Giant tiger prawns are native to the coasts of the Arabian peninsula and the Pacific and Indian Ocean coasts of Australia, Indonesia, south and southeast Asia, and South Africa. They were accidentally introduced to the United States off the coast of South Carolina in 1988, by an unexpected release from an aquaculture center. They had spread as far south as Florida's coastline by 1990 and, since 2006, have been found in the Gulf of Mexico; they are found along the coastlines of Alabama, Florida, Georgia, Louisiana, Mississippi, North Carolina, South Carolina and Texas. (Duda and Palumbi, 1999; FAO Fisheries and Aquaculture Department, 2012; Knott, et al., 2011; South Carolina Department of Natural Resources, 2008; Xu, et al., 2001)
Young giant tiger prawns are most commonly found in estuaries, lagoons and mangroves; they are very tolerant to a range of salinity levels from 2-30 ppt. Adults move into deeper waters and live on rocky or muddy bottoms, ranging in depth from 0-110 m (most commonly at 20-50 m). These shrimps may bury themselves in the substrate during the day, emerging to feed at night. They live in waters ranging from 28-33°C and are unlikely to survive in waters colder than 13°C. (FAO Fisheries and Aquaculture Department, 2012; Knott, et al., 2011)
Giant tiger prawns have a typical prawn body plan including a head, tail, five pairs of swimming legs (pleopods) and five pairs of walking legs (pereopods), as well as numerous head appendages. A carapace (hard exoskeleton) encloses the cephalothorax. Their heads have a rostrum (an extension of the carapace in front of the eyes) and six to eight dorsal teeth, as well as two to four sigmoidally-shaped ventral teeth. A posterior ridge called the adrostral carina extends from the rostrum to the edge of the epigastric spine, which reaches to the posterior end of the carapace. Their first three pairs of pereopods have claws and they are distinguished from other shrimp species by the lack of an exopod (an external branch) on their fifth pleopodia. The telson at the posterior end of the prawn is unarmed, with no spines. (Dall, et al., 1991; FAO Fisheries and Aquaculture Department, 2012; State of New South Wales through Department of Industry and Investment, 2010)
Giant tiger prawns are identified by distinct black and white stripes on their backs and tails; on their abdomens, these stripes alternate black/yellow or blue/yellow. Base body color varies from green, brown, red, grey, or blue. These prawns are very large, reaching 330 mm or greater in length (largest individual found at 336 mm total length) and are sexually dimorphic, with females are larger than males. At sexual maturity, female carapace lengths range from 47-164 mm and their total lengths from 164-190 mm, while male carapace lengths fall between 37 and 71 mm, with total lengths of up to 134 mm. On average, females weigh 200-320 g and males weigh 100-170 g. (Dall, et al., 1991; Environmental Defense Fund, 2011; FAO Fisheries and Aquaculture Department, 2012; FAO-FIRA, 2010; Knott, et al., 2011; Primavera, et al., 1998; Vainio and Lagerspetz, 2006)
Females have a sperm receptacle (thyelycum) located ventrally on the last thoracic segment. After mating, sperm remain in this receptacle until eggs are released. Females have a pair of internal fused ovaries that extend almost the entire length of their bodies, from the cardiac region of the stomach to the anterior portion of the telson. Males have a copulatory organ (petasma, formed by the longitudinally folded endopods of the first pair of pleopods. The presence of an appendix masculina (an oval flap on the second pleopod) can distinguish males from females. Testes are unpigmented/translucent and are found dorsal to the hepatopancreas under the carapace. The vas deferens is also internal, and arises from the posterior margins of the main axis of the testes. Sperm are released through genital pores on the fifth pereopod. (Dall, et al., 1991; FAO-FIRA, 2010)
Eggs begin development by slowly sinking to the bottom of outer littoral areas. Giant tiger prawns develop through a complex life cycle beginning with three larval stages. Naupilii hatch twelve to fifteen hours after spawning is completed and look like tiny spiders. Larvae at this stage do not feed, instead surviving on their yolks as they are carried by tidal currents from open ocean towards shore. Naupilii larvae pass through six quick molts, increasing their body size. Individuals in the next larval stage, called protozoea, are identified by increased body size and length, the appearance of feathery appendages and, though still planktonic, beginning to feed. After molting three more times, protozoea proceed into the mysis larval stage. At this stage, they begin to have characteristics of adult prawns including segmented bodies, eye stalks, and tails. Mysis larvae molt three more times, becoming postlarvae. At this point in the life cycle, they change from planktonic to benthic feeding. This entire process takes two to three weeks. Prawns continue to molt through a juvenile phase, lasting 1-6 months. Juveniles and adults are distinguished mainly by location and carapace length. Carapace lengths of juveniles range from 2.2-11 mm and they are found mainly in estuarine areas located at the mouth or middle of bays and mangroves while adults are found in outer littoral areas of full salinity, and have carapace lengths ranging from 37-81 mm. (Dall, et al., 1991; FAO Fisheries and Aquaculture Department, 2012; FAO-FIRA, 2010; Motoh, 1981; State of New South Wales through Department of Industry and Investment, 2010)
Giant tiger prawns are known to mate prior to ovarian maturation; females store sperm in sacs within their closed thelycum until eggs are fully mature. Although little is known regarding specific mating behaviors, it has been noted that this species mates nocturnally, in off-shore waters, shortly after females have molted and their carapaces are still soft (males typically still have hard carapaces during breeding). Copulation begins with a male swimming parallel to a female. The male bends his body and first pair of pleopods with the petasma (caught by the appendix masculina) stretched vertically down, in order to facilitate the forward swinging of the second pair of pleopods. The first pair of pleopods pulls apart the petasmal halves, preventing the loss of sperm during copulation. The pair then takes an abdomen-to-abdomen position. The female exerts pressure on the male's petasma using her 4th pair of pereiopods and a spermatophore (sac of sperm) is thrust into her thyelycum, after which the pair separate. A majority of adult individuals copulate more than once; females are known to spawn 4 times during their lives, at carapace lengths of 50, 62, 66, and 72 mm. (Dall, et al., 1991; FAO Fisheries and Aquaculture Department, 2012; Motoh, 1981; Yano, et al., 1988)
It is difficult to estimate age at sexual maturity, but males become mature upon reaching an average carapace size of 37 mm, females at 47 mm. Females can produce 248,000-810,000 eggs at a time and are known to spawn up to four times during their lifespan. Once eggs are mature, they are expelled in a greenish-white cloud, along with stored spermatophores, into the ocean where external fertilization occurs. Eggs range in size from 0.27-0.31 mm. (Dall, et al., 1991; FAO Fisheries and Aquaculture Department, 2012; Motoh, 1981; New South Wales Government, 2009; State of New South Wales through Department of Industry and Investment, 2010; Yano, et al., 1988)
Males exhibit no parental involvement after mating. Females invest by yolking and protecting eggs while they are still in their bodies. They exhibit no further parental involvement once eggs and sperm have been released. (Dall, et al., 1991; Motoh, 1981)
The lifespan for wild and captive giant tiger prawns is about 2 years, though it has been suggested that individuals introduced into the Gulf of Mexico have a lifespan closer to 3 years. (Dall, et al., 1991; Institute for the Study of Invasive Species, 2011)
Giant tiger prawns are nocturnal feeders who often burrow into substrate during the day. They move about the ocean floor searching for food, which is picked up and manipulated by their pereopods and mouthparts. No published information regarding their social behaviors is currently available. (Dall, et al., 1991; FAO-FIRA, 2010)
No information regarding the average territory size for this species is currently available.
Giant tiger prawns have eyestalks on their heads which enable them to detect predators and search out prey. The eyes are called ommatidia, and are composed of clusters of photoreceptors. Since giant tiger prawns are nocturnal, they must have very good vision at night to detect predators and prey, but can also see well in daylight. Eyestalks have the ability to change their optical properties based on light-dark adaptations. In dark light, eyestalks receive light from a wide angle and create a superposition image, formed by mirrors in the sides of the cornea instead of by lenses. This superposition image is very effective at detecting movement. In bright light, eyestalks have the ability to see almost 360 degrees and form apposition images, a more efficient detector of light than superposition images. Molting Inhibition Hormone (MIH), which controls the molting cycle, is produced in the eyestalks; a recent study showed that when eyestalks are ablated, molting is accelerated. It is also known that ablating eyestalks in this species induces ovulation and jeopardizes growth. Giant tiger prawns also have flagellae on their antennae, which detect predators and prey through vibrations. These flagellae also have chemosensors, which detect amino acids and differences in pH, salinity and food stimulants. (Dall, et al., 1991; Uawisetwathana, et al., 2011)
In their first larval stage, giant tiger prawns feed on their yolk reserves. Later larval stages filter feed on plankton, diatoms, and other small organisms in the water column before becoming benthic feeders with a diet composed of organisms such as polycheate worms (Sabellaridae, Spionidae, Unicidae), as well as detritus. In the wild, adult giant tiger prawns feed on mollusks (including squid, blood clams (Arca sp.) and oysters), small crustaceans (including isopods, crabs and their eggs, and young penaeid prawns, including their own species). In aquaculture, these prawns feed on artificial diets consisting mainly of fishmeal; it has been noted that individuals grow more quickly when fed this diet. (Abu Hena and Hishamuddin, 2012; Dall, et al., 1991; FAO Fisheries and Aquaculture Department, 2012; FAO-FIRA, 2010; Thomas, 1972)
Throughout their lifetimes, giant tiger prawns face a variety of predators, including birds, comb jellies, crustaceans, and fishes. When adult prawns move from shallow inshore areas to deeper water, their rate of mortality drops. (Dall, et al., 1991; Primavera, 1997; Rajagopal, et al., 1995)
Giant tiger prawns have developed a variety of defenses to protect themselves from predation. Prawns have spines on either end of their body (a rostrum above the mouth, and a telson located at the dorsal end of the body). Their distinctive stripes and body color, which is similar to their muddy environment, help to camouflage them from predators. These prawns also bury themselves in substrate, not only hiding their bodies but also masking their waste, which would otherwise likely be detected by potential fish predators' chemosensory systems. (Dall, et al., 1991)
Giant tiger prawns are a host for a variety of viruses, all of which are extremely contagious within populations and cause high mortality rates. The Yellowhead virus, originally isolated from this species, causes the hepatopancreas and cephalothorax to become discoloured and swollen. WSSV (White Spot Syndrome Virus) causes white spot disease, symptoms of which include lesions and white deposits on the skin and connective tissue. There are two types of Baculovirus infections commonly seen in these prawns: Baculoviral Midgut Gland Necrosis, which affects mainly larvae, and Monodon baculovirus disease, which is typically followed by secondary bacterial infections. These diseases are of particular concern in aquaculture environments and in areas where this species has been introduced. (Crockford, 2008; FAO, 2001; Nadala, Jr., et al., 1997)
Giant tiger prawns are also host to a number of protozoan ectoparasites and endoparasites. Their ectoparasites attach themselves to the gills and limbs, potentially interfering with breathing and motility, while their endoparasites live in the gut and can affect nutrient absorption. This species is also known to host of a number of fungal microsporidians. (Chakraborti and Bandyapadhyay, 2011; Dash, et al., 2010; Toubiana, et al., 2004; Tourtip, et al., 2009)
Farming of Giant tiger prawns constitutes 47% of total world shrimp production giving it significant economic importance, particularly in Asian countries. With a high demand in Asian and international markets, building and running farms to produce these shrimp can be highly profitable and create many jobs. (FAO Fisheries and Aquaculture Department, 2012; Monterey Bay Aquarium, 2012)
This species is invasive in waters around the United States. Diseases carried by giant tiger prawns are highly contagious and can infect native shrimp populations, harming local fishing industries. (Institute for the Study of Invasive Species, 2011)
It has been estimated that up to 38% of native mangrove forests in Asia have been destroyed to be converted into ponds for shrimp farming, triggering erosion and harming habitat for mollusks and many other species, including shorebirds. Farming pools are sprayed with many chemicals and antibiotics to maximize shrimp production and these chemicals can enter natural waterways, harming animals and humans alike. These pools are often abandoned after a few years and there is typically no effort to return these lands to their original conditions. (FAO, 2001; GreenPeace, 2012)
This species is known by a variety of common names. The most common name is giant tiger prawn (shrimp). However, they are also called Asian prawn shrimp, ghost prawn, and grass shrimp. (FAO Fisheries and Aquaculture Department, 2012)
Jennifer Kiel (author), University of Michigan-Ann Arbor, Jeremy Wright (editor), University of Michigan-Ann Arbor.
the body of water between Africa, Europe, the southern ocean (above 60 degrees south latitude), and the western hemisphere. It is the second largest ocean in the world after the Pacific Ocean.
Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.
living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.
living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.
body of water between the southern ocean (above 60 degrees south latitude), Australia, Asia, and the western hemisphere. This is the world's largest ocean, covering about 28% of the world's surface.
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
Referring to an animal that lives on or near the bottom of a body of water. Also an aquatic biome consisting of the ocean bottom below the pelagic and coastal zones. Bottom habitats in the very deepest oceans (below 9000 m) are sometimes referred to as the abyssal zone. see also oceanic vent.
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
areas with salty water, usually in coastal marshes and estuaries.
an animal that mainly eats meat
flesh of dead animals.
uses smells or other chemicals to communicate
the nearshore aquatic habitats near a coast, or shoreline.
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
a substantial delay (longer than the minimum time required for sperm to travel to the egg) takes place between copulation and fertilization, used to describe female sperm storage.
an animal that mainly eats decomposed plants and/or animals
particles of organic material from dead and decomposing organisms. Detritus is the result of the activity of decomposers (organisms that decompose organic material).
animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature
an area where a freshwater river meets the ocean and tidal influences result in fluctuations in salinity.
fertilization takes place outside the female's body
parental care is carried out by females
union of egg and spermatozoan
a method of feeding where small food particles are filtered from the surrounding water by various mechanisms. Used mainly by aquatic invertebrates, especially plankton, but also by baleen whales.
A substance that provides both nutrients and energy to a living thing.
the area of shoreline influenced mainly by the tides, between the highest and lowest reaches of the tide. An aquatic habitat.
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.
eats mollusks, members of Phylum Mollusca
having the capacity to move from one place to another.
specialized for swimming
the area in which the animal is naturally found, the region in which it is endemic.
active during the night
an animal that mainly eats all kinds of things, including plants and animals
found in the oriental region of the world. In other words, India and southeast Asia.
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
photosynthetic or plant constituent of plankton; mainly unicellular algae. (Compare to zooplankton.)
an animal that mainly eats plankton
the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.
mainly lives in oceans, seas, or other bodies of salt water.
an animal that mainly eats dead animals
reproduction that includes combining the genetic contribution of two individuals, a male and a female
mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.
uses touch to communicate
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
movements of a hard surface that are produced by animals as signals to others
uses sight to communicate
breeding takes place throughout the year
2013. "Penaeus monodon Fabricius, 1798" (On-line). IUCN Red List of Threatened Species. Accessed January 30, 2013 at http://www.iucnredlist.org/search.
Abu Hena, M., O. Hishamuddin. 2012. Food selection preference of different ages and sizes of black tiger shrimp, Penaeus monodon Fabricius, in tropical aquaculture ponds in Malaysia. African Journal of Biotechnology, 11/22: 6153-6159. Accessed January 30, 2013 at http://www.academicjournals.org/ajb/PDF/pdf2012/15Mar/Abu%20Hena%20and%20Hishamuddin.pdf.
Chakraborti, J., P. Bandyapadhyay. 2011. Seasonal incidence of protozoan parasites of the black tiger shrimp (Penaeus monodon) of Sundarbans, West Bengal, India. Journal of Parasitic Diseases, 35/1: 61-65. Accessed January 30, 2013 at http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3114977/.
Crockford, M. 2008. "White Spot Disease" (On-line pdf). Australia and New Zealand Standard Diagnostic Procedures. Accessed January 30, 2013 at http://www.scahls.org.au/__data/assets/pdf_file/0009/1516518/White_Spot_Syndrome_Virus.pdf.
Dall, W., B. Hill, P. Rothlisberg, D. Sharples. 1991. Advances in Marine Biology. Queensland, Australia: Academic Press. Accessed March 15, 2012 at http://www.sciencedirect.com.proxy.lib.umich.edu/science/bookseries/00652881/27.
Dash, G., P. Yonzone, A. Roy. 2010. PREVALENCE AND SEASONAL ABUNDANCE OF PROTOZOAN PARASITES IN PENAEID SHRIMP PENAEUS MONODON IN HIGH SALINE BHERIES OF WEST BENGAL. Journal of Experimental Zoology of India, 13/2: 427-430. Accessed January 30, 2013 at http://www.connectjournals.com/file_html_pdf/787602H_j18_427a.pdf.
Duda, T., S. Palumbi. 1999. Population structure of the black tiger prawn, Penaeus monodon, among western Indian Ocean and western Pacific populations. Marine Biology, 134: 705-710. Accessed February 01, 2012 at http://www.springerlink.com/content/4avy2kyfja0967uf/.
Environmental Defense Fund, 2011. "Giant Tiger Prawn" (On-line). Accessed February 09, 2012 at http://apps.edf.org/page.cfm?tagID=15754.
FAO Fisheries and Aquaculture Department, 2012. "Fisheries and Aquaculture Department. About us - Fisheries and Aquaculture Department" (On-line). Accessed February 01, 2012 at http://www.fao.org/fishery/culturedspecies/Penaeus_monodon/en.
FAO, 2001. "Crustacean Disease" (On-line). Accessed February 22, 2012 at http://www.fao.org/docrep/005/y1679e/y1679e00.htm.
FAO-FIRA, 2010. "Giant Tiger Prawn Home" (On-line). Accessed February 10, 2012 at http://affris.org/giant_tiger_prawn/overview.php.
GreenPeace, 2012. "Shrimp Farming" (On-line). Accessed February 22, 2012 at http://www.greenpeace.org/international/en/campaigns/oceans/aquaculture/shrimp-farming/.
IUCN, 2012. "The IUCN Red List of Threatened Species" (On-line). Accessed February 04, 2013 at www.iucnredlist.org.
Institute for the Study of Invasive Species, 2011. "Penaeus monodon" (On-line). Accessed February 24, 2012 at http://www.tsusinvasives.org/database/black-tiger-shrimp.html.
Johnson, S. 1995. Handbook of Shrimp Diseases. Bryan, TX: Texas A&M University Sea Grant College Program. Accessed January 30, 2013 at http://nsgl.gso.uri.edu/tamu/tamuh95001.pdf.
Knott, D., P. Fuller, A. Benson, M. Neilson. 2011. "NAS - Nonindigenous Aquatic Species" (On-line). Accessed February 09, 2012 at http://nas.er.usgs.gov/queries/FactSheet.aspx?speciesID=1209.
Monterey Bay Aquarium, 2012. "Farmed Shrimp Seafood Watch" (On-line). Accessed February 09, 2012 at http://www.montereybayaquarium.org/cr/SeafoodWatch/web/sfw_factsheet.aspx?gid=58.
Motoh, H. 1981. Studies on the fisheries biology of the giant tigen prawn, Penaeus monodon in the Philippines. Tigbauan, Philippines: Aquaculture Dept., Southeast Asian Fisheries Development Center.
Nadala, Jr., E., L. Tapay, P. Loh. 1997. Yellow-head virus: a rhabdovirus-like pathogen of penaeid shrimp. Diseases of Aquatic Organisms, 31: 141-146. Accessed January 30, 2013 at http://www.int-res.com/articles/dao/31/d031p141.pdf.
New South Wales Government, 2009. "Prawns - aquaculture prospects" (On-line). Accessed February 10, 2012 at http://www.dpi.nsw.gov.au/fisheries/aquaculture/publications/species-saltwater/prawns.
Primavera, H. 1997. Fish predation on mangrove-associated penaeids The role of structures and substrate. Journal of Experimental Marine Biology and Ecology, 215: 205-216. Accessed January 30, 2013 at http://mangroveweb.seafdec.org.ph/pubs/Fish%20predation.pdf.
Primavera, J., F. Parado-Estepa, J. Lebata. 1998. Morphometric relationship of length and weight of giant tiger prawn Penaeus monodon according to life stage, sex and source. Aquaculture, Volume 164. Issues 1-4: 67-75. Accessed February 01, 2012 at http://www.sciencedirect.com/science/article/pii/S004484869800177X.
Rajagopal, S., M. Srinivasan, S. Khan. 1995. Problems in Culturing the Black Tiger Shrimp (Naga, 3: 29-30. Accessed January 30, 2013 at http://www.worldfishcenter.org/Naga/na_2235.pdf.) the Semi-intensive way: An Indian Experience.
South Carolina Department of Natural Resources, 2008. "South Carolina Aquatic Invasive Species MANAGEMENT PLAN" (On-line). Accessed February 09, 2012 at http://www.dnr.sc.gov/invasiveweeds/aisfiles/SCAISplan.pdf.
State of New South Wales through Department of Industry and Investment, 2010. "Biology and Life cycle of prawns" (On-line). Accessed February 10, 2012 at http://www.dpi.nsw.gov.au/fisheries/aquaculture/publications/species-saltwater/prawns.
Thomas, M. 1972. FOOD AND FEEDING HABITS OF PENAEUS MONODON FABRICIUS FROM KORAPUZHA ESTUARY. Indian Journal of Fisheries, Volume 19. Issue 1&2: 202-204.
Toubiana, M., O. Guelorget, J. Bouchereau, H. Lucien-Brun, A. Marques. 2004. Microsporidians in penaeid shrimp along the west coast of Madagascar. Diseases of Aquatic Organisms, 58: 79–82. Accessed January 30, 2013 at http://www.int-res.com/articles/dao2004/58/d058p079.pdf.
Tourtip, S., S. Wongtripop, G. Stentiford, K. Bateman, S. Sriurairatana, J. Chavadej, K. Sritunyalucksana, B. Withyachumnarnkul. 2009. Enterocytozoon hepatopenaei sp. nov. (Microsporida: Enterocytozoonidae), a parasite of the black tiger shrimp Penaeus monodon (Decapoda: Penaeidae): Fine structure and phylogenetic relationships. Journal of Invertebrate Pathology, 102: 21-29. Accessed January 30, 2013 at http://www.crustaceancrl.eu/publications/2009_%20JIP%20Enterocytozoon%20hepatopanaei.pdf.
Uawisetwathana, U., R. Leelatanawit, A. Klanchui, J. Prommoon, S. Klinbunga, N. Karoonuthaisiri. 2011. Insights into Eyestalk Ablation Mechanism to Induce Ovarian Maturation in the Black Tiger Shrimp. PLoS ONE, 6/9: doi:10.1371/journal.pone.0024427. Accessed January 29, 2012 at http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0024427.
Vainio, L., K. Lagerspetz. 2006. Thermal Behavior of Crustaceans. Biological Reviews, Vol. 81, Issue 2: 237-258. Accessed March 25, 2012 at http://www.aseanbiodiversity.info/Abstract/51008099.pdf.
Xu, Z., J. Primavera, L. de la Pena, P. Pettit, J. Belak, A. Alcivar-Warren. 2001. Genetic diversity of wild and cultured Black Tiger Shrimp (Penaeus monodon) in the Philippines using microsatellites. Aquaculture, 199: 13-40. Accessed February 01, 2012 at http://www.sciencedirect.com/science/article/pii/S0044848600005354.
Yano, I., R. Kanna, R. Oyama, J. Wyban. 1988. Mating behavior in the penaeid shrimp Pennaeus vannamei. Marine Biology, 97: 171-175. Accessed March 21, 2012 at http://wenku.baidu.com/view/6a6c9c6eaf1ffc4ffe47ac5f.html.