Dendrobranchiata is a suborder of

decapods The Decapoda or decapods (literally "ten-footed") are an order of crustaceans within the class Malacostraca, including many familiar groups, such as crabs, lobsters, crayfish, shrimp and prawns. Most decapods are scavengers. The order is estim ...

, commonly known as prawns. There are 540 extant species in seven families, and a

fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...

extending back to the

Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...

. They differ from related animals, such as

Caridea The Caridea, commonly known as caridean shrimp or true shrimp, are an infraorder of shrimp within the order Decapoda. This infraorder contains all species of true shrimp. They are found widely around the world in both fresh and salt water. Many ot ...

and

Stenopodidea The Stenopodidea or boxer shrimps are a small group of decapod crustaceans. Often confused with Caridea shrimp or Dendrobranchiata prawns, they are neither, belonging to their own group. Anatomy They can be differentiated from the Dendrobranc ...

, by the branching form of the

gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...

s and by the fact that they do not brood their eggs, but release them directly into the water. They may reach a length of over and a mass of , and are widely fished and

farmed Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people t ...

for human consumption.

Shrimp and prawns

While Dendrobranchiata and

belong to different

suborder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...

s of

Decapoda The Decapoda or decapods (literally "ten-footed") are an order of crustaceans within the class Malacostraca, including many familiar groups, such as crabs, lobsters, crayfish, shrimp and prawns. Most decapods are scavengers. The order is esti ...

, they are very similar in appearance, and in many contexts such as commercial

farming Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people to ...

and

fisheries Fishery can mean either the enterprise of raising or harvesting fish and other aquatic life; or more commonly, the site where such enterprise takes place ( a.k.a. fishing ground). Commercial fisheries include wild fisheries and fish farms, both ...

, they are both often referred to as "shrimp" and "prawn" interchangeably. In the

United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the continental mainland. It comprises England, Scotland, Wales and North ...

, the word "prawn" is more common on menus than "shrimp", while the opposite is the case in

North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...

. The term "prawn" is also loosely used to describe any large shrimp, especially those that come 15 (or fewer) to the pound (such as "king prawns", yet sometimes known as "jumbo shrimp").

Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...

and some other

Commonwealth A commonwealth is a traditional English term for a political community founded for the common good. Historically, it has been synonymous with "republic". The noun "commonwealth", meaning "public welfare, general good or advantage", dates from the ...

nations follow this British usage to an even greater extent, using the word "prawn" almost exclusively. When Australian comedian

Paul Hogan Paul Hogan (born 8 October 1939) is an Australian actor and comedian. He was nominated for the Academy Award for Best Original Screenplay and won the Golden Globe Award for Best Actor – Motion Picture Musical or Comedy for his performance as ...

used the phrase, "I'll slip an extra shrimp on the barbie for you" in an American television advertisement, Baker & Bendel, 2007 it was intended to make what he was saying easier for his American audience to understand, and was thus a deliberate distortion of what an Australian would typically say.

Description

Together with other swimming Decapoda, Dendrobranchiata show the "caridoid facies", or shrimp-like form. Tavares & Martin, 2010, p. 100 The body is typically robust, and can be divided into a

cephalothorax The cephalothorax, also called prosoma in some groups, is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind. (The terms ''prosoma'' and ''opisthosoma'' are equivalent to ''cepha ...

(head and thorax fused together) and a

pleon The decapod ( crustaceans such as a crab, lobster, shrimp or prawn) is made up of 20 body segments grouped into two main body parts: the cephalothorax and the pleon (abdomen). Each segment may possess one pair of appendages, although in various ...

(abdomen). The body is generally slightly flattened side-to-side. The largest species, ''

Penaeus monodon ''Penaeus monodon'', commonly known as the giant tiger prawn, Asian tiger shrimp, black tiger shrimp, and other names, is a marine crustacean that is widely reared for food. Taxonomy ''Penaeus monodon'' was alpha taxonomy, first described by J ...

'', can reach a

mass Mass is an intrinsic property of a body. It was traditionally believed to be related to the quantity of matter in a physical body, until the discovery of the atom and particle physics. It was found that different atoms and different elementar ...

of and a length of . Dall, 1990, pp. 3–4

Head

The most conspicuous appendages arising from the head are the antennae. The first pair are biramous (having two

flagella A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...

), except in

Luciferidae Luciferidae is a family of prawns. These prawns are small, characterised by bioluminescence and the loss or reduction of some appendages. They are predators of tiny planktonic crustaceans for which their third pereiopod is adapted to capture b ...

, and are relatively small. Tavares & Martin, 2010, p. 106 The second pair can be 2–3 times the length of the body and are always uniramous (having a single flagellum). The mouthparts comprise pairs of

mandibles In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...

, maxillules and maxillae, arising from the head, and three pairs of maxillipeds, arising from the thorax. Tavares & Martin, 2010, pp. 106–108 A pair of stalked

eye Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect light and conv ...

s points forwards from the head. Tavares & Martin, 2010, p. 102

Thorax

The

carapace A carapace is a Dorsum (biology), dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tor ...

grows from the thorax to cover the cephalothorax, and extends forwards between the eyes into a

rostrum Rostrum may refer to: * Any kind of a platform for a speaker: **dais **pulpit * Rostrum (anatomy), a beak, or anatomical structure resembling a beak, as in the mouthparts of many sucking insects * Rostrum (ship), a form of bow on naval ships * Ros ...

. This is only as long as the stalked eyes in

Benthesicymidae Benthesicymidae is a family of shrimps in the suborder Dendrobranchiata. References External links * * Benthesicymidaeat WoRMS Dendrobranchiata Decapod families {{Dendrobranchiata-stub ...

and

Sergestidae Sergestidae is a family of prawns which have lived since at least the Middle Jurassic (Bajocian/Bathonian). It contains the following genera: *'' Acetes'' H. Milne-Edwards, 1830 *'' Allosergestes'' Judkins & Kensley, 2008 *'' Casertanus'' Bravi ...

, but considerably longer in

Aristeidae Aristeidae is a family of Dendrobranchiata decapod crustaceans known as deep-sea shrimps, gamba prawns or gamba shrimps. Some species are subject to commercial fisheries. Genera The following genera are classified under the Aristeidae: *'' Ar ...

. As well as the three pairs of maxillipeds, the thorax also bears five pairs of

pereiopod The decapod (crustaceans such as a crab, lobster, shrimp or prawn) is made up of 20 body segments grouped into two main body parts: the cephalothorax and the pleon (abdomen). Each segment may possess one pair of appendages, although in various g ...

s, or walking legs; the first three of these end in small

chelae A chela ()also called a claw, nipper, or pinceris a pincer (biology), pincer-like organ at the end of certain limbs of some arthropods. The name comes from Ancient Greek , through New Latin '. The plural form is chelae. Legs bearing a chela are ...

(pincers). Tavares & Martin, 2010, pp. 108–110 The last two pereiopods are absent in

and ''

Acetes ''Acetes'' is a genus of small shrimp that resemble krill, which is native to the western and central Indo-Pacific, the Atlantic coast of the Americas, Pacific coast of South America and inland waters of South America. Although most are from mar ...

'', but much longer than the preceding pereiopods in '' Hymenopenaeus'' and ''

Xiphopenaeus ''Xiphopenaeus'' is a genus of crustaceans in the suborder Dendrobranchiata, the shrimps and prawns. Two species are in this genus: *''Xiphopenaeus kroyeri'' *'' Xiphopenaeus riveti'' Likely, more species have not yet been described and named. ...

''. Tavares & Martin, 2010, p. 110 The thoracic appendages carry

s, which are protected beneath the carapace. Tavares & Martin, 2010, pp. 103–105 The gills are typically branched, and so resemble trees, lending the group its scientific name, Dendrobranchiata, from the

Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...

words (',

tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...

) and (', gills). Tavares & Martin, 2010, p. 103

Pleon

The

, or abdomen, is similar in length to the cephalothorax. Tavares & Martin, 2010, p. 105 It has six segments, the first five bearing lamellar

pleopod The decapod ( crustaceans such as a crab, lobster, shrimp or prawn) is made up of 20 body segments grouped into two main body parts: the cephalothorax and the pleon (abdomen). Each segment may possess one pair of appendages, although in various ...

s, and the last one bearing

uropod Uropods are posterior appendages found on a wide variety of crustaceans. They typically have functions in locomotion. Definition Uropods are often defined as the appendages of the last body segment of a crustacean. An alternative definition sugge ...

s. Tavares & Martin, 2010, pp. 110–111 The pleopods are biramous, except in

Sicyoniidae ''Sicyonia'' is a genus of prawns, placed in its own family, Sicyoniidae. It differs from other prawns in that the last three pairs of its pleopods are uniramous, rather than biramous as seen in all other prawns. ''Sicyonia'' contains 52 extant ...

, where they are uniramous. The uropods and

telson The telson () is the posterior-most division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment of the arthropod body, or an additional division that is not a true segment on accou ...

collectively form the tail fan; the uropods are not divided by a diaeresis, as they are in many other decapods. Tavares & Martin, 2010, p. 111 The telson is pointed and is usually armed with four pairs of

seta In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...

e or spines.

Internal anatomy

Most of the

musculature Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of muscle ...

of a prawn is used for bending the pleon, and almost all the space in the pleon is filled by muscle. Tavares & Martin, 2010, p. 113 More than 17 muscles operate each of the pleopods, and a further 16 power the tail fan in the rapid backward movement of the

caridoid escape reaction The caridoid escape reaction, also known as lobstering or tail-flipping, refers to an innate escape mechanism in marine and freshwater crustaceans such as lobster Lobsters are a family (biology), family (Nephropidae, Synonym (taxonomy), sy ...

. Tavares & Martin, 2010, pp. 113–114 These muscles, collectively, are the meat for which prawns are commercially fished and farmed. The

nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes th ...

of prawns comprises a dorsal

brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...

, and a

ventral nerve cord The ventral nerve cord is a major structure of the invertebrate central nervous system. It is the functional equivalent of the vertebrate spinal cord. The ventral nerve cord coordinates neural signaling from the brain to the body and vice versa, in ...

, connected by two

commissure A commissure () is the location at which two objects abut or are joined. The term is used especially in the fields of anatomy and biology. * The most common usage of the term refers to the brain's commissures, of which there are five. Such a commi ...

s around the

oesophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...

. Tavares & Martin, 2010, p. 114 The chief sensory inputs are visual input from the eyes,

chemoreceptor A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance (endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorecept ...

s on the antennae and in the mouth, and

mechanoreceptor A mechanoreceptor, also called mechanoceptor, is a sensory receptor that responds to mechanical pressure or distortion. Mechanoreceptors are innervated by sensory neurons that convert mechanical pressure into electrical signals that, in animals, ...

s on the antennae and elsewhere. Tavares & Martin, 2010, pp. 116–118 The

digestive system The human digestive system consists of the gastrointestinal tract plus the accessory organs of digestion (the tongue, salivary glands, pancreas, liver, and gallbladder). Digestion involves the breakdown of food into smaller and smaller compone ...

comprises a foregut, a midgut and a hindgut, and is situated dorsally. Tavares & Martin, 2010, p. 118 The foregut begins at the mouth, passes through the oesophagus, and opens into a sac that contains the grinding apparatus of the

gastric mill The gizzard, also referred to as the ventriculus, gastric mill, and gigerium, is an organ found in the digestive tract of some animals, including archosaurs (pterosaurs, crocodiles, alligators, dinosaurs, birds), earthworms, some gastropods, so ...

. The

hepatopancreas The hepatopancreas, digestive gland or midgut gland is an organ of the digestive tract of arthropods and molluscs. It provides the functions which in mammals are provided separately by the liver and pancreas, including the production of digestive ...

feeds into the midgut, where

digestive enzyme Digestive enzymes are a group of enzymes that break down polymeric macromolecules into their smaller building blocks, in order to facilitate their absorption into the cells of the body. Digestive enzymes are found in the digestive tracts of anim ...

s are released, and nutrients taken up. The hindgut forms faecal pellets, which are then passed out through the muscular

anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, d ...

. Tavares & Martin, 2010, pp. 118–119 The

circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...

is based around a compact, triangular

heart The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide t ...

, which pumps blood into three main

arteries An artery (plural arteries) () is a blood vessel in humans and most animals that takes blood away from the heart to one or more parts of the body (tissues, lungs, brain etc.). Most arteries carry oxygenated blood; the two exceptions are the pul ...

. Tavares & Martin, 2010, p. 120

Excretion Excretion is a process in which metabolic waste is eliminated from an organism. In vertebrates this is primarily carried out by the lungs, kidneys, and skin. This is in contrast with secretion, where the substance may have specific tasks after lea ...

is carried out through the gills, and by specialised

gland In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream (endocrine gland) or into cavities inside the body or its outer surface (exocrine gland). Structure De ...

s located at the base of the antennae, and is mostly in the form of

ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous was ...

. Tavares & Martin, 2010, pp. 120–121

Life cycle

Prawns may be divided into two groups: those with an open thelycum (female genitalia) and those with a closed thelycum. Tavares & Martin, 2010, p. 125 In the open–thelycum species, mating takes place towards the end of the moulting cycle, and usually at sunset. Tavares & Martin, 2010, pp. 125–126 In closed–thelycum species, mating takes place shortly after moulting, when the

exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...

is still soft, and usually occurs in the night. Tavares & Martin, 2010, p. 126

Courtship Courtship is the period wherein some couples get to know each other prior to a possible marriage. Courtship traditionally may begin after a betrothal and may conclude with the celebration of marriage. A courtship may be an informal and private m ...

and mating may take up to 3 hours in ''

'', while in ''

Farfantepenaeus paulensis ''Farfantepenaeus'' is a genus of prawns in the family Penaeidae. Its eight species were formerly included in the genus ''Penaeus''. It was first published as a genus name in 1972 by Rudolf N. Burukovsky, but without the necessary designation of ...

'', mating lasts just 4–5 seconds. Spawning may occur several times during the moulting cycle, and usually occurs at night. Tavares & Martin, 2010, p. 127 With the exception of

, the

eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...

of prawns are shed directly into the water, rather than being brooded. Tavares & Martin, 2010, p. 130 The eggs hatch into nauplius

larvae A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...

, which are followed by

zoea Crustaceans may pass through a number of larval and immature stages between hatching from their eggs and reaching their adult form. Each of the stages is separated by a moult, in which the hard exoskeleton is shed to allow the animal to grow. The ...

larvae (initially protozoea, and later mysis) and then a postlarva, before reaching adulthood. The changes between moults are gradual, and so the development is

anamorphic Anamorphic format is the cinematography technique of shooting a widescreen picture on standard 35 mm film or other visual recording media with a non-widescreen native aspect ratio. It also refers to the projection format in which a distorted ...

rather than

metamorphic Metamorphic rocks arise from the transformation of existing rock to new types of rock in a process called metamorphism. The original rock (protolith) is subjected to temperatures greater than and, often, elevated pressure of or more, causin ...

. Uniquely among the Decapoda, the nauplii of Dendrobranchiata are free-swimming. There are five to eight naupliar stages. Tavares & Martin, 2010, p. 131 The earlier stages have three pairs of appendages that are used for locomotion – two pairs of antennae and the

. Later stages also have rudiments of other mouthparts, but the nauplius is unable to feed, and only lasts 24 to 68 hours. The body ends at a two-lobed

, and the beginnings of a

emerge at this stage. There are typically 5 or 6 zoea stages in Dendrobranchiata, divided into protozoea and mysis. In the protozoea larvae, the antennae are still used for locomotion, but the mandibles become specialised for

mastication Chewing or mastication is the process by which food is crushed and ground by teeth. It is the first step of digestion, and it increases the surface area of foods to allow a more efficient break down by enzymes. During the mastication process, th ...

. All the thoracic somites (body segments) have formed, and a

is present, covering part of the thorax. It is smooth in the family

Penaeidae Penaeidae is a family of marine crustaceans in the suborder Dendrobranchiata, which are often referred to as penaeid shrimp or penaeid prawns. The Penaeidae contain many species of economic importance, such as the tiger prawn, whiteleg shrimp, ...

, but bears many spines in the family

Solenoceridae Solenoceridae is a family of decapods The Decapoda or decapods (literally "ten-footed") are an order (biology), order of crustaceans within the class Malacostraca, including many familiar groups, such as crabs, lobsters, crayfish, Caridea, s ...

. The

(abdomen) is unsegmented in the first protozoea, and ends in a bilobed telson, which may be used for cleaning other appendages, or for steering. By the second protozoea, segmentation appears on the pleon, and by the third protozoea, which may also be called the metazoea, the

s have appeared. By the mysis stages, the

s (thoracic appendages) start to be used instead of the antennae for locomotion. Tavares & Martin, 2010, p. 133 The larva swims backwards, with its tail upwards, spinning slowly as it goes. The carapace covers most of the segments of the thorax, and claws appear on the first three pereiopods. By the last mysis stage, the beginnings of

s have appeared on the first five segments of the abdomen. The post-larva or

juvenile Juvenile may refer to: *Juvenile status, or minor (law), prior to adulthood *Juvenile (organism) *Juvenile (rapper) (born 1975), American rapper * ''Juvenile'' (2000 film), Japanese film * ''Juvenile'' (2017 film) *Juvenile (greyhounds), a greyho ...

stage is characterised by the use of the pleopods for locomotion. Tavares & Martin, 2010, p. 134 The claws become functional, but the

s are still rudimentary. The telson is narrower and only retains traces of its two-lobed development. Through a series of gradual changes over following moults, the animal takes on its adult form.

Systematics

Dendrobranchiata were traditionally grouped together with Caridea as "Natantia" (the swimming decapoda), as opposed to the

Reptantia Reptantia is a clade of decapod crustaceans named in 1880 which includes lobsters, crabs and many other well-known crustaceans. Classification In older classifications, Reptantia was one of the two sub-orders of Decapoda alongside Natantia, wi ...

(the walking decapods). In 1888,

Charles Spence Bate Charles Spence Bate, FRS (March 16, 1819 – July 29, 1889) was a British zoologist and dentist. Life He was born at Trenick House near Truro, the son of Charles Bate (1789–1872) and Harriet Spence (1788–1879). Charles adopted "Spence Bate" ...

recognised the differences in gill morphology, and separated Natantia into Dendrobranchiata, Phyllobranchiata and Trichobranchiata. Tavares & Martin, 2010, p. 99 Recent analyses using

cladistics Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...

and

molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

recognise Dendrobranchiata as the

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...

to all other Decapoda, collectively called

Pleocyemata Pleocyemata is a suborder of Decapoda, decapod crustaceans, erected by Martin Burkenroad in 1963. Burkenroad's classification replaced the earlier sub-orders of Natantia and Reptantia with the monophyletic groups Dendrobranchiata (prawns) and Ple ...

. Tavares & Martin, 2010, p. 137 The

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...

below shows Dendrobranchiata's placement within the larger

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...

, from analysis by Wolfe ''et al.'', 2019. 9150 - Milano - Museo storia naturale - Aeger - Foto Giovanni Dall'Orto 22-Apr-2007 cropped

Before 2010, the earliest known fossil prawns come from rocks in

Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...

of Permo-

Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...

age, . Schram ''et al.'', 2000 In 2010, however, the discovery of ''

Aciculopoda ''Aciculopoda'' is an extinct prawn which existed in what is now Oklahoma approximately . It was described in 2010 on the basis of a single fossil from Oklahoma. The single species, ''Aciculopoda mapesi'', was named by Rodney Feldmann and Carr ...

'' from

Famennian The Famennian is the latter of two faunal stages in the Late Devonian Epoch. The most recent estimate for its duration estimates that it lasted from around 371.1 million years ago to 359.3 million years ago. An earlier 2012 estimate, still used b ...

–stage rocks in

Oklahoma Oklahoma (; Choctaw language, Choctaw: ; chr, ᎣᎧᎳᎰᎹ, ''Okalahoma'' ) is a U.S. state, state in the South Central United States, South Central region of the United States, bordered by Texas on the south and west, Kansas on the nor ...

extended the group's fossil record back to . Feldmann & Schweitzer, 2010 The best known fossil prawns are from the

Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...

Solnhofen limestones from

Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...

. Living prawns are divided among seven families, five in the

superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...

Penaeoidea Penaeoidea is the larger of the two superfamilies of prawns. It comprises eight families, three of which are known only from fossils. The fossil record of the group stretches back to '' Aciculopoda'', discovered in Famennian sediments in Oklahom ...

, and two in the

Sergestoidea Sergestoidea is a superfamily of prawns, divided into two families – the Luciferidae and the Sergestidae Sergestidae is a family of prawns which have lived since at least the Middle Jurassic (Bajocian/Bathonian). It contains the following g ...

, although molecular evidence disagrees with some aspects of the current classifications. Ma ''et al.'', 2009 Collectively, these include 540

extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...

species, and nearly 100 exclusively fossil species. De Grave ''et al.'', 2009 A further two families are known only from fossils. The cladogram below shows Dendrobranchiata's internal relationships of

families (excluding

):
Dendrobranchiata comprises the following superfamilies and

families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideall ...

:
:Superfamily

Rafinesque-Schmaltz, 1815 ::† Aciculopodidae Feldmann & Schweitzer, 2010 – a single

species, '' Aciculopoda mapesi'' ::†

Aegeridae Aegeridae is a family of fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of ani ...

Burkenroad, 1963 – two Mesozoic genera: ''

Aeger ''Aeger'' is a genus of fossil prawns. They first occur in the Middle Triassic, and died out at the end of the Late Cretaceous. A total of 21 species are known. Species * '' Aeger brevirostris'' * '' Aeger brodiei'' * ''Aeger elegans'' * '' Aeg ...

'' and ''

Acanthochirana ''Acanthochirana'' is an extinct genus of prawn that existed during the upper Jurassic period. It was named by E. Strand in 1928, and its type species is ''Acanthochirana cordata''. They are distinguished from the related genus '' Aeger'' by the ...

'' Tavares & Martin, 2010, p. 151 ::

Wood-Mason, 1891 – 26 extant species in 9 genera, and one fossil genus Tavares & Martin, 2010, p. 152 ::

Wood-Mason, 1891 – 41 species in 4 genera ::† Carpopenaeidae Garassino, 1994 – two

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...

species of ''

Carpopenaeus ''Carpopenaeus'' is an extinct genus of prawn, which existed during the Upper Jurassic and Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Me ...

'' Tavares & Martin, 2010, pp. 152–153 ::

Rafinesque-Schmaltz, 1815 – 216 extant species in 26 genera, and several extinct genera, mostly Mesozoic Tavares & Martin, 2010, p. 153 ::

Ortmann, 1898 – 43 species of ''

Sicyonia ''Sicyonia'' is a genus of prawns, placed in its own family, Sicyoniidae. It differs from other prawns in that the last three pairs of its pleopods are uniramous, rather than biramous as seen in all other prawns. ''Sicyonia'' contains 52 extant ...

'' Tavares & Martin, 2010, p. 154 ::

Wood-Mason, 1891 – 81 species in 9 genera Tavares & Martin, 2010, p. 155 :Superfamily

Dana, 1852 ::

De Haan, 1849 – 7 species in 2 genera ::

Dana, 1852 – 90 extant species in six genera, and two extinct monotypic genera Tavares & Martin, 2010, p. 156

Distribution

The

biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') l ...

of Dendrobranchiata decreases markedly at increasing

latitude In geography, latitude is a coordinate that specifies the north– south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from –90° at the south pole to 90° at the north pol ...

s; most species are only found in a region between 40° north and 40° south. Tavares & Martin, 2010, p. 145 Some species may occur at higher latitudes. For instance, '' Bentheogennema borealis'' is abundant at 57° north in the

Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the continen ...

, while collections of '' Gennadas kempi'' have been made as far south as 61° south in the

Antarctic Ocean The Southern Ocean, also known as the Antarctic Ocean, comprises the southernmost waters of the World Ocean, generally taken to be south of 60° S latitude and encircling Antarctica. With a size of , it is regarded as the second-small ...

Ecology and behaviour

Messina Straits Argyropelecus hemigymnus

There is a great deal of ecological variation within the suborder Dendrobranchiata. Some species of Sergestidae live in

fresh water Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does include ...

, but most prawns are exclusively marine. Species of

and

mostly live in deep water, and

species live offshore, while most

species live in shallow inshore waters, and ''

Lucifer Lucifer is one of various figures in folklore associated with the planet Venus. The entity's name was subsequently absorbed into Christianity as a name for the devil. Modern scholarship generally translates the term in the relevant Bible passage ...

'' is

plankton Plankton are the diverse collection of organisms found in Hydrosphere, water (or atmosphere, air) that are unable to propel themselves against a Ocean current, current (or wind). The individual organisms constituting plankton are called plankt ...

ic. Some species burrow in mud on the sea floor during the day and emerge at night to feed. Prawns are "opportunistic omnivores", and their

diet Diet may refer to: Food * Diet (nutrition), the sum of the food consumed by an organism or group * Dieting, the deliberate selection of food to control body weight or nutrient intake ** Diet food, foods that aid in creating a diet for weight loss ...

can include a range of food items from fine particles to large organisms. These may include

fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...

chaetognath The Chaetognatha or chaetognaths (meaning ''bristle-jaws'') are a phylum of predatory marine worms that are a major component of plankton worldwide. Commonly known as arrow worms, about 20% of the known Chaetognatha species are benthic, and can ...

krill Krill are small crustaceans of the order Euphausiacea, and are found in all the world's oceans. The name "krill" comes from the Norwegian Norwegian, Norwayan, or Norsk may refer to: *Something of, from, or related to Norway, a country in n ...

copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...

radiolaria The Radiolaria, also called Radiozoa, are protozoa of diameter 0.1–0.2 mm that produce intricate mineral skeletons, typically with a central capsule dividing the cell (biology), cell into the inner and outer portions of endoplasm and Ecto ...

ns,

phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. Ph ...

nematocyst A cnidocyte (also known as a cnidoblast or nematocyte) is an explosive cell containing one large secretory organelle called a cnidocyst (also known as a cnida () or nematocyst) that can deliver a sting to other organisms. The presence of this ce ...

ostracod Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several orders. They are small crustaceans, typic ...

s and

detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts commun ...

. Tavares & Martin, 2010, p. 135 Prawns eat less around the time of

ecdysis Ecdysis is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remna ...

(moulting), probably because of the softness of the mouthparts, and must eat more than usual to compensate, once ecdysis is complete. Prawns are an attractive food for

predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...

s, with a higher

energy content In physics, energy density is the amount of energy stored in a given system or region of space per unit volume. It is sometimes confused with energy per unit mass which is properly called specific energy or . Often only the ''useful'' or extrac ...

than most other invertebrates. Dall, 1990, p. 357 The larvae are prey to

comb jellies Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), an ...

jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbrella- ...

and other

crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...

s (such as

mantis shrimp Mantis shrimp, or stomatopods, are carnivorous marine crustaceans of the order Stomatopoda (). Stomatopods branched off from other members of the class Malacostraca around 340 million years ago. Mantis shrimp typically grow to around in lengt ...

and

crab Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen) ( el, βραχύς , translit=brachys = short, / = tail), usually hidden entirely under the thorax. They live in all the ...

s), and only a tiny proportion survive. Dall, 1990, p. 358 Juveniles are targeted by a number of fish,

cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...

s and

bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...

s; ''

Litopenaeus vannamei Whiteleg shrimp (''Litopenaeus vannamei'', synonym ''Penaeus vannamei''), also known as Pacific white shrimp or King prawn, is a species of prawn of the eastern Pacific Ocean commonly caught or farmed for food. Description ''L. vannamei'' grows ...

'' juveniles experience 90% mortality in the 6–12 weeks they spend in Mexican

lagoon A lagoon is a shallow body of water separated from a larger body of water by a narrow landform, such as reefs, barrier islands, barrier peninsulas, or isthmuses. Lagoons are commonly divided into ''coastal lagoons'' (or ''barrier lagoons'') a ...

s, and this is thought to be due almost entirely to predation. Adult prawns are less susceptible to predation, but can fall prey to some

. Dall, 1990, p. 359

Economic importance

Dendrobranchiata are of huge importance. While in some countries, such as the

United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...

, production is almost entirely through

, other countries have concentrated on

aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. lot ...

(

shrimp farm Shrimp farming is an aquaculture business that exists in either a marine or freshwater environment, producing shrimp or prawns (crustaceans of the groups Caridea or Dendrobranchiata) for human consumption. Marine Commercial marine shrimp farming ...

s), including

Ecuador Ecuador ( ; ; Quechua: ''Ikwayur''; Shuar: ''Ecuador'' or ''Ekuatur''), officially the Republic of Ecuador ( es, República del Ecuador, which literally translates as "Republic of the Equator"; Quechua: ''Ikwadur Ripuwlika''; Shuar: ''Eku ...

where 95% of production is

; some countries produce similar amounts from fisheries and aquaculture, including

Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...

China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...

India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...

and

Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...

. Tavares & Martin, 2010, p. 136 Species from the family

are important to deep-water fisheries, particularly in the

Mediterranean Sea The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the ea ...

, where ''

Aristaeomorpha foliacea ''Aristaeomorpha foliacea'', the giant red shrimp or giant gamba prawn, is a species of deep water benthopelagic decapod crustacean. It is found in all the world's oceans in the temperate and tropical zones. It is subject to some commercial fish ...

'' is caught by trawlers. In

Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...

, ''Aristaeomorpha foliacea'', '' Aristaeopsis edwardsiana'' and '' Aristeus antillensis'' are of commercial importance. The shallow-water

are of greater importance, however, and the most important species for fisheries is ''

Fenneropenaeus chinensis The Chinese white shrimp, oriental shrimp, or fleshy prawn (''Fenneropenaeus chinensis'') is a species of shrimp. It is cultivated at an industrial level off mainland China. Production was devastated by a series of epidemics in the 1990s and ear ...

'', with a catch in 2005 of over 100,000 tons. The most important species for

are ''

Marsupenaeus japonicus ''Marsupenaeus'' is a monotypic genus of prawn. It contains a single species, ''Marsupenaeus japonicus'', known as the kuruma shrimp, kuruma prawn, or Japanese tiger prawn. It occurs naturally in bays and seas of the Indo-West Pacific, but has a ...

'' (Kuruma prawn), ''Fenneropenaeus chinensis'' (Chinese prawn), ''

'' (giant tiger prawn) and ''

'' (whiteleg prawn).

References

Bibliography

* * * * * * * * * * *

External links

* * * * * {{Taxonbar, from=Q6500145 Decapods Commercial crustaceans Seafood Extant Late Devonian first appearances Taxa named by Charles Spence Bate Arthropod suborders