Shrimp

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For other uses, see Shrimp (disambiguation).

The shrimp Palaemon serratus of the infraorder Caridea

A shrimp (pl.: shrimp (US) or shrimps (UK)) is a crustacean (a form of shellfish) with an elongated body and a primarily swimming mode of locomotion – typically belonging to the Caridea or Dendrobranchiata of the order Decapoda, although some crustaceans outside of this order are also referred to as "shrimp".

More narrow definitions may be restricted to Caridea, to smaller species of either group or to only the marine species. Under a broader definition, shrimp may be synonymous with

swimmerets on the underside of their abdomens, although their escape response is typically repeated flicks with the tail driving them backwards very quickly. Crabs and lobsters have strong walking legs, whereas shrimp have thin, fragile legs which they use primarily for perching.[3]

Shrimp are widespread and abundant. There are thousands of species adapted to a wide range of habitats. They can be found feeding near the seafloor on most coasts and estuaries, as well as in rivers and lakes. To escape predators, some species flip off the seafloor and dive into the sediment.[3] They usually live from one to seven years.[4] Shrimp are often solitary, though they can form large schools during the spawning season.[3][5]

They play important roles in the

estuaries
when they are used to support shrimp farming.

Many shrimp species are small as the term shrimp suggests, about 2 cm (0.79 in) long, but some shrimp exceed 25 cm (9.8 in). Larger shrimp are more likely to be targeted commercially and are often referred to as prawns, particularly in the Commonwealth of Nations and former British colonies.

Classification

Shrimp and prawn

From Raymond Bauer in Remarkable Shrimps:[6]

  • Shrimp is characteristically used to refer to those crustaceans with long antennae, slender legs, and a laterally compressed, muscular abdomen that is highly adapted for both forward swimming and a backward (retrograde) escape response.
  • Prawn is often used as a synonym of shrimp for
    penaeoidean and caridean shrimp
    , especially those of large size.

From the English Oxford Dictionaries:

Shrimp are swimming

pleopods (swimmerets) and slender walking legs; they are more adapted for swimming than walking. Historically, it was the distinction between walking and swimming that formed the primary taxonomic division into the former suborders Natantia and Reptantia. Members of the Natantia (shrimp in the broader sense) were adapted for swimming while the Reptantia (crabs, lobsters, etc.) were adapted for crawling or walking.[9] Some other groups also have common names that include the word "shrimp";[10] any small swimming crustacean resembling a shrimp tends to be called one.[3]

Differences between shrimp, lobsters and crabs
Shrimp     shrimp
Clawed lobster     lobsters     Spiny lobster
Mud crab     crabs
Shrimp are slender with long muscular abdomens. They look somewhat like small lobsters, but not like crabs. The abdomens of crabs are small and short, whereas the abdomens of lobsters and shrimp are large and long. The lower abdomens of shrimp support pleopods which are well-adapted for swimming. The carapaces of crabs are wide and flat, whereas the carapaces of lobsters and shrimp are more cylindrical. The antennae of crabs are short, whereas the antennae of lobsters and shrimp are usually long, reaching more than twice the body length in some shrimp species.[3][10][11][12]
Clawed lobsters (pictured left) and spiny lobsters (pictured right) are an intermediate evolutionary development between shrimp and crabs. They look somewhat like large versions of shrimp. Clawed lobsters have large claws while spiny lobsters do not, having instead spiny antennae and carapace. Some of the biggest decapods are lobsters. Like crabs, lobsters have robust legs and are highly adapted for walking on the seafloor, though they do not walk sideways. Some species have rudimentary pleopods, which give them some ability to swim, and like shrimp they can lobster with their tail to escape predators, but their primary mode of locomotion is walking, not swimming.[3][10][11][13]
mineralized.[3][10][11][14][15]

Description

External anatomy of the common European shrimp, Crangon crangon
Shrimp can dart backwards by lobstering.

The following description refers mainly to the external anatomy of the common European shrimp,

compound eyes which have panoramic vision and are very good at detecting movement. Two pairs of whiskers (antennae) also issue from the head. One of these pairs is very long and can be twice the length of the shrimp, while the other pair is quite short. The antennae have sensors on them which allow the shrimp to feel where they touch, and also allow them to "smell" or "taste" things by sampling the chemicals in the water. The long antennae help the shrimp orient itself with regard to its immediate surroundings, while the short antennae help assess the suitability of prey.[3][10]

Eight pairs of appendages issue from the cephalothorax. The first three pairs, the

pereiopods. These form the ten decapod legs. In Crangon crangon, the first two pairs of pereiopods have claws or chela. The chela can grasp food items and bring them to the mouth. They can also be used for fighting and grooming. The remaining four legs are long and slender, and are used for walking or perching.[3][10][12][17]

The muscular abdomen has six segments and has a thinner shell than the carapace. Each segment has a separate overlapping shell, which can be transparent. The first five segments each have a pair of appendages on the underside, which are shaped like paddles and are used for swimming forward. The appendages are called

pleopods or swimmerets, and can be used for purposes other than swimming. Some shrimp species use them for brooding eggs, others have gills on them for breathing, and the males in some species use the first pair or two for insemination. The sixth segment terminates in the telson flanked by two pairs of appendages called the uropods. The uropods allow the shrimp to swim backward, and function like rudders, steering the shrimp when it swims forward. Together, the telson and uropods form a splayed tail fan. If a shrimp is alarmed, it can flex its tail fan in a rapid movement. This results in a backward dart called the caridoid escape reaction (lobstering).[3][10][12]

Habitat

Shrimp are widespread, and can be found near the seafloor of most coasts and estuaries, as well as in rivers and lakes. There are numerous species, and usually there is a species adapted to any particular habitat.

semi-terrestrial and spend a significant part of their life on land in mangrove.[19][20]

Habitats
Most shrimp such as Lysmata amboinensis live in fairly shallow waters and use their "walking legs" to perch on the sea bottom.
cherry shrimp
, perch on plant leaves

Red Cherry Shrimp – YouTube
Some shrimp live in deep and dark waters, such as this Heterocarpus ensifer.
Pederson's shrimp lives with anemone hosts, and cleans parasites from fish.[21]
 Scorpion Fish being cleaned by Pederson Shrimp – YouTube

Behaviour

There are many variations in the ways different types of shrimp look and behave. Even within the core group of

symbiotic relationship with sea slugs and sea cucumbers, and may help keep them clear of ectoparasites.[26]

Behaviour
Pistol shrimp
stun prey with "sonic booms", producing some of the loudest sounds in the ocean.

symbiotic
relationship.
yellow watchman goby
. The shrimp maintains a burrow for the pair and the goby is the lookout.
Living Together: Shrimp and Fish BBC: Trials of Life

Most shrimp are

necrotic tissue of the reef fish they groom.[16] Some species of shrimp are known to cannibalize others as well if other food sources are not readily available. In turn, shrimp are eaten by various animals, particularly fish and seabirds, and frequently host bopyrid parasites.[16]

Mating

Females of the freshwater shrimp Caridina ensifera are capable of storing sperm from multiple partners, and thus can produce progeny with different paternities.[27] Reproductive success of sires was found to correlate inversely with their genetic relatedness to the mother.[27] This finding suggests that sperm competition and/or pre- and post-copulatory female choice occurs. Female choice may increase the fitness of progeny by reducing inbreeding depression that ordinarily results from the expression of homozygous deleterious recessive mutations.[28]

Species

Decapods

There is little agreement among taxonomists concerning the

phylogeny of crustaceans.[29] Within the decapods, "every study gives totally different results. Nor do even one of these studies match any of the rival morphology studies".[30] Some taxonomists identify shrimp with the infraorder Caridea and prawns with the suborder Dendrobranchiata.[31] While different experts give different answers, there is no disagreement that the caridean species are shrimp.[3] There are over 3000 caridean species.[32] Occasionally they are referred to as "true shrimp".[33]

Traditionally,

paraphyletic; that is, it was thought that originally all decapods were like shrimp.[34]

However, classifications are now based on clades, and the paraphyletic suborder Natantia has been discontinued. "On this basis, taxonomic classifications now divide the order Decapoda into the two suborders: Dendrobranchiata for the largest shrimp clade, and Pleocyemata for all other decapods. The Pleocyemata are in turn divided into half a dozen infra-orders"[34]

Major shrimp groups of the Natantia
Order Suborder Infraorder Image
Extant species [32]
 Description
Decapoda Dendrobranchiata Dendrobranchiata: Giant tiger prawn (Penaeus monodon) 533
Penaeid shrimp

A particularly significant family in this suborder is the Penaeidae, often referred to as penaeid shrimp or penaeid prawn. Most commercially important species are in this family. See below.

The species in this suborder tend to be larger than the caridean shrimp species below, and many are commercially important. They are sometimes referred to as prawns. Dendrobranchiata, such as the
lamellar. The segments on their abdomens are even-sized, and there is no pronounced bend in the abdomen.[3][6][13][39]
Pleocyemata Caridea Caridea: Pink shrimp (Pandalus borealis) 3438 The numerous species in this infraorder are known as caridean shrimp, though only a few are commercially important. They are usually small, nocturnal, difficult to find (they burrow in the sediment), and of interest mainly to marine biologists. Caridean shrimp, such as the pink shrimp pictured, typically have two pairs of claws. Female carideans attach eggs to their pleopods and brood them there. The second abdominal segment overlaps both the first and the third segment, and the abdomen shows a pronounced caridean bend.[3][9][39][40]
Procarididea Procarididea: Procaris ascensionis 6 A minor sister group to the Caridea (immediately above)
Stenopodidea 71 Known as
banded coral shrimp (pictured) is popular in aquariums. The Stenopodidea are a much smaller group than the Dendrobranchia and Caridea, and have no commercial importance.[41]

Other decapod crustaceans also called shrimp, are the ghost or mud shrimp belonging to the infra-order

monophyly of the group is not certain; recent studies have suggested dividing the group into two infraorders, Gebiidea and Axiidea.[35]

Non-decapods

A shrimp seems to be almost any crustacean that isn't a lobster, barnacle, or crab

– Greg Jensen [3]

A wide variety of non-decapod crustaceans are also commonly referred to as shrimp. This includes the

seed shrimp have bivalved carapaces which they can open or close.[12] Krill resemble miniature shrimp, and are sometimes called "krill shrimp".[43][44]

Other species groups commonly known as shrimp
Class Image Group
Extant species
 Description
Branchiopoda Branchiopoda comes from the Greek branchia meaning gills, and pous meaning feet.[45] They have gills on their feet or mouthparts.[46]
brine shrimp 8 Brine shrimp belong to the genus
fish feed in aquaculture.[47] Brine shrimp are sold as novelty gifts under the marketing name Sea-Monkeys
.
clam shrimp 150 Clam shrimp belong to the group
bivalved
carapace which can open and close.
fairy shrimp
 300 Fairy shrimp belong to the class
fairy shrimp are herbivores, and eat only the algae in the plankton. Their eggs can survive drought and temperature extremes for years, reviving and hatching after the rain returns.[48]
tadpole shrimp
 20
longtail tadpole shrimp (pictured) has three eyes and up to 120 legs with gills on them.[50] It lives for 20–90 days. Different populations can be bisexual, unisexual or hermaphroditic
.
Malacostraca Malacostraca comes from the Greek malakós meaning soft and óstrakon meaning shell.[51] The name is misleading, since normally the shell is hard, and is soft only briefly after moulting.[52]
Lophogastrida 56 These marine
opossum shrimp, females lophogastrida carry a brood pouch.[53]
mantis shrimp 400 Mantis shrimp, so called because they resemble a
Stomatopoda. They grow up to 38 cm (15 in) long, and can be vividly coloured. Some have powerful spiked claws which they punch into their prey, stunning, spearing and dismembering them. They have been called "thumb splitters" because of the severe gashes they can inflict if handled carelessly.[54]
opossum shrimp
 1,000 Opossum shrimp belong to the order
anchialine caves.[53][55]
skeleton shrimp
 Skeleton shrimp, sometimes known as ghost shrimp, are amphipods. Their threadlike slender bodies allow them to virtually disappear among fine filaments in seaweed. Males are usually much larger than females.[56][57] For a good account of a specific species, see Caprella mutica.
Ostracoda
bivalves or clams
.
seed shrimp
 13,000
Ostracoda. This is a class of numerous small crustacean species which look like seeds, typically about one millimetre (0.04 in) in size. Their carapace looks like a clam shell, with two parts held together by a hinge to allow the shell to open and close. Some marine seed shrimp drift as pelagic plankton, but most live on the sea floor and burrow in the upper sediment layer. There are also freshwater and terrestrial species. The class includes carnivores, herbivores, filter feeders and scavengers.[58]

Some mantis shrimp are a foot long, and have bulging eyes, a flattened tail and formidable claws equipped with clubs or sharp spikes, which it can use to knock out its opponents.[12][54]

Examples of non-decapod "shrimp"
Some mantis shrimp knock prey out with powerful punches, and can crack a clam open.[12][54]
Skeleton shrimp
have threadlike slender bodies

Tadpole shrimp
can have up to 120 swimming legs

branchiopods, like this brine shrimp
, have eggs which survive prolonged drought conditions.
Branchiopod shrimp – YouTube

Human uses

History

FAO, 1950–2009[59]

In 1991, archeologists suggested that ancient raised paved areas near the coast in Chiapas, Mexico, were platforms used for drying shrimp in the sun, and that adjacent clay hearths were used to dry the shrimp when there was no sun.[60][61] The evidence was circumstantial, because the chitinous shells of shrimp are so thin they degrade rapidly, leaving no fossil remains. In 1985 Quitmyer and others found direct evidence dating back to 600 AD for shrimping off the southeastern coast of North America, by successfully identifying shrimp from the archaeological remains of their mandibles (jaws).[62][63][64] Clay vessels with shrimp decorations have been found in the ruins of Pompeii.[64] In the 3rd century AD, the Greek author Athenaeus wrote in his literary work, Deipnosophistae; "... of all fish the daintiest is a young shrimp in fig leaves."[65]

In North America,

moults. Modern industrial shrimping methods originated in this area.[64]

""For shrimp to develop into one of the world's most popular foods, it took the simultaneous development of the

Edward III received a request that he ban this new and destructive way of fishing.[66] In 1583, the Dutch banned shrimp trawling in estuaries.[67]

In the 1920s, diesel engines were adapted for use in shrimp boats.

Power winches were connected to the engines, and only small crews were needed to rapidly lift heavy nets on board and empty them. Shrimp boats became larger, faster, and more capable. New fishing grounds could be explored, trawls could be deployed in deeper offshore waters, and shrimp could be tracked and caught round the year, instead of seasonally as in earlier times. Larger boats trawled offshore and smaller boats worked bays and estuaries. By the 1960s, steel and fibreglass hulls further strengthened shrimp boats, so they could trawl heavier nets, and steady advances in electronics, radar, sonar, and GPS resulted in more sophisticated and capable shrimp fleets.[64]

As shrimp fishing methods industrialised, parallel changes were happening in the way shrimp were processed. "In the 19th century, sun dried shrimp were largely replaced by canneries. In the 20th century, the canneries were replaced with freezers."[64]

In the 1970s, significant shrimp farming was initiated, particularly in China. The farming accelerated during the 1980s as the quantity of shrimp demand exceeded the quantity supplied, and as excessive bycatch and threats to endangered sea turtles became associated with trawling for wild shrimp.[64] In 2007, the production of farmed shrimp exceeded the capture of wild shrimp.[59]

Commercial species

Although there are thousands of species of shrimp worldwide, only about 20 of these species are commercially significant. The following table contains the principal commercial shrimp, the seven most harvested species. All of them are decapods; most of them belong to the

penaeid shrimp
.

Principal commercial shrimp species
Group Common name Scientific name Description Max length (mm) Depth (m) Habitat
FAO
WoRMS
 2010 production (thousand tonnes)
wild farmed total
Dendrobranchiata Whiteleg shrimp
Litopenaeus vannamei
 (Boone, 1931) 		The most extensively farmed species of shrimp. 230 0–72 marine, estuarine [68][69] [70] 1 2721 2722
Giant tiger prawn Penaeus monodon Fabricius, 1798 336 0–110 marine, estuarine [71][72] [73] 210 782 992
Akiami paste shrimp Acetes japonicus Kishinouye, 1905 Most intensively fished species. They are small with black eyes and red spots on the
fermented.[74]
 30 shallow marine [75][76] [77] 574 574
Southern rough shrimp Trachysalambria curvirostris (Stimpson, 1860) Easier to catch at night, and fished only in waters less than 60 m (200 ft) deep.[78] Most of the harvest is landed in China.[79] 98 13–150 marine [80][81] [82] 294 294
Fleshy prawn
Fenneropenaeus chinensis
 (Osbeck, 1765) 		Trawled in Asia where it is sold frozen. Exported to Western Europe. Cultured by Japan and South Korea in ponds.[83] 183 90–180 marine [83][84] [85] 108 45 153
Banana prawn Fenneropenaeus merguiensis (De Man, 1888) Typically trawled in the wild and frozen, with most catches made by Indonesia. Commercially important in Australia, Pakistan and the Persian Gulf. Cultured in Indonesia and Thailand. In India it tends to be confused with
Fenneropenaeus indicus, so its economic status is unclear.[86]
 240 10–45 marine, estuarine [86][87] [88] 93 20 113
Caridea Northern prawn Pandalus borealis (Krøyer, 1838) Widely fished since the early 1900s in Norway, and later in other countries following Johan Hjort's practical discoveries of how to locate them. They have a short life which contributes to a variable stock on a yearly basis. They are not considered overfished. 165 20–1380 marine [89][90] [91] 361 361
All other species
1490
220
1710
Combined total 3129 3788 6917

Fishing

Double-rigged shrimp trawler with one net up and the other being brought aboard
Traditional shrimper with a shrimp net
FAO[59]
Main article: Shrimp fishery

Commercial techniques for catching wild shrimp include otter trawls, seines and shrimp baiting. A system of nets is used when trawling. Baited traps are common in parts of the Pacific Northwest.

Shrimp trawling can result in very high

cetaceans.[93] Bycatch is often discarded dead or dying by the time it is returned to the sea, and may alter the ecological balance in discarded regions.[94]
Worldwide, shrimp trawl fisheries generate about 2% of the world's catch of fish in weight, but result in more than one third of the global bycatch total.

The most extensively fished species are the

giant tiger prawn. Together these four species account for nearly half of the total wild capture. In recent years, the global capture of wild shrimp has been overtaken by the harvest from farmed shrimp.[59]

Farming

Tanks in a shrimp hatchery
The whiteleg shrimp (juvenile shown) has become the preferred species for shrimp farming.
FAO[59]
Main articles:
Prawn farming

A shrimp farm is an

China, Thailand, Indonesia, India and Vietnam. The other 25% are produced mainly in Latin America, where Brazil is the largest producer. By 2016, the largest exporting nation is India, followed by Ecuador, Thailand, Indonesia and China.[95]

As can be seen from the global production chart on the left, significant aquaculture production started slowly in the 1970s and then rapidly expanded during the 1980s. After a lull in growth during the 1990s, due to pathogens, production took off again and by 2007 exceeded the capture from wild fisheries. By 2010, the aquaculture harvest was 3.9 million tonnes, compared to 3.1 million tonnes for the capture of wild shrimp.[59]

In the earlier years of marine shrimp farming the preferred species was the large

sea cucumbers and some fish species, in integrated multi-trophic systems
.

The major producer of farmed shrimp is China. Other significant producers are Thailand, Indonesia, India, Vietnam, Brazil, Ecuador and Bangladesh. Most farmed shrimp is exported to the United States, the European Union and Japan,[98] also other Asian markets, including South Korea, Hong Kong, Taiwan and Singapore.[95]

Investigations by

giant tiger shrimp.[101]

As food

Shrimp tail ready for eating
Main article:
Shrimp as food
Wikibooks Cookbook has a recipe/module on

Shrimp are

meat of shrimp
is marketed.

As with other seafood, shrimp is high in

triglycerides.[105]

Ebiko - shrimp roe, sometimes translated as "shrimp flakes" - is used as one of the ingredients in the preparation of sushi.[106]

Shrimp and other

kosher and thus are forbidden in Jewish cuisine
.

Aquaria

Several types of shrimp are kept in home

Hymenocera picta
).

Freshwater aquaria variant shrimp come in many colours
cherry shrimp
.
Caridina cantonensis
crystal red bee shrimp has broad red and white stripes.

Shrimp versus prawn

The terms shrimp and prawn are

taxa, but are terms of convenience with little circumscriptional significance. There is no reason to avoid using the terms shrimp or prawn when convenient, but it is important not to confuse them with the names or relationships of actual taxa.[12]

According to the crustacean taxonomist Tin-Yam Chan, "The terms shrimp and prawn have no definite reference to any known taxonomic groups. Although the term shrimp is sometimes applied to smaller species, while prawn is more often used for larger forms, there is no clear distinction between both terms and their usage is often confused or even reverse in different countries or regions."

L. B. Holthuis noted that the terms prawn and shrimp were used inconsistently "even within a single region", generalising that larger species fished commercially were generally called shrimp in the United States, and prawns in other English-speaking countries, although not without exceptions.[110]

caridean shrimp
, and is rarely referred to as a prawn.

A lot of confusion surrounds the scope of the term shrimp. Part of the confusion originates with the association of smallness. That creates problems with shrimp-like species that are not small. The expression "jumbo shrimp" can be viewed as an oxymoron, a problem that does not exist with the commercial designation "jumbo prawns".[111]

The term shrimp originated around the 14th century with the Middle English shrimpe, akin to the Middle Low German schrempen, and meaning to contract or wrinkle; and the Old Norse skorpna, meaning to shrivel up, or skreppa, meaning a thin person.[112][113] It is not clear where the term prawn originated, but early forms of the word surfaced in England in the early 15th century as prayne, praine and prane.[114][115][116] According to the linguist Anatoly Liberman it is unclear how shrimp, in English, came to be associated with small. "No Germanic language associates the shrimp with its size... The same holds for Romance... it remains unclear in what circumstances the name was applied to the crustacean."[117]

Taxonomic studies in Europe on shrimp and prawns were shaped by the

common prawn, both found in huge numbers along the European coastlines. The common shrimp, Crangon crangon, was categorised in 1758 by Carl Linnaeus, and the common prawn, Palaemon serratus, was categorised in 1777 by Thomas Pennant. The common shrimp is a small burrowing species aligned with the notion of a shrimp as being something small, whereas the common prawn is much larger. The terms true shrimp or true prawn are sometimes used to mean what a particular person thinks is a shrimp or prawn.[12] This varies with the person using the terms. But such terms are not normally used in the scientific literature, because the terms shrimp and prawn themselves lack scientific standing. Over the years the way shrimp and prawn are used has changed, and nowadays the terms are almost interchangeable. Although from time to time some biologists declare that certain common names should be confined to specific taxa, the popular use of these names seems to continue unchanged.[12][118]


Fossils

Only 57 exclusively fossil species are known in the shrimp

Lower Jurassic, followed by specimens from the Cretaceous.[119]

See also

References

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Further reading

External links

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