Dendrobranchiata: Difference between revisions

Dendrobranchiata; Temporal range: Famennian–Recent PreꞒ Ꞓ O S D C P T J K Pg N
	Penaeus monodon
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Malacostraca
Order:	Decapoda
Suborder:	Dendrobranchiata; Bate, 1888
Superfamilies and families
	Penaeoidea Aciculopodidae † Aegeridae † Aristeidae Benthesicymidae Carpopenaeidae † Penaeidae Sicyoniidae Solenoceridae Sergestoidea Luciferidae Sergestidae
Synonyms
	Penaeidea Dana, 1852

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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.

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, Australia and some other Commonwealth, the word "prawn" is used almost exclusively, 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").

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

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

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 five or six 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)

Pleocyemata

Stenopodidea (boxer shrimp)

	Procarididea

	Caridea ("true" shrimp)

Reptantia

Achelata (spiny lobsters and slipper lobsters)

	Polychelida (benthic crustaceans)

	Astacidea (lobsters and crayfish)

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

Gebiidea (mud lobsters and mud shrimp)

	Anomura (hermit crabs and allies)

	Brachyura ("true" crabs)

(crawling /
walking
decapods)

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

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

Dendrobranchiata are of huge economic 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]

References

^ ^a ^b ^c De Grave et al., 2009
^ ^a ^b Martin & Davis, 2001
^ ^a ^b ^c Tavares & Martin, 2010, p. 100
^ Dall, 1990, pp. 3–4
^ ^a ^b Tavares & Martin, 2010, p. 106
^ Tavares & Martin, 2010, pp. 106–108
^ ^a ^b ^c Tavares & Martin, 2010, p. 102
^ Tavares & Martin, 2010, pp. 108–110
^ ^a ^b Tavares & Martin, 2010, p. 110
^ Tavares & Martin, 2010, pp. 103–105
^ Tavares & Martin, 2010, p. 103
^ Tavares & Martin, 2010, p. 105
^ Tavares & Martin, 2010, pp. 110–111
^ ^a ^b Tavares & Martin, 2010, p. 111
^ Tavares & Martin, 2010, p. 113
^ Tavares & Martin, 2010, pp. 113–114
^ Kanduri & Eckhardt, 2002, p. 42
^ Tavares & Martin, 2010, p. 114
^ Tavares & Martin, 2010, pp. 116–118
^ ^a ^b ^c Tavares & Martin, 2010, p. 118
^ Tavares & Martin, 2010, pp. 118–119
^ Tavares & Martin, 2010, p. 120
^ Tavares & Martin, 2010, pp. 120–121
^ Tavares & Martin, 2010, p. 125
^ Tavares & Martin, 2010, pp. 125–126
^ ^a ^b Tavares & Martin, 2010, p. 126
^ Tavares & Martin, 2010, p. 127
^ ^a ^b ^c ^d ^e ^f Tavares & Martin, 2010, p. 130
^ ^a ^b ^c ^d ^e ^f ^g Tavares & Martin, 2010, p. 131
^ ^a ^b ^c ^d ^e Tavares & Martin, 2010, p. 133
^ ^a ^b ^c ^d ^e ^f ^g Tavares & Martin, 2010, p. 134
^ Tavares & Martin, 2010, p. 99
^ ^a ^b ^c Tavares & Martin, 2010, p. 137
^ ^a ^b Wolfe, Joanna M.; Breinholt, Jesse W.; Crandall, Keith A.; Lemmon, Alan R.; Lemmon, Emily Moriarty; Timm, Laura E.; Siddall, Mark E.; Bracken-Grissom, Heather D. (24 April 2019). "A phylogenomic framework, evolutionary timeline and genomic resources for comparative studies of decapod crustaceans". Proceedings of the Royal Society B. 286 (1901). doi:10.1098/rspb.2019.0079. PMC 6501934. PMID 31014217.
^ Crean, 2004
^ ^a ^b Schram et al., 2000
^ ^a ^b Feldmann & Schweitzer, 2010
^ Ma et al., 2009
^ Tavares & Martin, 2010, p. 151
^ ^a ^b Tavares & Martin, 2010, p. 152
^ Tavares & Martin, 2010, pp. 152–153
^ Tavares & Martin, 2010, p. 153
^ Tavares & Martin, 2010, p. 154
^ Tavares & Martin, 2010, p. 155
^ Tavares & Martin, 2010, p. 156
^ ^a ^b Tavares & Martin, 2010, p. 145
^ ^a ^b ^c Tavares & Martin, 2010, p. 135
^ Dall, 1990, p. 357
^ ^a ^b Dall, 1990, p. 358
^ Dall, 1990, p. 359
^ ^a ^b ^c Tavares & Martin, 2010, p. 136

Bibliography

Bill Baker; Peggy Bendel. "Come and Say G'Day!". Travel Marketing Decisions (Summer 2005). Archived from the original (PDF) on November 4, 2007. Retrieved December 21, 2007.
Robert P. D. Crean (November 14, 2004). "Dendrobranchiata". Order Decapoda. University of Bristol. Archived from the original on February 29, 2012. Retrieved December 26, 2009.
William Dall (1990). The Biology of the Penaeidae. Advances in Marine Biology. Vol. 27. Academic Press. ISBN 978-0-12-026127-7.
Sammy De Grave; N. Dean Pentcheff; Shane T. Ahyong; et al. (2009). "A classification of living and fossil genera of decapod crustaceans" (PDF). Raffles Bulletin of Zoology. Suppl. 21: 1–109. Archived from the original (PDF) on 2011-06-06.
Rodney Feldmann; Carrie Schweitzer (2010). "The oldest shrimp (Devonian: Famennian) and remarkable preservation of soft tissue". Journal of Crustacean Biology. 30 (4): 629–635. doi:10.1651/09-3268.1.
Indian Aquaculture Authority (2001). "Shrimp Aquaculture and the Environment - An Environment Impact Assessment Report, chapter 2; IAA report" (PDF). Archived from the original (PDF) on 2011-07-16.
Laxman Kanduri; Ronald A. Eckhardt (2002). "HACCP in shrimp processing". Food Safety in Shrimp Processing: a Handbook for Shrimp Processors, Importers, Exporters and Retailers. John Wiley and Sons. pp. 40–64. ISBN 978-0-85238-270-7.
K. Y. Ma; T.-Y. Chan; K. H. Chu (2009). "Phylogeny of penaeoid shrimps (Decapoda: Penaeoidea) inferred from nuclear protein-coding genes". Molecular Phylogenetics and Evolution. 53 (1): 45–55. doi:10.1016/j.ympev.2009.05.019. PMID 19477284.
J. W. Martin; G. E. Davis (2001). An Updated Classification of the Recent Crustacea (PDF). Natural History Museum of Los Angeles County. pp. 1–132. Archived from the original (PDF) on 2013-05-12. Retrieved 2009-12-26.
Frederick R. Schram; Shen Yanbin; Ronald Vonk; Rodney S. Taylor (2000). "The first fossil stenopodidean" (PDF). Crustaceana. 73 (2): 235–242. CiteSeerX 10.1.1.611.5275. doi:10.1163/156854000504183. JSTOR 20106269.
Carolina Tavares; Joel W. Martin (2010). "Suborder Dendrobranchiata Bate, 1888" (PDF). In F. R. Schram; J. C. von Vaupel Klein; J. Forest; M. Charmantier-Daures (eds.). Eucarida: Euphausiacea, Amphionidacea, and Decapoda (partim). Treatise on Zoology – Anatomy, Taxonomy, Biology – The Crustacea. Vol. 9A. Brill Publishers. pp. 99–164. ISBN 978-90-04-16441-3.

External links

Michael Türkay (2011). "Dendrobranchiata". World Register of Marine Species.
J. K. Lowry. "Dendrobranchiata (Decapoda, Eucarida, Malacostraca)". Crustacea, the Higher Taxa. Australian Museum. Archived from the original on 2012-03-23. Retrieved 2011-06-11.
Arthropod Genomic Consortium. "Dendrobranchiata". ArthropodBase wiki. Archived from the original on 2013-04-14.
Media related to Dendrobranchiata at Wikimedia Commons
Data related to Dendrobranchiata at Wikispecies

[Grave-1] De Grave et al., 2009

[M&D-2] Martin & Davis, 2001

[TM100-3] Tavares & Martin, 2010, p. 100

[Dall3-4-4] Dall, 1990, pp. 3–4

[TM106-5] Tavares & Martin, 2010, p. 106

[TM106-8-6] Tavares & Martin, 2010, pp. 106–108

[TM102-7] Tavares & Martin, 2010, p. 102

[TM108-10-8] Tavares & Martin, 2010, pp. 108–110

[TM110-9] Tavares & Martin, 2010, p. 110

[TM103-5-10] Tavares & Martin, 2010, pp. 103–105

[TM103-11] Tavares & Martin, 2010, p. 103

[TM105-12] Tavares & Martin, 2010, p. 105

[TM110-11-13] Tavares & Martin, 2010, pp. 110–111

[TM111-14] Tavares & Martin, 2010, p. 111

[TM113-15] Tavares & Martin, 2010, p. 113

[TM113-114-16] Tavares & Martin, 2010, pp. 113–114

[17] Kanduri & Eckhardt, 2002, p. 42

[TM114-18] Tavares & Martin, 2010, p. 114

[TM116-8-19] Tavares & Martin, 2010, pp. 116–118

[TM118-20] Tavares & Martin, 2010, p. 118

[TM118-9-21] Tavares & Martin, 2010, pp. 118–119

[TM120-22] Tavares & Martin, 2010, p. 120

[TM120-1-23] Tavares & Martin, 2010, pp. 120–121

[TM125-24] Tavares & Martin, 2010, p. 125

[TM125-6-25] Tavares & Martin, 2010, pp. 125–126

[TM126-26] Tavares & Martin, 2010, p. 126

[TM127-27] Tavares & Martin, 2010, p. 127

[TM130-28] ^ ^a ^b ^c ^d ^e ^f Tavares & Martin, 2010, p. 130

[TM131-29] ^ ^a ^b ^c ^d ^e ^f ^g Tavares & Martin, 2010, p. 131

[TM133-30] Tavares & Martin, 2010, p. 133

[TM134-31] ^ ^a ^b ^c ^d ^e ^f ^g Tavares & Martin, 2010, p. 134

[TM99-32] Tavares & Martin, 2010, p. 99

[TM137-33] Tavares & Martin, 2010, p. 137

[Wolfe2019-34] Wolfe, Joanna M.; Breinholt, Jesse W.; Crandall, Keith A.; Lemmon, Alan R.; Lemmon, Emily Moriarty; Timm, Laura E.; Siddall, Mark E.; Bracken-Grissom, Heather D. (24 April 2019). "A phylogenomic framework, evolutionary timeline and genomic resources for comparative studies of decapod crustaceans". Proceedings of the Royal Society B. 286 (1901). doi:10.1098/rspb.2019.0079. PMC 6501934. PMID 31014217.

[35] Crean, 2004

[Schram-36] Schram et al., 2000

[Aciculopoda-37] Feldmann & Schweitzer, 2010

[38] Ma et al., 2009

[TM151-39] Tavares & Martin, 2010, p. 151

[TM152-40] Tavares & Martin, 2010, p. 152

[TM152-3-41] Tavares & Martin, 2010, pp. 152–153

[TM153-42] Tavares & Martin, 2010, p. 153

[TM154-43] Tavares & Martin, 2010, p. 154

[TM155-44] Tavares & Martin, 2010, p. 155

[TM156-45] Tavares & Martin, 2010, p. 156

[TM145-46] Tavares & Martin, 2010, p. 145

[TM135-47] Tavares & Martin, 2010, p. 135

[Dall357-48] Dall, 1990, p. 357

[Dall358-49] Dall, 1990, p. 358

[Dall359-50] Dall, 1990, p. 359

[TM136-51] Tavares & Martin, 2010, p. 136

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@@ Line 1: / Line 1: @@
+{{Short description|Suborder of prawns}}
 {{Automatic taxobox
 | fossil_range = {{fossil range|Famennian|Recent}}
-| image = Penaeus monodon.jpg
+| image = Shrimp body.jpg
 | image_caption = ''[[Penaeus monodon]]''
 | taxon = Dendrobranchiata
 | authority = [[Charles Spence Bate|Bate]], 1888
+| subdivision_ranks = Superfamilies and families
-| synonyms = Penaeidea <small>Dana, 1852</small>
-| synonyms_ref = &nbsp;<ref name="M&D">[[#MartinDavis2001|Martin & Davis, 2001]]</ref>
+| subdivision_ref = <ref name="Grave"/>
-| subdivision_ranks = Superfamilies and families&nbsp;<ref name="Grave"/>
 | subdivision =
 [[Penaeoidea]]
@@ Line 21: / Line 21: @@
 :[[Luciferidae]]
 :[[Sergestidae]]
+| synonyms = Penaeidea <small>Dana, 1852</small>
+| synonyms_ref = <ref name="M&D">[[#MartinDavis2001|Martin & Davis, 2001]]</ref>
 }}
-'''Dendrobranchiata''' is a suborder of [[Decapoda|decapod]] shrimps, commonly known as '''prawns'''.{{cn|date=March 2019}} 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 [[gill]]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 {{convert|330|mm}} and a mass of {{convert|450|g|lb|1}}, and are widely [[shrimp fishery|fished]] and [[shrimp farm|farmed]] for human consumption.
+'''Dendrobranchiata''' is a suborder of [[Decapoda|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 [[gill]]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 {{convert|330|mm}} and a mass of {{convert|450|g|lb|1}}, and are widely [[shrimp fishery|fished]] and [[shrimp farm|farmed]] for human consumption.
 ==Shrimp and prawns==
+While Dendrobranchiata and [[Caridea]] belong to different [[Order (biology)|suborder]]s of [[Decapoda]], they are very similar in appearance, and in many contexts such as commercial [[farming]] and [[Fishery|fisheries]], they are both often referred to as "shrimp" and "prawn" interchangeably. In the [[United Kingdom]], [[Australia]] and some other [[Commonwealth of Nations|Commonwealth]], the word "prawn" is used almost exclusively, 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").
-{{Refimprove section|date=March 2019}}
-While Dendrobranchiata and [[Caridea]] belong to different [[Order (biology)|suborder]]s of [[Decapoda]], they are very similar in appearance, and in many contexts such as commercial [[farming]] and [[Fishery|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 of Nations|Commonwealth]] nations follow this British usage to an even greater extent, using the word "prawn" almost exclusively. When Australian comedian [[Paul Hogan]] used the phrase, [[Shrimp on the barbie|"I'll slip an extra shrimp on the barbie for you"]] in an American television advertisement,<ref name="BakerBendel">[[#BakerBendel2007|Baker & Bendel, 2007]]</ref> 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==
@@ Line 33: / Line 34: @@
 ===Head===
+[[File:Shrimp head.jpg|right|thumb|Head]]
 The most conspicuous appendages arising from the head are the [[antenna (biology)|antennae]]. The first pair are biramous (having two [[flagellum|flagella]]), except in [[Luciferidae]], and are relatively small.<ref name="TM106">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 106</ref> The second pair can be 2–3 times the length of the body and are always uniramous (having a single flagellum).<ref name="TM106"/> The [[arthropod mouthparts|mouthparts]] comprise pairs of [[Mandible (arthropod mouthpart)|mandibles]], maxillules and maxillae, arising from the head, and three pairs of maxillipeds, arising from the thorax.<ref name="TM106-8">[[#TavaresMartin2010|Tavares & Martin, 2010]], pp. 106–108</ref> A pair of stalked [[eye]]s points forwards from the head.<ref name="TM102">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 102</ref>
@@ Line 38: / Line 40: @@
 The [[carapace]] grows from the thorax to cover the cephalothorax, and extends forwards between the eyes into a [[rostrum (anatomy)|rostrum]].<ref name="TM102"/> This is only as long as the stalked eyes in [[Benthesicymidae]], [[Luciferidae]] and [[Sergestidae]], but considerably longer in [[Aristeidae]].<ref name="TM102"/>
-As well as the three pairs of maxillipeds, the thorax also bears five pairs of [[pereiopod]]s, or walking legs; the first three of these end in small [[claw]]s.<ref name="TM108-10">[[#TavaresMartin2010|Tavares & Martin, 2010]], pp. 108–110</ref> The last two pereiopods are absent in [[Luciferidae]] and ''[[Acetes]]'', but much longer than the preceding pereiopods in ''[[Hymenopenaeus]]'' and ''[[Xiphopenaeus]]''.<ref name="TM110">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 110</ref>
+As well as the three pairs of maxillipeds, the thorax also bears five pairs of [[pereiopod]]s, or walking legs; the first three of these end in small [[Chela (organ)|chelae]] (pincers).<ref name="TM108-10">[[#TavaresMartin2010|Tavares & Martin, 2010]], pp. 108–110</ref> The last two pereiopods are absent in [[Luciferidae]] and ''[[Acetes]]'', but much longer than the preceding pereiopods in ''[[Hymenopenaeus]]'' and ''[[Xiphopenaeus]]''.<ref name="TM110">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 110</ref>
 The thoracic appendages carry [[gill]]s, which are protected beneath the carapace.<ref name="TM103-5">[[#TavaresMartin2010|Tavares & Martin, 2010]], pp. 103–105</ref> The gills are typically branched, and so resemble trees, lending the group its scientific name, Dendrobranchiata, from the [[Ancient Greek language|Greek]] words {{lang|el|[[:wikt:δένδρον|δένδρον]]}} (''{{lang|el|dendron}}'', [[tree]]) and {{lang|el|βράγχια}} (''{{lang|el|[[:wikt:branchia|branchia]]}}'', gills).<ref name="TM103">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 103</ref>
@@ Line 50: / Line 52: @@
 The [[nervous system]] of prawns comprises a dorsal [[brain]], and a [[ventral nerve cord]], connected by two [[commissure]]s around the [[oesophagus]].<ref name="TM114">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 114</ref> The chief sensory inputs are visual input from the eyes, [[chemoreceptor]]s on the antennae and in the mouth, and [[mechanoreceptor]]s on the antennae and elsewhere.<ref name="TM116-8">[[#TavaresMartin2010|Tavares & Martin, 2010]], pp. 116–118</ref>
-The [[digestive system]] comprises a foregut, a midgut and a hindgut, and is situated dorsally.<ref name="TM118">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 118</ref> The foregut begins at the mouth, passes through the oesophagus, and opens into a sac which contains the grinding apparatus of the [[gastric mill]].<ref name="TM118"/> The [[hepatopancreas]] feeds into the midgut, where [[digestive enzyme]]s are released, and nutrients taken up.<ref name="TM118"/> The hindgut forms faecal pellets, which are then passed out through the muscular [[anus]].<ref name="TM118-9">[[#TavaresMartin2010|Tavares & Martin, 2010]], pp. 118–119</ref>
+The [[digestive system]] comprises a foregut, a midgut and a hindgut, and is situated dorsally.<ref name="TM118">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 118</ref> The foregut begins at the mouth, passes through the oesophagus, and opens into a sac that contains the grinding apparatus of the [[gastric mill]].<ref name="TM118"/> The [[hepatopancreas]] feeds into the midgut, where [[digestive enzyme]]s are released, and nutrients taken up.<ref name="TM118"/> The hindgut forms faecal pellets, which are then passed out through the muscular [[anus]].<ref name="TM118-9">[[#TavaresMartin2010|Tavares & Martin, 2010]], pp. 118–119</ref>
 The [[circulatory system]] is based around a compact, triangular [[heart]], which pumps blood into three main [[artery|arteries]].<ref name="TM120">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 120</ref> [[Excretion]] is carried out through the gills, and by specialised [[gland]]s located at the base of the antennae, and is mostly in the form of [[ammonia]].<ref name="TM120-1">[[#TavaresMartin2010|Tavares & Martin, 2010]], pp. 120–121</ref>
@@ Line 57: / Line 59: @@
 [[File:Shrimp nauplius.jpg|thumb|[[Nauplius (larva)|Nauplius]] of an unidentified prawn from the family [[Penaeidae]]]]
 [[File:Penaeus vannamei 01.jpg|thumb|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.<ref name="TM125">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 125</ref> In the open–thelycum species, mating takes place towards the end of the moulting cycle, and usually at sunset.<ref name="TM125-6">[[#TavaresMartin2010|Tavares & Martin, 2010]], pp. 125–126</ref> In closed–thelycum species, mating takes place shortly after moulting, when the [[exoskeleton]] is still soft, and usually occurs in the night.<ref name="TM126">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 126</ref> [[Courtship]] and mating may take up to 3 hours in ''[[Penaeus monodon]]'', while in ''[[Farfantepenaeus paulensis]]'', mating lasts just 4–5 seconds.<ref name="TM126"/> Spawning may occur several times during the moulting cycle, and usually occurs at night.<ref name="TM127">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 127</ref>
+Prawns may be divided into two groups: those with an open thelycum (female genitalia) and those with a closed thelycum.<ref name="TM125">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 125</ref> In the open–thelycum species, mating takes place towards the end of the moulting cycle, and usually at sunset.<ref name="TM125-6">[[#TavaresMartin2010|Tavares & Martin, 2010]], pp. 125–126</ref> In closed–thelycum species, mating takes place shortly after moulting, when the [[exoskeleton]] is still soft, and usually occurs in the night.<ref name="TM126">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 126</ref> [[courtship display|Courtship]] and mating may take up to 3 hours in ''[[Penaeus monodon]]'', while in ''[[Farfantepenaeus paulensis]]'', mating lasts just 4–5 seconds.<ref name="TM126"/> Spawning may occur several times during the moulting cycle, and usually occurs at night.<ref name="TM127">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 127</ref>
 With the exception of [[Luciferidae]], the [[egg (biology)|eggs]] of prawns are shed directly into the water, rather than being brooded.<ref name="TM130">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 130</ref> The eggs hatch into [[nauplius (larva)|nauplius]] [[crustacean larvae|larvae]], which are followed by [[zoea]] larvae (initially protozoea, and later mysis) and then a postlarva, before reaching adulthood.<ref name="TM130"/> The changes between [[ecdysis|moults]] are gradual, and so the development is [[Anamorphosis (biology)|anamorphic]] rather than [[metamorphosis|metamorphic]].<ref name="TM130"/>
-Uniquely among the Decapoda, the nauplii of Dendrobranchiata are free-swimming.<ref name="TM130"/> There are five to eight naupliar stages.<ref name="TM131">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 131</ref> The earlier stages have three pairs of appendages which are used for locomotion – two pairs of [[antenna (biology)|antennae]] and the [[Mandible (arthropod mouthpart)|mandibles]]. Later stages also have rudiments of other [[arthropod mouthparts|mouthparts]], but the nauplius is unable to feed, and only lasts 24 to 68 hours.<ref name="TM130"/> The body ends at a two-lobed [[telson]], and the beginnings of a [[carapace]] emerge at this stage.<ref name="TM131"/>
+Uniquely among the Decapoda, the nauplii of Dendrobranchiata are free-swimming.<ref name="TM130"/> There are five to eight naupliar stages.<ref name="TM131">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 131</ref> The earlier stages have three pairs of appendages that are used for locomotion – two pairs of [[antenna (biology)|antennae]] and the [[Mandible (arthropod mouthpart)|mandibles]]. Later stages also have rudiments of other [[arthropod mouthparts|mouthparts]], but the nauplius is unable to feed, and only lasts 24 to 68 hours.<ref name="TM130"/> The body ends at a two-lobed [[telson]], and the beginnings of a [[carapace]] emerge at this stage.<ref name="TM131"/>
-There are typically 5 or 6 zoea stages in Dendrobranchiata, divided into protozoea and mysis.<ref name="TM130"/> In the protozoea larvae, the antennae are still used for locomotion, but the mandibles become specialised for [[mastication]].<ref name="TM131"/> All the thoracic somites (body segments) have formed, and a [[carapace]] is present, covering part of the thorax.<ref name="TM131"/> It is smooth in the family [[Penaeidae]], but bears many spines in the family [[Solenoceridae]].<ref name="TM131"/> 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.<ref name="TM131"/> By the second protozoea, segmentation appears on the pleon,<ref name="TM131"/> and by the third protozoea, which may also be called the metazoea, the [[uropod]]s have appeared.<ref name="TM133"/>
+There are typically five or six zoea stages in Dendrobranchiata, divided into protozoea and mysis.<ref name="TM130"/> In the protozoea larvae, the antennae are still used for locomotion, but the mandibles become specialised for [[mastication]].<ref name="TM131"/> All the thoracic somites (body segments) have formed, and a [[carapace]] is present, covering part of the thorax.<ref name="TM131"/> It is smooth in the family [[Penaeidae]], but bears many spines in the family [[Solenoceridae]].<ref name="TM131"/> 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.<ref name="TM131"/> By the second protozoea, segmentation appears on the pleon,<ref name="TM131"/> and by the third protozoea, which may also be called the metazoea, the [[uropod]]s have appeared.<ref name="TM133"/>
 By the mysis stages, the [[pereiopod]]s (thoracic appendages) start to be used instead of the antennae for locomotion.<ref name="TM133">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 133</ref> The larva swims backwards, with its tail upwards, spinning slowly as it goes.<ref name="TM133"/> The carapace covers most of the segments of the thorax, and claws appear on the first three pereiopods.<ref name="TM133"/> By the last mysis stage, the beginnings of [[pleopod]]s have appeared on the first five segments of the abdomen.<ref name="TM133"/>
@@ Line 71: / Line 73: @@
 ==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.<ref name="TM99">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 99</ref> Recent analyses using [[cladistics]] and [[molecular phylogenetics]] recognise Dendrobranchiata as the [[sister group]] to all other Decapoda, collectively called [[Pleocyemata]].<ref name="TM137">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 137</ref>
+The [[cladogram]] below shows Dendrobranchiata's placement within the larger [[order (biology)|order]] [[Decapoda]], from analysis by Wolfe ''et al.'', 2019.<ref name="Wolfe2019">{{cite journal |last1=Wolfe |first1=Joanna M. |last2=Breinholt |first2=Jesse W. |last3=Crandall |first3=Keith A. |last4=Lemmon |first4=Alan R. |last5=Lemmon |first5=Emily Moriarty |last6=Timm |first6=Laura E. |last7=Siddall |first7=Mark E. |last8=Bracken-Grissom |first8=Heather D. |url=https://royalsocietypublishing.org/doi/10.1098/rspb.2019.0079 |title=A phylogenomic framework, evolutionary timeline and genomic resources for comparative studies of decapod crustaceans |journal=Proceedings of the Royal Society B | date=24 April 2019 |volume=286 |issue=1901 |doi=10.1098/rspb.2019.0079 |pmid=31014217 |doi-access=free |pmc=6501934 }}</ref>
+{{Decapoda cladogram}}
 [[File:9150 - Milano - Museo storia naturale - Aeger - Foto Giovanni Dall'Orto 22-Apr-2007 cropped.jpg|thumb|Fossil ''[[Aeger]]'' ([[Aegeridae]])]]
 Before 2010, the earliest known fossil prawns come from rocks in [[Madagascar]] of [[Permian|Permo]]-[[Triassic]] age, {{Ma|250}}.<ref>[[#Crean2004|Crean, 2004]]</ref><ref name="Schram">[[#Schram2000|Schram ''et al.'', 2000]]</ref> In 2010, however, the discovery of ''[[Aciculopoda]]'' from [[Famennian]]–stage rocks in [[Oklahoma]] extended the group's fossil record back to {{Ma|360}}.<ref name="Aciculopoda">[[#FeldmannSchweitzer2010|Feldmann & Schweitzer, 2010]]</ref> The best known fossil prawns are from the [[Jurassic]] [[Solnhofen limestone]]s from [[Germany]].<ref name="Schram"/>
-Living prawns are divided among seven families, five in the superfamily [[Penaeoidea]], and two in the [[Sergestoidea]],<ref name="M&D"/> although molecular evidence disagrees with some aspects of the current classifications.<ref>[[#Ma2009|Ma ''et al.'', 2009]]</ref> Collectively, these include 540 extant species, and nearly 100 exclusively fossil species.<ref name="Grave">[[#DeGrave2009|De Grave ''et al.'', 2009]]</ref> A further two families are known only from fossils.<ref name="Grave"/>
+Living prawns are divided among seven families, five in the [[superfamily (taxonomy)|superfamily]] [[Penaeoidea]], and two in the [[Sergestoidea]],<ref name="M&D"/> although molecular evidence disagrees with some aspects of the current classifications.<ref>[[#Ma2009|Ma ''et al.'', 2009]]</ref> Collectively, these include 540 [[extant taxon|extant]] species, and nearly 100 exclusively fossil species.<ref name="Grave">[[#DeGrave2009|De Grave ''et al.'', 2009]]</ref> A further two families are known only from fossils.<ref name="Grave"/>
+The cladogram below shows Dendrobranchiata's internal relationships of [[extant taxon|extant]] families (excluding [[Solenoceridae]]):<ref name="Wolfe2019"/>
+{{clade
+|style=font-size:85%; line-height:85%
+|label1= '''Dendrobranchiata'''
+|1={{clade
+ |label1= [[Sergestoidea]]
+ |1={{clade
+  |1=[[Luciferidae]]
+  |2=[[Sergestidae]] }}
+ |label2= [[Penaeoidea]]
+ |2={{clade
+  |1={{clade
+   |1=[[Sicyoniidae]]
+   |2=[[Penaeidae]] }}
+  |2={{clade
+   |1=[[Benthesicymidae]]
+   |2=[[Aristeidae]] }} }} }} }}
+<br>
-Suborder [[Penaeoidea]] <small>Rafinesque-Schmaltz, 1815</small>
+Dendrobranchiata comprises the following [[superfamily (taxonomy)|superfamilies]] and [[family (biology)|families]]:<br>
-:† [[Aciculopoda|Aciculopodidae]] <small>Feldmann & Schweitzer, 2010</small> – a single [[Famennian]] species, ''[[Aciculopoda|Aciculopoda mapesi]]'' <ref name="Aciculopoda"/>
+:Superfamily [[Penaeoidea]] <small>Rafinesque-Schmaltz, 1815</small>
-:† [[Aegeridae]] <small>Burkenroad, 1963</small> – two Mesozoic genera: ''[[Aeger]]'' and ''[[Acanthochirana]]'' <ref name="TM151">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 151</ref>
-:[[Aristeidae]] <small>Wood-Mason, 1891</small> – 26 extant species in 9 genera, and one fossil genus <ref name="TM152">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 152</ref>
+::† [[Aciculopoda|Aciculopodidae]] <small>Feldmann & Schweitzer, 2010</small> – a single [[Famennian]] species, ''[[Aciculopoda|Aciculopoda mapesi]]'' <ref name="Aciculopoda"/>
-:[[Benthesicymidae]] <small>Wood-Mason, 1891</small> – 41 species in 4 genera <ref name="TM152"/>
+::† [[Aegeridae]] <small>Burkenroad, 1963</small> – two Mesozoic genera: ''[[Aeger]]'' and ''[[Acanthochirana]]'' <ref name="TM151">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 151</ref>
-:† [[Carpopenaeus|Carpopenaeidae]] <small>Garassino, 1994</small> – two [[Cretaceous]] species of ''[[Carpopenaeus]]'' <ref name="TM152-3">[[#TavaresMartin2010|Tavares & Martin, 2010]], pp. 152–153</ref>
+::[[Aristeidae]] <small>Wood-Mason, 1891</small> – 26 extant species in 9 genera, and one fossil genus <ref name="TM152">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 152</ref>
-:[[Penaeidae]] <small>Rafinesque-Schmaltz, 1815</small> – 216 extant species in 26 genera, and several extinct genera, mostly Mesozoic <ref name="TM153">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 153</ref>
+::[[Benthesicymidae]] <small>Wood-Mason, 1891</small> – 41 species in 4 genera <ref name="TM152"/>
-:[[Sicyoniidae]] <small>Ortmann, 1898</small> – 43 species of ''[[Sicyonia]]'' <ref name="TM154">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 154</ref>
+::† [[Carpopenaeus|Carpopenaeidae]] <small>Garassino, 1994</small> – two [[Cretaceous]] species of ''[[Carpopenaeus]]'' <ref name="TM152-3">[[#TavaresMartin2010|Tavares & Martin, 2010]], pp. 152–153</ref>
-:[[Solenoceridae]] <small>Wood-Mason, 1891</small> – 81 species in 9 genera <ref name="TM155">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 155</ref>
+::[[Penaeidae]] <small>Rafinesque-Schmaltz, 1815</small> – 216 extant species in 26 genera, and several extinct genera, mostly Mesozoic <ref name="TM153">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 153</ref>
+::[[Sicyoniidae]] <small>Ortmann, 1898</small> – 43 species of ''[[Sicyonia]]'' <ref name="TM154">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 154</ref>
-Suborder [[Sergestoidea]] <small>Dana, 1852</small>
-:[[Luciferidae]] <small>De Haan, 1849</small> – 7 species in 2 genera
+::[[Solenoceridae]] <small>Wood-Mason, 1891</small> – 81 species in 9 genera <ref name="TM155">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 155</ref>
+:Superfamily [[Sergestoidea]] <small>Dana, 1852</small>
-:[[Sergestidae]] <small>Dana, 1852</small> – 90 extant species in six genera, and two extinct monotypic genera <ref name="TM156">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 156</ref>
+::[[Luciferidae]] <small>De Haan, 1849</small> – 7 species in 2 genera
+::[[Sergestidae]] <small>Dana, 1852</small> – 90 extant species in six genera, and two extinct monotypic genera <ref name="TM156">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 156</ref>
 ==Distribution==
@@ Line 95: / Line 122: @@
 ==Ecology and behaviour==
 [[File:Messina Straits Argyropelecus hemigymnus.jpg|thumb|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.<ref name="TM134"/> 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 (prawn)|Lucifer]]'' is [[plankton]]ic.<ref name="TM134"/> Some species burrow in mud on the sea floor during the day and emerge at night to feed.<ref name="TM134"/>
+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.<ref name="TM134"/> 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 (crustacean)|Lucifer]]'' is [[plankton]]ic.<ref name="TM134"/> Some species burrow in mud on the sea floor during the day and emerge at night to feed.<ref name="TM134"/>
 Prawns are "opportunistic omnivores",<ref name="TM135"/> and their [[diet (nutrition)|diet]] can include a range of food items from fine particles to large organisms. These may include [[fish]], [[chaetognath]]s, [[krill]], [[copepod]]s, [[radiolaria]]ns, [[phytoplankton]], [[nematocyst]]s, [[ostracod]]s and [[detritus]].<ref name="TM135">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 135</ref> 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.<ref name="TM135"/>
@@ Line 102: / Line 129: @@
 ==Economic importance==
-[[File:Penaeus japonicus.jpg|thumb|''[[Marsupenaeus japonicus]]'' (Kuruma prawn) in an [[aquaculture]] facility in [[Taiwan]]]]
+[[File:Marsupenaeus japonicus.jpg|thumb|''[[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 [[shrimp fishery|fisheries]], other countries have concentrated on [[aquaculture]] ([[shrimp farm]]s), including [[Ecuador]] where 95% of production is [[shrimp farm|farmed]]; some countries produce similar amounts from fisheries and aquaculture, including [[Mexico]], [[China]], [[India]] and [[Indonesia]].<ref name="TM136">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 136</ref>
+Dendrobranchiata are of huge economic importance. While in some countries, such as the [[United States]], production is almost entirely through [[shrimp fishery|fisheries]], other countries have concentrated on [[aquaculture]] ([[shrimp farm]]s), including [[Ecuador]] where 95% of production is [[shrimp farm|farmed]]; some countries produce similar amounts from fisheries and aquaculture, including [[Mexico]], [[China]], [[India]] and [[Indonesia]].<ref name="TM136">[[#TavaresMartin2010|Tavares & Martin, 2010]], p. 136</ref>
 Species from the family [[Aristeidae]] are important to deep-water fisheries, particularly in the [[Mediterranean Sea]], where ''[[Aristaeomorpha foliacea]]'' is caught by trawlers.<ref name="TM136"/> In [[Brazil]], ''Aristaeomorpha foliacea'', ''[[Aristaeopsis edwardsiana]]'' and ''[[Aristeus antillensis]]'' are of commercial importance.<ref name="TM136"/> 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.<ref name="TM137" />
@@ Line 114: / Line 141: @@
 ===Bibliography===
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-*{{cite journal |author1=Bill Baker |author2=Peggy Bendel |issue=Summer 2005 |url=http://www.atme.org/pubs/archives/77_1898_11926.cfm |title=Come and Say G'Day! |format=[[Portable Document Format|PDF]] |journal=Travel Marketing Decisions |accessdate=December 21, 2007 |ref=BakerBendel2007 |deadurl=yes |archiveurl=https://web.archive.org/web/20071104153701/http://www.atme.org/pubs/archives/77_1898_11926.cfm |archivedate=November 4, 2007 |df= }}
+*{{cite journal |author1=Bill Baker |author2=Peggy Bendel |issue=Summer 2005 |url=http://www.atme.org/pubs/archives/77_1898_11926.cfm |title=Come and Say G'Day! |format=[[Portable Document Format|PDF]] |journal=Travel Marketing Decisions |access-date=December 21, 2007 |ref=BakerBendel2007 |url-status=dead |archive-url=https://web.archive.org/web/20071104153701/http://www.atme.org/pubs/archives/77_1898_11926.cfm |archive-date=November 4, 2007 }}
-*{{cite web |url=http://palaeo.gly.bris.ac.uk/Palaeofiles/Fossilgroups/Decapoda/dendrobranchiata.html |title=Dendrobranchiata |publisher=[[University of Bristol]] |work=Order Decapoda |author=Robert P. D. Crean |date=November 14, 2004 |ref=Crean2004 |access-date=December 26, 2009 |archive-url=https://web.archive.org/web/20120229170005/http://palaeo.gly.bris.ac.uk/Palaeofiles/Fossilgroups/Decapoda/dendrobranchiata.html |archive-date=February 29, 2012 |dead-url=yes }}
+*{{cite web |url=http://palaeo.gly.bris.ac.uk/Palaeofiles/Fossilgroups/Decapoda/dendrobranchiata.html |title=Dendrobranchiata |publisher=[[University of Bristol]] |work=Order Decapoda |author=Robert P. D. Crean |date=November 14, 2004 |ref=Crean2004 |access-date=December 26, 2009 |archive-url=https://web.archive.org/web/20120229170005/http://palaeo.gly.bris.ac.uk/Palaeofiles/Fossilgroups/Decapoda/dendrobranchiata.html |archive-date=February 29, 2012 |url-status=dead }}
-*{{cite book |author=William Dall |year=1990 |title=The Biology of the Penaeidae |publisher=[[Academic Press]] |series=Advances in Marine Biology |volume=27 |isbn=978-0-12-026127-7 |url=https://books.google.com/books?id=BcXoGjvrPC4C&pg=PA357 |ref=Dall1990}}
+*{{cite book |author=William Dall |year=1990 |title=The Biology of the Penaeidae |publisher=[[Academic Press]] |series=Advances in Marine Biology |volume=27 |isbn=978-0-12-026127-7 |url=https://books.google.com/books?id=BcXoGjvrPC4C&pg=PA357 |ref=Dall1990 }}
-*{{cite journal|journal=[[Raffles Bulletin of Zoology]] |year=2009 |volume=Suppl. 21 |pages=1–109 |title=A classification of living and fossil genera of decapod crustaceans |author1=Sammy De Grave |author2=N. Dean Pentcheff |author3=Shane T. Ahyong |url=http://rmbr.nus.edu.sg/rbz/biblio/s21/s21rbz1-109.pdf |ref=DeGrave2009 |display-authors=etal |deadurl=yes |archiveurl=https://web.archive.org/web/20110606064728/http://rmbr.nus.edu.sg/rbz/biblio/s21/s21rbz1-109.pdf |archivedate=2011-06-06 |df= }}
+*{{cite journal |journal=[[Raffles Bulletin of Zoology]] |year=2009 |volume=Suppl. 21 |pages=1–109 |title=A classification of living and fossil genera of decapod crustaceans |author1=Sammy De Grave |author2=N. Dean Pentcheff |author3=Shane T. Ahyong |url=http://rmbr.nus.edu.sg/rbz/biblio/s21/s21rbz1-109.pdf |ref=DeGrave2009 |display-authors=etal |url-status=dead |archive-url=https://web.archive.org/web/20110606064728/http://rmbr.nus.edu.sg/rbz/biblio/s21/s21rbz1-109.pdf |archive-date=2011-06-06 }}
-*{{cite journal |author1=Rodney Feldmann |author2=Carrie Schweitzer |year=2010 |title=The oldest shrimp (Devonian: Famennian) and remarkable preservation of soft tissue |journal=[[Journal of Crustacean Biology]] |volume=30 |issue=4 |pages=629–635 |doi=10.1651/09-3268.1 |ref=FeldmannSchweitzer2010}}
+*{{cite journal |author1=Rodney Feldmann |author2=Carrie Schweitzer |year=2010 |title=The oldest shrimp (Devonian: Famennian) and remarkable preservation of soft tissue |journal=[[Journal of Crustacean Biology]] |volume=30 |issue=4 |pages=629–635 |doi=10.1651/09-3268.1 |ref=FeldmannSchweitzer2010|doi-access=free }}
-*{{cite web |author=Indian Aquaculture Authority |url=http://aquaculture.tn.nic.in/pdf/farming.pdf |title=Shrimp Aquaculture and the Environment - An Environment Impact Assessment Report, chapter 2; IAA report |year=2001 |ref=IAA2001 |deadurl=yes |archiveurl=https://web.archive.org/web/20110716081932/http://aquaculture.tn.nic.in/pdf/farming.pdf |archivedate=2011-07-16 |df= }}
+*{{cite web |author=Indian Aquaculture Authority |url=http://aquaculture.tn.nic.in/pdf/farming.pdf |title=Shrimp Aquaculture and the Environment - An Environment Impact Assessment Report, chapter 2; IAA report |year=2001 |ref=IAA2001 |url-status=dead |archive-url=https://web.archive.org/web/20110716081932/http://aquaculture.tn.nic.in/pdf/farming.pdf |archive-date=2011-07-16 }}
-*{{cite book |author1=Laxman Kanduri |author2=Ronald A. Eckhardt |year=2002 |title=Food Safety in Shrimp Processing: a Handbook for Shrimp Processors, Importers, Exporters and Retailers |publisher=[[John Wiley and Sons]] |isbn=978-0-85238-270-7 |chapter=HACCP in shrimp processing |pages=40–64 |chapter-url=https://books.google.com/books?id=UCeZbFp6YM0C&pg=PA42 |ref=KanduriEckhardt2002}}
+*{{cite book |author1=Laxman Kanduri |author2=Ronald A. Eckhardt |year=2002 |title=Food Safety in Shrimp Processing: a Handbook for Shrimp Processors, Importers, Exporters and Retailers |publisher=[[John Wiley and Sons]] |isbn=978-0-85238-270-7 |chapter=HACCP in shrimp processing |pages=40–64 |chapter-url=https://books.google.com/books?id=UCeZbFp6YM0C&pg=PA42 |ref=KanduriEckhardt2002 }}
 *{{cite journal |doi=10.1016/j.ympev.2009.05.019 |title=Phylogeny of penaeoid shrimps (Decapoda: Penaeoidea) inferred from nuclear protein-coding genes |author1=K. Y. Ma |author2=T.-Y. Chan |author3=K. H. Chu |journal=[[Molecular Phylogenetics and Evolution]] |volume=53 |issue=1 |year=2009 |pages=45–55 |pmid=19477284 |ref=Ma2009}}
-*{{cite book |url=http://atiniui.nhm.org/pdfs/3839/3839.pdf |title=An Updated Classification of the Recent Crustacea |author1=J. W. Martin |author2=G. E. Davis |year=2001 |pages=1–132 |publisher=[[Natural History Museum of Los Angeles County]] |ref=MartinDavis2001}}
+*{{cite book |url=http://atiniui.nhm.org/pdfs/3839/3839.pdf |title=An Updated Classification of the Recent Crustacea |author1=J. W. Martin |author2=G. E. Davis |year=2001 |pages=1–132 |publisher=[[Natural History Museum of Los Angeles County]] |ref=MartinDavis2001 |access-date=2009-12-26 |archive-date=2013-05-12 |archive-url=https://web.archive.org/web/20130512091254/http://atiniui.nhm.org/pdfs/3839/3839.pdf |url-status=dead }}
-*{{cite journal |title=The first fossil stenopodidean |author1=Frederick R. Schram |author2=Shen Yanbin |author3=Ronald Vonk |author4=Rodney S. Taylor |journal=[[Crustaceana]] |volume=73 |issue=2 |year=2000 |pages=235–242 |doi=10.1163/156854000504183 |ref=Schram2000 |jstor=20106269|url=http://decapoda.nhm.org/pdfs/11778/11778.pdf |citeseerx=10.1.1.611.5275 }}
+*{{cite journal |title=The first fossil stenopodidean |author1=Frederick R. Schram |author2=Shen Yanbin |author3=Ronald Vonk |author4=Rodney S. Taylor |journal=[[Crustaceana]] |volume=73 |issue=2 |year=2000 |pages=235–242 |doi=10.1163/156854000504183 |ref=Schram2000 |jstor=20106269 |url=http://decapoda.nhm.org/pdfs/11778/11778.pdf |citeseerx=10.1.1.611.5275 }}
-*{{cite book |author1=Carolina Tavares |author2=Joel W. Martin |year=2010 |chapter=Suborder Dendrobranchiata Bate, 1888 |editor=F. R. Schram |editor2=J. C. von Vaupel Klein |editor3=J. Forest |editor4=M. Charmantier-Daures |series=Treatise on Zoology – Anatomy, Taxonomy, Biology – The Crustacea |isbn=978-90-04-16441-3 |publisher=[[Brill Publishers]] |title=Eucarida: Euphausiacea, Amphionidacea, and Decapoda (partim) |volume=9A |pages=99–164 |chapter-url=http://decapoda.nhm.org/pdfs/31616/31616.pdf |ref=TavaresMartin2010}}
+*{{cite book |author1=Carolina Tavares |author2=Joel W. Martin |year=2010 |chapter=Suborder Dendrobranchiata Bate, 1888 |editor=F. R. Schram |editor2=J. C. von Vaupel Klein |editor3=J. Forest |editor4=M. Charmantier-Daures |series=Treatise on Zoology – Anatomy, Taxonomy, Biology – The Crustacea |isbn=978-90-04-16441-3 |publisher=[[Brill Publishers]] |title=Eucarida: Euphausiacea, Amphionidacea, and Decapoda (partim) |volume=9A |pages=99–164 |chapter-url=http://decapoda.nhm.org/pdfs/31616/31616.pdf |ref=TavaresMartin2010 }}
 {{Refend}}
@@ Line 130: / Line 157: @@
 {{Portal|Crustaceans|Arthropods}}
 *{{cite web |author=Michael Türkay |year=2011 |title=Dendrobranchiata |publisher=[[World Register of Marine Species]] |url=http://www.marinespecies.org/aphia.php?p=taxdetails&id=106669}}
-*{{cite web |url=http://www.crustacea.net/crustace/www/dendrobr.htm |title=Dendrobranchiata (Decapoda, Eucarida, Malacostraca) |work=Crustacea, the Higher Taxa |author=J. K. Lowry |publisher=[[Australian Museum]] |access-date=2011-06-11 |archive-url=https://web.archive.org/web/20120323014244/http://www.crustacea.net/crustace/www/dendrobr.htm |archive-date=2012-03-23 |dead-url=yes }}
+*{{cite web |url=http://www.crustacea.net/crustace/www/dendrobr.htm |title=Dendrobranchiata (Decapoda, Eucarida, Malacostraca) |work=Crustacea, the Higher Taxa |author=J. K. Lowry |publisher=[[Australian Museum]] |access-date=2011-06-11 |archive-url=https://web.archive.org/web/20120323014244/http://www.crustacea.net/crustace/www/dendrobr.htm |archive-date=2012-03-23 |url-status=dead }}
-*{{cite web |url=http://arthropodgenomes.org/wiki/Dendrobranchiata |title=Dendrobranchiata |work=ArthropodBase wiki |author=Arthropod Genomic Consortium }}{{Dead link|date=July 2019 |bot=InternetArchiveBot |fix-attempted=yes }}
+*{{cite web |url=http://arthropodgenomes.org/wiki/Dendrobranchiata |archive-url=https://archive.today/20130414060140/http://arthropodgenomes.org/wiki/Dendrobranchiata |url-status=dead |archive-date=2013-04-14 |title=Dendrobranchiata |work=ArthropodBase wiki |author=Arthropod Genomic Consortium }}
 *{{Commons category-inline|Dendrobranchiata}}
 *{{Wikispecies-inline|Dendrobranchiata}}
+{{Decapoda}}
 {{commercial fish topics}}
 {{Edible crustaceans}}
@@ Line 140: / Line 168: @@
 [[Category:Dendrobranchiata| ]]
-[[Category:Decapods]]
 [[Category:Commercial crustaceans]]
 [[Category:Seafood]]
 [[Category:Extant Late Devonian first appearances]]
 [[Category:Taxa named by Charles Spence Bate]]
+[[Category:Arthropod suborders]]

v t e Edible crustaceans
Shrimp/ prawns	Acetes Crangon crangon Cryphiops caementarius Dried shrimp Indian prawn Litopenaeus setiferus Macrobrachium rosenbergii Palaemon serratus Pandalus borealis Penaeus esculentus Penaeus monodon Processa edulis Shrimp paste Whiteleg shrimp Xiphopenaeus kroyeri
Lobsters (incl. slipper & spiny)	American lobster Arctides guineensis California spiny lobster Homarus gammarus Ibacus peronii Japanese spiny lobster Jasus Jasus edwardsii Jasus lalandii Metanephrops challengeri Thenus orientalis Nephrops norvegicus Palinurus elephas Panulirus argus Panulirus cygnus Panulirus echinatus Panulirus guttatus Panulirus homarus Panulirus longipes Panulirus ornatus Panulirus pascuensis Panulirus penicillatus Panulirus versicolor Parribacus japonicus Sagmariasus Scyllarides herklotsii Scyllarides latus Scyllarus arctus Thymops birsteini Tristan rock lobster
Crabs	Callinectes sapidus Callinectes similis Cancer irroratus Cancer bellianus Cancer pagurus Cancer productus Chaceon fenneri Chaceon quinquedens Chinese mitten crab Chionoecetes Dungeness crab Florida stone crab Gecarcinus ruricola Horsehair crab Hypothalassia acerba Jonah crab Maja squinado Menippe adina Orithyia sinica Ovalipes australiensis Ovalipes catharus Pie crust crab Portunus pelagicus Portunus trituberculatus Ranina ranina Scylla paramamosain Scylla serrata Soft-shell crab Tasmanian giant crab
Crayfish	Acocil Astacus astacus Marron Cherax Paranephrops Procambarus clarkii Orconectes virilis Signal crayfish
Others	Austromegabalanus psittacus Coconut crab Goose barnacle King crab Krill Langostino Mantis shrimp Squat lobster Thalassina
Category