Dendrobranchiata

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Dendrobranchiata
Temporal range: Famennian–Recent
Shrimp body.jpg
Penaeus monodon
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Malacostraca
Order: Decapoda
Suborder: Dendrobranchiata
Bate, 1888
Superfamilies and families [1]

Penaeoidea

Aciculopodidae
Aegeridae
Aristeidae
Benthesicymidae
Carpopenaeidae
Penaeidae
Sicyoniidae
Solenoceridae

Sergestoidea

Luciferidae
Sergestidae
Synonyms   [2]

Penaeidea Dana, 1852

Dendrobranchiata is a suborder of decapods, commonly known as prawns. There are 540 extant species in seven families, and a fossil record extending back to the Devonian. They differ from related animals, such as Caridea and Stenopodidea, by the branching form of the gills 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 330 millimetres (13 in) and a mass of 450 grams (1.0 lb), and are widely fished and farmed for human consumption.

Contents

Shrimp and prawns

While Dendrobranchiata and Caridea belong to different suborders of Decapoda, they are very similar in appearance, and in many contexts such as commercial farming and fisheries, they are both often referred to as "shrimp" and "prawn" interchangeably. In the United Kingdom, the word "prawn" is more common on menus than "shrimp", while the opposite is the case in North America. 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 and some other Commonwealth nations follow this British usage to an even greater extent, using the word "prawn" almost exclusively.

Description

Together with other swimming Decapoda, Dendrobranchiata show the "caridoid facies", or shrimp-like form. [3] The body is typically robust, and can be divided into a cephalothorax (head and thorax fused together) and a pleon (abdomen). [3] The body is generally slightly flattened side-to-side. [3] The largest species, Penaeus monodon , can reach a mass of 450 grams (16 oz) and a length of 336 millimetres (13.2 in). [4]

Head Shrimp head.jpg
Head

The most conspicuous appendages arising from the head are the antennae. The first pair are biramous (having two flagella), except in Luciferidae, and are relatively small. [5] The second pair can be 2–3 times the length of the body and are always uniramous (having a single flagellum). [5] The mouthparts comprise pairs of mandibles, maxillules and maxillae, arising from the head, and three pairs of maxillipeds, arising from the thorax. [6] A pair of stalked eyes points forwards from the head. [7]

Thorax

The carapace grows from the thorax to cover the cephalothorax, and extends forwards between the eyes into a rostrum. [7] This is only as long as the stalked eyes in Benthesicymidae, Luciferidae and Sergestidae, but considerably longer in Aristeidae. [7]

As well as the three pairs of maxillipeds, the thorax also bears five pairs of pereiopods, or walking legs; the first three of these end in small chelae (pincers). [8] The last two pereiopods are absent in Luciferidae and Acetes , but much longer than the preceding pereiopods in Hymenopenaeus and Xiphopenaeus . [9]

The thoracic appendages carry gills, which are protected beneath the carapace. [10] The gills are typically branched, and so resemble trees, lending the group its scientific name, Dendrobranchiata, from the Greek words δένδρον (dendron, tree) and βράγχια ( branchia , gills). [11]

Pleon

The pleon, or abdomen, is similar in length to the cephalothorax. [12] It has six segments, the first five bearing lamellar pleopods, and the last one bearing uropods. [13] The pleopods are biramous, except in Sicyoniidae, where they are uniramous. [9] The uropods and telson collectively form the tail fan; the uropods are not divided by a diaeresis, as they are in many other decapods. [14] The telson is pointed and is usually armed with four pairs of setae or spines. [14]

Internal anatomy

Most of the musculature of a prawn is used for bending the pleon, and almost all the space in the pleon is filled by muscle. [15] 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. [16] These muscles, collectively, are the meat for which prawns are commercially fished and farmed. [17]

The nervous system of prawns comprises a dorsal brain, and a ventral nerve cord, connected by two commissures around the oesophagus. [18] The chief sensory inputs are visual input from the eyes, chemoreceptors on the antennae and in the mouth, and mechanoreceptors on the antennae and elsewhere. [19]

The digestive system comprises a foregut, a midgut and a hindgut, and is situated dorsally. [20] The foregut begins at the mouth, passes through the oesophagus, and opens into a sac that contains the grinding apparatus of the gastric mill. [20] The hepatopancreas feeds into the midgut, where digestive enzymes are released, and nutrients taken up. [20] The hindgut forms faecal pellets, which are then passed out through the muscular anus. [21]

The circulatory system is based around a compact, triangular heart, which pumps blood into three main arteries. [22] Excretion is carried out through the gills, and by specialised glands located at the base of the antennae, and is mostly in the form of ammonia. [23]

Life cycle

Nauplius of an unidentified prawn from the family Penaeidae Shrimp nauplius.jpg
Nauplius of an unidentified prawn from the family Penaeidae
A juvenile Litopenaeus vannamei (whiteleg prawn) Penaeus vannamei 01.jpg
A juvenile Litopenaeus vannamei (whiteleg prawn)

Prawns may be divided into two groups: those with an open thelycum (female genitalia) and those with a closed thelycum. [24] In the open–thelycum species, mating takes place towards the end of the moulting cycle, and usually at sunset. [25] In closed–thelycum species, mating takes place shortly after moulting, when the exoskeleton is still soft, and usually occurs in the night. [26] Courtship and mating may take up to 3 hours in Penaeus monodon , while in Farfantepenaeus paulensis , mating lasts just 4–5 seconds. [26] Spawning may occur several times during the moulting cycle, and usually occurs at night. [27]

With the exception of Luciferidae, the eggs of prawns are shed directly into the water, rather than being brooded. [28] The eggs hatch into nauplius larvae, which are followed by zoea larvae (initially protozoea, and later mysis) and then a postlarva, before reaching adulthood. [28] The changes between moults are gradual, and so the development is anamorphic rather than metamorphic. [28]

Uniquely among the Decapoda, the nauplii of Dendrobranchiata are free-swimming. [28] There are five to eight naupliar stages. [29] The earlier stages have three pairs of appendages that are used for locomotion – two pairs of antennae and the mandibles. Later stages also have rudiments of other mouthparts, but the nauplius is unable to feed, and only lasts 24 to 68 hours. [28] The body ends at a two-lobed telson, and the beginnings of a carapace emerge at this stage. [29]

There are typically 5 or 6 zoea stages in Dendrobranchiata, divided into protozoea and mysis. [28] In the protozoea larvae, the antennae are still used for locomotion, but the mandibles become specialised for mastication. [29] All the thoracic somites (body segments) have formed, and a carapace is present, covering part of the thorax. [29] It is smooth in the family Penaeidae, but bears many spines in the family Solenoceridae. [29] The pleon (abdomen) is unsegmented in the first protozoea, and ends in a bilobed telson, which may be used for cleaning other appendages, or for steering. [29] By the second protozoea, segmentation appears on the pleon, [29] and by the third protozoea, which may also be called the metazoea, the uropods have appeared. [30]

By the mysis stages, the pereiopods (thoracic appendages) start to be used instead of the antennae for locomotion. [30] The larva swims backwards, with its tail upwards, spinning slowly as it goes. [30] The carapace covers most of the segments of the thorax, and claws appear on the first three pereiopods. [30] By the last mysis stage, the beginnings of pleopods have appeared on the first five segments of the abdomen. [30]

The post-larva or juvenile stage is characterised by the use of the pleopods for locomotion. [31] The claws become functional, but the gills are still rudimentary. [31] The telson is narrower and only retains traces of its two-lobed development. [31] Through a series of gradual changes over following moults, the animal takes on its adult form. [31]

Systematics

Dendrobranchiata were traditionally grouped together with Caridea as "Natantia" (the swimming decapoda), as opposed to the Reptantia (the walking decapods). In 1888, Charles Spence Bate recognised the differences in gill morphology, and separated Natantia into Dendrobranchiata, Phyllobranchiata and Trichobranchiata. [32] Recent analyses using cladistics and molecular phylogenetics recognise Dendrobranchiata as the sister group to all other Decapoda, collectively called Pleocyemata. [33]

The cladogram below shows Dendrobranchiata's placement within the larger order Decapoda, from analysis by Wolfe et al., 2019. [34]

Decapoda

Dendrobranchiata (prawns) Litopenaeus setiferus.png

Pleocyemata

Stenopodidea (boxer shrimp) Spongicola venustus.png

Procarididea

Caridea (true shrimp) Macrobrachium sp.jpg

Reptantia

Achelata (spiny lobsters, slipper lobsters) Panulirus argus.png

Polychelida (benthic crustaceans)

Astacidea (lobsters, crayfish) Lobster NSRW rotated2.jpg

Axiidea (mud shrimp, ghost shrimp, or burrowing shrimp)

Gebiidea (mud lobsters and mud shrimp)

Anomura (hermit crabs and others) Coenobita variabilis.jpg

Brachyura (crabs) Charybdis japonica.jpg

 (crawling / walking decapods) 
Fossil Aeger (Aegeridae) 9150 - Milano - Museo storia naturale - Aeger - Foto Giovanni Dall'Orto 22-Apr-2007 cropped.jpg
Fossil Aeger (Aegeridae)

Before 2010, the earliest known fossil prawns come from rocks in Madagascar of Permo-Triassic age, 250  million years ago. [35] [36] In 2010, however, the discovery of Aciculopoda from Famennian–stage rocks in Oklahoma extended the group's fossil record back to 360  million years ago. [37] The best known fossil prawns are from the Jurassic Solnhofen limestones from Germany. [36]

Living prawns are divided among seven families, five in the superfamily Penaeoidea, and two in the Sergestoidea, [2] although molecular evidence disagrees with some aspects of the current classifications. [38] Collectively, these include 540 extant species, and nearly 100 exclusively fossil species. [1] A further two families are known only from fossils. [1]

The cladogram below shows Dendrobranchiata's internal relationships of extant families (excluding Solenoceridae): [34]

Dendrobranchiata
Sergestoidea

Luciferidae

Sergestidae

Penaeoidea

Sicyoniidae

Penaeidae

Benthesicymidae

Aristeidae


Dendrobranchiata comprises the following superfamilies and families:

Superfamily Penaeoidea Rafinesque-Schmaltz, 1815
Aciculopodidae Feldmann & Schweitzer, 2010 – a single Famennian species, Aciculopoda mapesi [37]
Aegeridae Burkenroad, 1963 – two Mesozoic genera: Aeger and Acanthochirana [39]
Aristeidae Wood-Mason, 1891 – 26 extant species in 9 genera, and one fossil genus [40]
Benthesicymidae Wood-Mason, 1891 – 41 species in 4 genera [40]
Carpopenaeidae Garassino, 1994 – two Cretaceous species of Carpopenaeus [41]
Penaeidae Rafinesque-Schmaltz, 1815 – 216 extant species in 26 genera, and several extinct genera, mostly Mesozoic [42]
Sicyoniidae Ortmann, 1898 – 43 species of Sicyonia [43]
Solenoceridae Wood-Mason, 1891 – 81 species in 9 genera [44]
Superfamily Sergestoidea Dana, 1852
Luciferidae De Haan, 1849 – 7 species in 2 genera
Sergestidae Dana, 1852 – 90 extant species in six genera, and two extinct monotypic genera [45]

Distribution

The biodiversity of Dendrobranchiata decreases markedly at increasing latitudes; most species are only found in a region between 40° north and 40° south. [46] Some species may occur at higher latitudes. For instance, Bentheogennema borealis is abundant at 57° north in the Pacific Ocean, while collections of Gennadas kempi have been made as far south as 61° south in the Antarctic Ocean. [46]

Ecology and behaviour

Fish, such as this Argyropelecus hemigymnus, are important predators of juvenile prawns, such as this Sicyonia. Messina Straits Argyropelecus hemigymnus.jpg
Fish, such as this Argyropelecus hemigymnus , are important predators of juvenile prawns, such as this Sicyonia .

There is a great deal of ecological variation within the suborder Dendrobranchiata. Some species of Sergestidae live in fresh water, but most prawns are exclusively marine. [31] Species of Sergestidae and Benthesicymidae mostly live in deep water, and Solenoceridae species live offshore, while most Penaeidae species live in shallow inshore waters, and Lucifer is planktonic. [31] Some species burrow in mud on the sea floor during the day and emerge at night to feed. [31]

Prawns are "opportunistic omnivores", [47] and their diet can include a range of food items from fine particles to large organisms. These may include fish, chaetognaths, krill, copepods, radiolarians, phytoplankton, nematocysts, ostracods and detritus. [47] Prawns eat less around the time of ecdysis (moulting), probably because of the softness of the mouthparts, and must eat more than usual to compensate, once ecdysis is complete. [47]

Prawns are an attractive food for predators, with a higher energy content than most other invertebrates. [48] The larvae are prey to comb jellies, jellyfish, chaetognaths, fish and other crustaceans (such as mantis shrimp and crabs), and only a tiny proportion survive. [49] Juveniles are targeted by a number of fish, cephalopods and birds; Litopenaeus vannamei juveniles experience 90% mortality in the 6–12 weeks they spend in Mexican lagoons, and this is thought to be due almost entirely to predation. [49] Adult prawns are less susceptible to predation, but can fall prey to some fish. [50]

Economic importance

Marsupenaeus japonicus (Kuruma prawn) in an aquaculture facility in Taiwan Marsupenaeus japonicus.jpg
Marsupenaeus japonicus (Kuruma prawn) in an aquaculture facility in Taiwan

Dendrobranchiata are of huge importance. While in some countries, such as the United States, production is almost entirely through fisheries, other countries have concentrated on aquaculture (shrimp farms), including Ecuador where 95% of production is farmed; some countries produce similar amounts from fisheries and aquaculture, including Mexico, China, India and Indonesia. [51]

Species from the family Aristeidae are important to deep-water fisheries, particularly in the Mediterranean Sea, where Aristaeomorpha foliacea is caught by trawlers. [51] In Brazil, Aristaeomorpha foliacea, Aristaeopsis edwardsiana and Aristeus antillensis are of commercial importance. [51] The shallow-water Penaeidae are of greater importance, however, and the most important species for fisheries is Fenneropenaeus chinensis , with a catch in 2005 of over 100,000 tons. [33]

The most important species for aquaculture are Marsupenaeus japonicus (Kuruma prawn), Fenneropenaeus chinensis (Chinese prawn), Penaeus monodon (giant tiger prawn) and Litopenaeus vannamei (whiteleg prawn). [33]

Related Research Articles

<span class="mw-page-title-main">Caridea</span> Infraorder of shrimp

The Caridea, commonly known as caridean shrimp or true shrimp, from the Greek word καρίς, καρίδος, 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 other animals with similar names – such as the mud shrimp of Axiidea and the boxer shrimp of Stenopodidea – are not true shrimp, but many have evolved features similar to true shrimp.

<span class="mw-page-title-main">Malacostraca</span> Largest class of crustaceans

Malacostraca is the second largest of the six classes of pancrustaceans just behind hexapods, containing about 40,000 living species, divided among 16 orders. Its members, the malacostracans, display a great diversity of body forms and include crabs, lobsters, crayfish, shrimp, krill, prawns, woodlice, amphipods, mantis shrimp, tongue-eating lice and many other less familiar animals. They are abundant in all marine environments and have colonised freshwater and terrestrial habitats. They are segmented animals, united by a common body plan comprising 20 body segments, and divided into a head, thorax, and abdomen.

<span class="mw-page-title-main">Decapoda</span> Order of crustaceans

The Decapoda or decapods are an order of crustaceans within the class Malacostraca, and includes crabs, lobsters, crayfish, shrimp, and prawns. Most decapods are scavengers. The order is estimated to contain nearly 15,000 extant species in around 2,700 genera, with around 3,300 fossil species. Nearly half of these species are crabs, with the shrimp and Anomura including hermit crabs, porcelain crabs, squat lobsters making up the bulk of the remainder. The earliest fossils of the group date to the Devonian.

<span class="mw-page-title-main">Isopoda</span> Order of arthropods

Isopoda is an order of crustacean, which includes woodlice and their relatives. Members of this group are called Isopods and include both terrestrial and aquatic species. All have rigid, segmented exoskeletons, two pairs of antennae, seven pairs of jointed limbs on the thorax, and five pairs of branching appendages on the abdomen that are used in respiration. Females brood their young in a pouch under their thorax.

<span class="mw-page-title-main">Reptantia</span> Suborder of crustaceans

Reptantia is a clade of decapod crustaceans named in 1880 which includes lobsters, crabs and many other well-known crustaceans.

<span class="mw-page-title-main">Pleocyemata</span> Suborder of crustaceans

Pleocyemata is a suborder of 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 Pleocyemata. Pleocyemata contains all the members of the Reptantia, as well as the Stenopodidea, and Caridea, which contains the true shrimp.

<span class="mw-page-title-main">Squat lobster</span> Decapod crustaceans in the infraorder Anomura

Squat lobsters are dorsoventrally flattened crustaceans with long tails held curled beneath the cephalothorax. They are found in the two superfamilies Galatheoidea and Chirostyloidea, which form part of the decapod infraorder Anomura, alongside groups including the hermit crabs and mole crabs. They are distributed worldwide in the oceans, and occur from near the surface to deep sea hydrothermal vents, with one species occupying caves above sea level. More than 900 species have been described, in around 60 genera. Some species form dense aggregations, either on the sea floor or in the water column, and a small number are commercially fished.

<span class="mw-page-title-main">Decapod anatomy</span> Entire structure of a decapod crustacean

The decapod 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 groups these may be reduced or missing. They are, from head to tail:

<i>Lucifer</i> (crustacean) Genus of crustaceans

Lucifer is a little-known and degenerate genus of prawns, the type genus of the family Luciferidae. Lucifer has a long body, but many fewer appendages than other prawns, with only three pairs of pereiopods remaining, all without claws. It also bears no gills. The females, uniquely among prawns, carry the fertilised eggs on her pleopods until they are ready to hatch. This parallels the development of a similar system in pleocyemates, although the attachment is less strong in Lucifer. The length of the eye-stalks and the form of the petasma are used in distinguishing the eight species from each other.

<span class="mw-page-title-main">Penaeidae</span> Family of crustaceans

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, Atlantic white shrimp, and Indian prawn. Many prawns are the subject of commercial fishery, and farming, both in marine settings, and in freshwater farms. Lateral line–like sense organs on the antennae have been reported in some species of Penaeidae. At 210 metres per second (760 km/h), the myelinated giant interneurons of pelagic penaeid shrimp have the world record for impulse conduction speed in any animal.

<span class="mw-page-title-main">Cumacea</span> Order of crustacean

Cumacea is an order of small marine crustaceans of the superorder Peracarida, occasionally called hooded shrimp or comma shrimp. Their unique appearance and uniform body plan makes them easy to distinguish from other crustaceans. They live in soft-bottoms such as mud and sand, mostly in the marine environment. There are more than 1,500 species of cumaceans formally described. The species diversity of Cumacea increases with depth.

<i>Polycheles typhlops</i> Species of crustacean

Polycheles typhlops is a species of blind, deep water decapod crustacean with a cosmopolitan distribution. It is "one of the dominant and most characteristic crustaceans in deep-sea communities of the Mediterranean Sea".

<span class="mw-page-title-main">Crustacean larva</span> Crustacean larval and immature stages between hatching and adult form

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 of crustaceans often bear little resemblance to the adult, and there are still cases where it is not known what larvae will grow into what adults. This is especially true of crustaceans which live as benthic adults, more-so than where the larvae are planktonic, and thereby easily caught.

<i>Marsupenaeus</i> Genus of crustaceans

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 also reached the Mediterranean Sea as a Lessepsian migrant. It is one of the largest species of prawns, and is accordingly one of the most economically important species in the family.

<i>Trachysalambria curvirostris</i> Species of crustacean

Trachysalambria curvirostris is a species of prawn that lives in shallow waters of the Indo-West Pacific. It is one of the most important species targeted by prawn fishery, with annual harvests of more than 300,000 t, mostly landed in China.

<span class="mw-page-title-main">Shrimp</span> Decapod crustaceans

A shrimp is a crustacean with an elongated body and a primarily swimming mode of locomotion – typically belonging to the Caridea or Dendrobranchiata of the decapod order, although some crustaceans outside of this order are also referred to as "shrimp".

<i>Albunea carabus</i> Species of crustacean

Albunea carabus is a rare species of "sand crab" or "mole crab" in the genus Albunea. It lives in shallow, turbulent waters in sandy areas of the tropical eastern Atlantic Ocean and the Mediterranean Sea.

<span class="mw-page-title-main">Crustacean</span> Subphylum of arthropods

Crustaceans are invertebrate animals of the subphylum Crustacea, a large, diverse group of mainly aquatic arthropods including decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, opossum shrimps, amphipods and mantis shrimp. The crustacean group can be treated as a subphylum under the clade Mandibulata. It is now well accepted that the hexapods emerged deep in the Crustacean group, with the completed group referred to as Pancrustacea. The three classes Cephalocarida, Branchiopoda and Remipedia are more closely related to the hexapods than they are to any of the other crustaceans.

<i>Ogyrides</i> Genus of crustaceans

Ogyrides, also known as long eyed shrimps, is a genus of decapod crustaceans consisting of 13 species. It is the only genus in the monotypic family Ogyrididae.

<i>Belzebub hanseni</i> Species of crustacean

Belzebub hanseni, the ghost shrimp or ghost prawn, is a small planktonic and benthic species of prawn from the family Luciferidae.

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