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Dendrobranchiata is a suborder of , commonly known as prawns (though this may be ambiguous). There are 540 extant species in seven families, and a extending back to the . They differ from related animals, such as and , by the branching form of the (hence their ) 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 and for human consumption.


Nomenclature
While Dendrobranchiata and belong to different suborders of , they are very similar in appearance, and in many contexts such as commercial and , they are both often referred to as "shrimp" and "prawn" interchangeably. In the , and some other Commonwealth nations, the word "prawn" is used almost exclusively, while the opposite is the case in . 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").

Due to their superficial similarity, Dendrobranchiata and Caridea were historically grouped together, though it is currently understood that they are distantly related. The classification of these groups is explained below.


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 (fusion of the head or cephalon and ) and a (abdomen). The body is generally slightly flattened side-to-side. The largest species, , can reach a of and a length of .Dall, 1990, pp. 3–4


Head
The most conspicuous arising from the head are the antennae; the first pair are (having two ), except in , 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 (having a single flagellum). The mouthparts comprise pairs of mandibles, maxillules and maxillae, arising from the head, and three pairs of , arising from the thorax.Tavares & Martin, 2010, pp. 106–108 A pair of stalked points forwards from the head.Tavares & Martin, 2010, p. 102


Thorax
The originates from the thorax to cover the cephalothorax, and extends forwards between the eyes into a rostrum. This is only as long as the stalked eyes in , and , but considerably longer in .

As well as the three pairs of maxillipeds, the thorax also bears five pairs of , or walking legs; the first three of these end in small chelae (pincers).Tavares & Martin, 2010, pp. 108–110 The last two pereiopods are absent in and , but much longer than the preceding pereiopods in and .Tavares & Martin, 2010, p. 110

The thoracic appendages carry , 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 words δένδρον]] ( dendron, ) and βράγχια ( branchia]], 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 , and the last one bearing .Tavares & Martin, 2010, pp. 110–111 The pleopods are biramous, except in , where they are uniramous. The uropods and 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 or spines.


Internal anatomy
Most of the 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.Tavares & Martin, 2010, pp. 113–114 These muscles, collectively, are the meat for which prawns are commercially fished and farmed.Kanduri & Eckhardt, 2002, p. 42

The of prawns comprises a dorsal , and a ventral nerve cord, connected by two around the .Tavares & Martin, 2010, p. 114 The chief sensory inputs are visual input from the eyes, on the antennae and in the mouth, and on the antennae and elsewhere.Tavares & Martin, 2010, pp. 116–118

The 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 . The feeds into the midgut, where are released, and nutrients taken up. The hindgut forms faecal pellets, which are then passed out through the muscular .Tavares & Martin, 2010, pp. 118–119

The circulatory system is based around a compact, triangular , which pumps blood into three main .Tavares & Martin, 2010, p. 120 is carried out through the gills, and by specialised located at the base of the antennae, and is mostly in the form of .Tavares & Martin, 2010, pp. 120–121


Life cycle
Prawns may be divided into two groups based on the structure of the female genitalia, known as thelycum: those with an open thelycum and those with a closed thelycum.Tavares & Martin, 2010, p. 125 In the open-thelycum species, mating takes place towards the end of the cycle, and usually at sunset.Tavares & Martin, 2010, pp. 125–126 In closed-thelycum species, mating takes place shortly after moulting, when the is still soft, and usually occurs in the night.Tavares & Martin, 2010, p. 126 Courtship and mating may take up to 3 hours in , while in Farfantepenaeus paulensis, 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 of prawns are shed directly into the water, rather than being brooded.Tavares & Martin, 2010, p. 130 The eggs hatch into nauplius larvae, which are followed by larvae (initially protozoea, and later mysis) and then a postlarva, before reaching adulthood. The changes between are gradual, and so the development is anamorphic rather than .

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 mandibles. 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 five or six 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 . All the thoracic somites (body segments) have formed, and a is present, covering part of the thorax. It is smooth in the family , but bears many spines in the family . 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 have appeared.

By the mysis stages, the (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 have appeared on the first five segments of the abdomen.

The post-larva or juvenile stage is characterised by the use of the pleopods for locomotion.Tavares & Martin, 2010, p. 134 The claws become functional, but the 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 Caridean shrimp as "" (the swimming decapoda), as opposed to the (the walking decapods). In 1888, Charles Spence Bate recognised the differences in gill morphology, and separated Natantia into Dendrobranchiata, Phyllobranchiata and Trichobranchiata.Tavares & Martin, 2010, p. 99 Recent analyses using and molecular phylogenetics recognise Dendrobranchiata as the to all other Decapoda, collectively called .Tavares & Martin, 2010, p. 137

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

Before 2010, the earliest known fossil prawns come from rocks in of - age, .Crean, 2004Schram et al., 2000 In 2010, however, the discovery of from –stage rocks in extended the group's fossil record back to .Feldmann & Schweitzer, 2010 The best known fossil prawns are from the Solnhofen limestones from .

Living prawns are divided among seven families, five in the superfamily , and two in the , although molecular evidence disagrees with some aspects of the current classifications.Ma et al., 2009 Collectively, these include 540 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 ):


Diversity

Dendrobranchiata comprises the following superfamilies and families:
Superfamily Rafinesque-Schmaltz, 1815
: Feldmann & Schweitzer, 2010 – a single species,
: Burkenroad, 1963 – two Mesozoic genera: and 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
: Garassino, 1994 – two species of 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 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 of Dendrobranchiata decreases markedly at increasing ; 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 , while collections of have been made as far south as 61° south in the .


Ecology and behaviour
There is a great deal of ecological variation within the suborder Dendrobranchiata. Some species of Sergestidae live in , 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 is . 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 can include a range of food items from fine particles to large organisms. These may include , , , , , , , and .Tavares & Martin, 2010, p. 135 Prawns eat less around the time of (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 , with a higher than most other invertebrates.Dall, 1990, p. 357 The larvae are prey to , , , and other (such as and ), and only a tiny proportion survive.Dall, 1990, p. 358 Juveniles are targeted by a number of fish, and ; Litopenaeus vannamei juveniles experience 90% mortality in the 6–12 weeks they spend in Mexican , 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 economic importance. While in some countries, such as the , production is almost entirely through , other countries have concentrated on (), including where 95% of production is ; some countries produce similar amounts from fisheries and aquaculture, including , , and .Tavares & Martin, 2010, p. 136

Species from the family are important to deep-water fisheries, particularly in the Mediterranean Sea, where Aristaeomorpha foliacea is caught by trawlers. In , 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, with a catch in 2005 of over 100,000 tons.

The most important species for are Marsupenaeus japonicus (Kuruma prawn), Fenneropenaeus chinensis (Chinese prawn), (giant tiger prawn) and Litopenaeus vannamei (whiteleg prawn).


Bibliography


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