S1063074020050053.

Kurata, H. and T. Midorikawa. 1975. The larval stages of the swimming crabs, Portunus pelagicus and Portunus sanguinolentus reared in the laboratory. Bulletin of Nansei National Fisheries Research Institute 8: 29-38.

Lee, H.H. and C.C. Hsu. 2003. Population biology of the swimming crab Portunus sanguinolentus in the waters off northern Taiwan. Journal of Crustacean Biology 23: 691-699. DOI: 10.1651/C-2358.

Leffler, C.W. 1972. Some effects of temperature on the growth and metabolic rate of juvenile blue crabs, Callinectes sapidus, in the laboratory. Marine Biology 14: 258-272. DOI: 10.1007/BF00373209.

Liu, J.Y. 2008. Checklist of Marine Biota of China Seas. Science Press, Beijing, China. 1267 pp.

Macedo, D., I. Caballero, M. Mateos, R. Leblois, S, McCay and L.A. Hurtado. 2019. Population genetics and historical demographic inferences of the blue crab Callinectes sapidus in the US based on microsatellites. PeerJ 7: e7780. DOI: 10.7717/peerj.7780. DOI: 10.7717/peerj.7780.

Madduppa, H., R. Martaulina, Z. Zairion, R.M. Renjani, M. Kawaroe, N.P. Anggraini, B. Subhan, I. Verawati and L.M.I. Sani. 2021. Genetic population subdivision of the blue swimming crab (Portunus pelagicus) across Indonesia inferred from mitochondrial DNA: Implication to sustainable fishery. PLoS ONE 16: e0240951. DOI: 10.1371/journal.pone.0240951.

Marco-Herrero, E., J.A. Cuesta and J.I. González-Gordillo. 2021. DNA barcoding allows identification of undescribed crab megalopas from the open sea. Scientific Reports 11: 20573. DOI: 10.1038/s41598-021-99486-4.

Meng, X, F. Jia, X. Zhang, P. Liu and J. Li. 2016. Complete sequence and characterization of mitochondrial genome in the swimming crab Portunus sanguinolentus (Herbst, 1783) (Decapoda, Brachyura, Portunidae). Mitochondrial DNA Part A: DNA Mapping, Sequencing, and Analysis 27: 3052-3053. DOI: 10.3109/19401736.2015.1063130.

Meynecke, J.O. and R.G. Richards. 2014. A full life cycle and spatially explicit individual-based model for the giant mud crab (Scylla serrata): a case study from a marine protected area. ICES Journal of Marine Science 71: 484-498. DOI : 10.1093/icesjms/fst181.

Naimullah, M., Y.L. Wu, M.A. Lee and K.W. Lan. 2021. Effect of the El Niño-Southern oscillation (ENSO) cycle on the catches and habitat patterns of three swimming crabs in the Taiwan strait. Frontiers in Marine Science 8: 763543. DOI: 10.3389/fmars.2021.763543.

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

Naz, F., N.S. Saher and M. Kamal. 2016. Genetic diversity of the Portunus sanguinolentus (Herbst, 1783) (Decapoda, Brachyura, Portunidae) in Indo West Pacific region base on mitochondrial DNA 16S gene. Pakistan Journal of Marine Sciences 25: 59-68.

Ng, P.K.L. 1998. Crabs. In: FAO Species Identification Guide for Fishery Purposes (eds, K.E. Carpenter and V.H. Niem), pp. 1045-1055. Food and Agriculture Organisation of the United Nations, Rome, Italy.

O’Halloran, M.J. and R.K O’Dor. 1988. Molt cycle of male snow crabs, Chionoecetes opilio, from observations of external features, setal changes and feeding behavior. Journal of Crustacean Biology 8: 164-176. DOI: 10.2307/1548309.

Olsen, E.M., G.R. Lilly, M. Heino, M.J. Morgan, J. Brattey and U. Dieckmann. 2005. Assessing changes in age and size at maturation in collapsing populations of Atlantic cod (Gadus morhua). Canadian Journal of Fisheries and Aquatic Sciences 62: 811-823. DOI: 10.1139/f05-065.

Pauly, D. 1980. On the interrelationship between natural mortality, growth parameters and mean environmental temperature in 175 fish stocks. Journal du Conseil 39: 175-192. DOI : 10.1093/icesjms/39.2.175.

Pillai, S.L and P. Thirumilu. 2012. Fishery, biology and yield estimates of Portunus sanguinolentus off Chennai. Journal of the Marine Biological Association of India 54: 73-76.

Quinn, T.J.I.I and R.B. Deriso. 1999. Quantitative Fish Dynamics. Oxford University Press, New York, USA. 560 pp.

Rasheed, S. and J. Mustaquim. 2018. Natural diet of two commercial crab species, Portunus segnis (Forskål, 1775) and P. sanguinolentus (Herbst, 1783), in the Coastal Waters of Karachi. Animal and Veterinary Sciences 6: 35-42. DOI: 10.11648/j.avs.20180603.11.

Rasheed, S. and J. Mustaquim J. 2010. Size at sexual maturity, breeding season and fecundity of three-spot swimming crab Portunus sanguinolentus (Herbst, 1783) (Decapoda, Brachyura, Portunidae) occurring in the coastal waters of Karachi, Pakistan. Fisheries Research 103: 56-62. DOI: 10.1016/j.fishres.2010.02.002.

Reeby, J., P.N. Prasad and M.S. Kusuma. 1990. Size at maturity in the male crabs of Portunus sanguinolentus and P. pelagicus. Fishery Technology 27: 115-119.

Ren, G., H. Ma, C. Ma, W. Wang, W. Chen and L. Ma. 2017. Genetic diversity and population structure of Portunus sanguinolentus (Herbst, 1783) revealed by mtDNA COI sequences. Mitochondrial DNA Part A: DNA Mapping, Sequencing, and Analysis 28: 740-746. DOI: 10.1080/24701394.2016.1177040.

Rouf, M.A., S.I.M. Shahriar, A-H. Antu and M.N. Siddiqui. 2021. Population parameters of the orange mud crab Scylla olivacea (Herbst, 1796) from the Sundarban mangrove forest in Bangladesh. Heliyon 7: e06223. DOI: 10.1016/j.heliyon.2021.e06223.

Ryan, E.P. 1967. Structure and function of the reproductive system of the crab Portunus sanguinolentus (Herbst) (Brachyura: Portunidae). II. Female system. Journal of the Marine Biological Association of India. Symposium Series 2: 522-544.

Samuel, N.J. and P. Soundarapandian. 2009. Embryonic development of commercially important portunid crab Portunus sanguinolentus (Herbst). International Journal of Animal and Veterinary Advances 1: 32-38. DOI: 10.4172/2155-9910.1000122.

Samuel, N.J., P. Soundarapandian and T. Anand. 2011. Larval development of commercially important crab Portunus sanguinolentus (Herbst) reared under laboratory conditions. Research Journal of Fisheries and Hydrobiology 6: 22-40.

Sarada, P.T. 1998. Crab fishery of the Calicut coast with some aspects of the population characteristics of Portunus sanguinolentus, P. pelagicus and Charybdis cruciata. Indian Journal of Fisheries 45: 375-386.

Soundarapandian, P., D. Varadharajan and A. Boopathi. 2013. Reproductive biology of the commercially important portunid crab, Portunus Sanguinolentus (Herbst). Journal of Marine Science Research and Development 3: 124. DOI: 10.4172/2155-9910.1000124.

Stephenson, W. and B. Campbell. 1959. The Australian portunids (Crustacea: Portunidae). III. The genus Portunus. Marine and Freshwater Research 10: 84-123. DOI: 10.1071/MF9590084.

Sugumar, V., G. Vijayalakshmi and K. Saranya. 2013. Molt cycle related changes and effect of short term starvation on the biochemical constituents of the blue swimmer crab Portunus pelagicus. Saudi Journal of Biological Sciences 20: 93-103. DOI: 10.1016/j.sjbs.2012.10.003.

Sukumaran, K.K. and B. Neelakantan. 1997a. Age and growth in two marine portunid crabs, Portunus (Portunus) sanguinolentus (Herbst) and Portunus (Portunus) pelagicus (Linnaeus) along the southwest coast of India. Indian Journal of Fisheries 44: 111-131.

Sukumaran, K.K. and B. Neelakantan. 1997b. Food and feeding of Portunus (Portunus) sanguinolentus (Herbst) and Portunus (Portunus) pelagicus (Linnaeus) (Brachyura: Portunidae) along Karnataka coast. Indian Journal of Marine Sciences 26: 35-38.

Sukumaran, K.K. and B. Neelakantan. 1999. Spawning biology of two marine portunid crabs, Portunus (Portunus) sanguinolentus (Herbst) and Portunus (Portunus) pelagicus (Linnaeus) from the Karnataka Coast. Proceeding of the 4th Indian Fisheries Forum 1996: 35-38.

Sukumaran, K.K., K.Y. Telang and D. Thippeswamy. 1986. On the fishery and biology of the crab Portunus sanguinolentus (Herbst) along the south Kanara coast. Indian Journal of Fisheries 33: 188-200.

Sumpton, W.D., G.S. Smith and M.A. Potter. 1989. Notes on the biology of the Portunid crab, Portunus sanguinolentus (Herbst), in subtropical Queensland waters. Australian Journal of Marine and Freshwater Research 40: 711-717. DOI: 10.1071/MF9890711.

Takween, W. and N.A. Qureshi. 2005. Population structure and reproductive biology of four species of swimming crabs (Crustacea: Brachyura: Portunidae) from coastal area of Karachi, Pakistan. Pakistan Journal of Marine Sciences 14: 107-121.

Tang, B., D. Zhang, H. li, S. Jiang, H. Zhang, F. Xuan, B. Ge, Z. Wang, Y. Liu, S. Sha, Y. Cheng, W. Jiang, H. Jiang, Z. Wang, K. Wang, C. Li, Y. Sun, S. She, Q. Qiu, W. Wang, X. Li, Y, Li, Q. Liu and Y. Ren. 2020. Chromosome-level genome assembly reveals the unique genome evolution of the swimming crab (Portunus trituberculatus). GigaScience 9: giz161. DOI: 10.1093/gigascience/giz161.

Waiho, K., H. Fazhan and M. Ikhwanuddin. 2016. Size distribution, length-weight relationship and size at the onset of sexual maturity of the orange mud crab, Scylla olivacea, in Malaysian waters. Marine Biology Research 12: 726-738. DOI: 10.1080/17451000.2016.1200726.

Waiho, K., H. Fazhan, E.T. Quinitio, J.C. Baylon, Y. Fujaya, G. Azmie, Q. Wu, X. Shi, M. Ikhwanuddin and H. Ma. 2018. Larval rearing of mud crab (Scylla): What lies ahead. Aquaculture 493: 37-50. DOI: 10.1016/j.aquaculture.2018.04.047.

Waiho, K., H. Fazhan, J.C. Baylon, H. Madihah, S. Noorbaiduri, H. Ma and M. Ikhwanuddin. 2017a. On types of sexual maturity in brachyurans, with special reference to size at the onset of sexual maturity. Journal of Shellfish Research 36: 807-839. DOI: 10.2983/035.036.0330.

Waiho, K., H. Fazhan, S. Jasmani and M. Ikhwanuddin. 2017b. Gonadal development in males of the orange mud crab, Scylla olivacea (Herbst, 1796) (Decapoda, Brachyura, Portunidae). Crustaceana 90: 1-19. DOI: 10.1163/15685403-00003622.

Waiho, K., M. Ikhwanuddin, M.H. Abualreesh, A.C. Shu-Chien, S.D. Ishak, M. Jalilah, G. Azmie and H. Fazhan. 2021. Intra- and interspecific variation in sexual dimorphism patterns of mud crab genus Scylla along the equatorial region. Frontiers in Marine Science 8: 690836. DOI: 10.3389/fmars.2021.690836.

Waiho, K., M. Ikhwanuddin, N. Afiqah-Aleng, A.C. Shu-Chien, Y. Wang, H. Ma and H. Fazhan. 2022. Transcriptomics in advancing portunid aquaculture: A systematic review. Reviews in Aquaculture early view. DOI: 10.1111/raq.12689.

Waiho, K., M. Mustaqim, H. Fazhan, W.I. Wan Norfaizza, F.H. Megat and M. Ikhwanuddin. 2015. Mating behaviour of the orange mud crab, Scylla olivacea: The effect of sex ratio and stocking density on mating success. Aquaculture Reports 2: 50-57. DOI: 10.1016/j.aqrep.2015.08.004.

Wang, Z., L. Sun, W. Guan, C. Zhou, B. Tang, Y. Cheng, J. Huang and F. Xuan. 2018. De novo transcriptome sequencing and analysis of male and female swimming crab (Portunus trituberculatus) reproductive systems during mating embrace (stage II). BMC Genetics 19: 3. DOI: 10.1186/s12863-017-0592-5.

Warner, G.F. 1977. The Biology of Crabs. Elek Science London, London, UK. 202 pp.

Wenner, A.M. 1972. Sex ratio as a function of size in marine Crustacea. American Naturalist 106: 32-50. DOI: 10.1086/282774.

Williams, M.J. 1982. Natural food and feeding in the commercial sand crab Portunus pelagicus Linnaeus, 1766 (Crustacea: Decapoda: Portunidae) in Moreton Bay, Queensland. Journal of Experimental Marine Biology and Ecology 59: 165-176. DOI: 10.1016/0022-0981(82)90113-7.

Williams, M.J. and J.H. Primavera. 2001. Choosing tropical portunid species for culture, domestication and stock enhancement in the Indo-Pacific. Asian Fisheries Science 14: 121-142.

Wimalasiri, H.B.U.G.M. and D.C.T. Dissanayake. 2016. Reproductive biology of the three-spot swimming crab (Portunus sanguinolentus) from the west coast of Sri Lanka with a novel approach to determine the maturity stage of male gonads. Invertebrate Reproduction and Development 60: 243-253. DOI: 10.1080/07924259.2016.1202337.

Windsor, A.M., J.C.E. Mendoza and J.R. Deeds. 2019. Resolution of the Portunus gladiator species complex: taxonomic status and identity of Monomia gladiator (Fabricius, 1798) and Monomia haanii (Stimpson, 1858) (Brachyura, Decapoda, Portunidae). ZooKeys 858: 11-43. DOI: 10.3897/zookeys.858.33826.

World Register of Marine Species (WoRMS). 2022. Portunus sanguinolentus (Herbst, 1783 [in Herbst, 1782-1790]). https://www.marinespecies.org/aphia.php?p= taxdetails&id=1061758. Cited 6 Jan 2022.

Wu, R.S.S. and P.K.S. Shin. 1997. Food segregation in three species of portunid crabs. Hydrobiologia 362: 107-113. DOI: 10.1023/A:1003126415990.

Yang, C.P., H.X. Li, L. Li, J. Xu and Y. Yan. 2014a. Population structure, morphometric analysis and reproductive biology of Portunus Sanguinolentus (Decapoda: Brachyura: Portunidae) in Honghai Bay, South China Sea. Journal of Crustacean Biology 34: 722-730. DOI: 10.1163/1937240X-00002273.

Yang, L., Z. Tan, D. Wang, L. Xue, M-x. Guan, T. Huang and R. Li. 2014b. Species identification through mitochondrial rRNA genetic analysis. Scientific Reports 4: 4089. DOI: 10.1038/srep04089.

Yu, H., W. Xu, D. Zhang, H. Cheng, L. Wu and J. He. 2021. Development of embryos and larvae of Portunus trituberculatus (Decapoda, Brachyura) in off-season breeding mode. Crustaceana 94: 679-695. DOI: 10.1163/15685403-bja10126.

Zairion, Y. Wardiatno, M. Boer and A. Fahrudin. 2015. Reproductive biology of the blue swimming crab Portunus pelagicus (Brachyura: Portunidae) in East Lampung Waters, Indonesia: Fecundity and reproductive potential. Tropical Life Sciences Research 26: 67-85.

Zhang, Y., K. Waiho, M. Ikhwanuddin and H. Ma. 2021a. Identification of sex-related genes from the three-spot swimming crab Portunus sanguinolentus and comparative analysis with the crucifix crab Charybdis feriatus. Animals 11: 1946. DOI: 10.3390/ani11071946.

Zhang, M., X. Zhang, N.T. Tran, S. Sun, X. Zhang, H. Ye, Y. Zhang, H. Ma, J.J. Aweya and S. Li. 2021b. Molting alters the microbiome, immune response, and digestive enzyme activity in mud crab (Scylla paramamosain). mSystems 6: e00917-21. DOI: 10.1128/mSystems.00917-21.