Caridea

The Caridea, commonly known as caridean shrimp, are an infraorder of shrimp within the order Decapoda. They are found widely around the world in both fresh and salt water.

Biology

Carideans are found in every kind of aquatic habitat, with the majority of species being marine. Around a quarter of the described species are found in fresh water, however, including almost all the members of the species-rich family Atyidae and the Palaemonidae subfamily Palaemoninae.[1] They include several commercially important species, such as Macrobrachium rosenbergii, and are found on every continent except Antarctica.[1] The marine species are found at depths to 5,000 m (16,000 ft),[2] and from the tropics to the polar regions.

In addition to the great variety in habitat, carideans vary greatly in form, from species a few millimetres long when fully grown,[3] to those that grow to over a foot long.[2] Except where secondarily lost, shrimp have one pair of stalked eyes, although they are sometimes covered by the carapace, which protects the cephalothorax.[2] The carapace also surrounds the gills, through which water is pumped by the action of the mouthparts.[2]

Most carideans are omnivorous, but some are specialised for particular modes of feeding. Some are filter feeders, using their setose (bristly) legs as a sieve; some scrape algae from rocks. The snapping shrimp of the genus Alpheus snap their claws to create a shock wave that stuns prey. Many cleaner shrimp, which groom reef fish and feed on their parasites and necrotic tissue, are carideans.[2] In turn, carideans are eaten by various animals, particularly fish and seabirds, and frequently host bopyrid parasites.[2]

Lifecycle

In most species of caridean shrimp, the females lay 50,000 to 1 million eggs, which hatch after some 24 hours into tiny nauplii. These nauplii feed on yolk reserves within their bodies and then undergo a metamorphosis into zoeae. This second larval stage feeds in the wild on algae and after a few days metamorphoses again into the third stage to become myses. At this stage, the myses already begin to appear like tiny versions of fully developed adults and feed on algae and zooplankton. After another three to four days, they metamorphose a final time into postlarvae: young shrimp having all the characteristics of adults. The whole process takes about 12 days from hatching. In the wild, the marine postlarvae then migrate into estuaries, which are rich in nutrients and low in salinity. There they grow and eventually migrate back into open waters when they mature. Most adult carideans are benthic animals living primarily on the sea floor.

Common species include Pandalus borealis (the "pink shrimp"), Crangon crangon (the "brown shrimp") and the snapping shrimp of the genus Alpheus. Depending on the species and location, they grow from about 1.2 to 30 cm (0.47 to 11.81 in) long, and live between 1.0 and 6.5 years.[4]

Commercial fishing

See also: Shrimp fishery
Global wild capture, 1950–2010, in tonnes, of caridean shrimp[5]

The most significant commercial species among the carideans is Pandalus borealis,[6] followed by Crangon crangon.[7] The wild capture production of P. borealis is about ten times that of C. crangon. In 1950, the position was reversed, with the capture of C. crangon about ten times that of P. borealis.[5]

In 2010, the global aquaculture of all shrimp and prawn species (3.5 million tonnes) slightly exceeded the global wild capture (3.2 million tonnes).[5] No carideans were significantly involved in aquaculture, but about 430,000 tonnes were captured in the wild. That is, about 13% of the global wild capture, or about 6% of the total production of all shrimp and prawns, were carideans.[5]

Taxonomy

The infraorder Caridea is divided into 15 superfamilies:[8]

Superfamily Image Description
Alpheoidea
Lysmata debelius		 Contains four families, including Alpheidae, the family of pistol or snapping shrimp, and Hippolytidae, a family of cleaner shrimp.[9][10]
Atyoidea
Atya gabonensis		 Contains one family, Atyidae, with 42 genera.[8] They are present in all tropical and most temperate waters. Adults of this family are almost always confined to fresh water.
Bresilioidea
Rimicaris kairei		 Likely to be an artificial group, containing five families[8] which may or may not be related.[11]
Campylonotoidea Contains two families. Chace considered it to be a sister group to the much larger superfamily Palaemonoidea (below) with which it shares the absence of endopods on the pereiopods, and a first pereiopod that is thinner than the second.[12] Using molecular phylogenetics, Bracken et al. proposed that Campylonotoidea may be closer to Atyoidea (above).[8][13]
Crangonoidea
Crangon crangon		 Contains two families: including the family Crangonidae.[8] Crangon crangon is abundant around the European coast has a sandy brown colour which it can change to match it environment. It lives in shallow water which can be slightly brackish, and it feeds nocturnally. During the day, it stays buried in the sand to escape predatory birds and fish, with only its antennae protruding.[14]
Galatheacaridoidea Contains only one species, the rare Galatheacaris abyssalis. Described in 1997 on the basis of what was then a single specimen, it was seen to be so different from previously known shrimp species that a new family Galatheacarididae and superfamily Galatheacaridoidea were erected for it.[15] Molecular phylogenetic analyses has indicated that Galatheacaris abyssalis is the larval stage of Eugonatonotus.[16]
Nematocarcinoidea
Rhynchocinetes durbanensis		 Contains four families.[9][17] They share the presence of strap-like epipods on at least the first three pairs of pereiopods, and a blunt molar process.[18] One of the families, Rhynchocinetidae, are a group of small, reclusive red-and-white shrimp. This family typically has an upward-hinged foldable rostrum,[19] hence its taxon name "Rhynchocinetidae", which means "movable beak".[19] Pictured is Rhynchocinetes durbanensis.
Oplophoroidea
Hymenodora glacialis		 There is only one family, Oplophoridae, of this pelagic shrimp, which contains 12 genera.[8]
Palaemonoidea
Palaemon elegans		 Contains 8 families and nearly 1,000 species.[8] The position of the family Typhlocarididae is unclear, although the monophyly of a group containing the remaining seven families is well supported.[13]
Pandaloidea
Heterocarpus ensifer		 Contains two family. The larger family Pandalidae has 23 genera and about 200 species, including some of commercial significance.[8]
Pasiphaeoidea Contains one family with seven extant genera.[8]
Physetocaridoidea Contains a single family with only one rare species, Physetocaris microphthalma.[8] Adult Physetocaris microphthalma have no eyes, and cannot form a claw because they are missing the last segment of the first pereiopod. They also have reduced gills and mouthparts, and no exopods on the pereiopods.[20]
Processoidea Contains a single family comprising 65 species in 5 genera.[8] These small nocturnal shrimp live mostly in shallow seas, particularly on grass flats. The first pereiopods are usually asymmetrical, with a claw on one but not the other. The rostrum is generally a simple projection from the front of the carapace, with two teeth, one at the tip, and one further back.[21]
Psalidopodoidea
Psalidopus huxleyi		 Contains a single family comprising three species, one in the western Atlantic Ocean, and two in the Indo-Pacific.[8][22][23]
Stylodactyloidea Contains a single family made up of five genera.[8]
Difference between carideans and dendrobranchiates
Carideans, such as Pandalus borealis, typically have two pairs of claws, and the second segment of the abdomen overlaps the segments on either side. The abdomen shows a pronounced caridean bend.
Dendrobranchiata, such as Penaeus monodon, typically have three pairs of claws, and even-sized segments on the abdomen. There is no pronounced bend in the abdomen.

Shrimp of the infraorder Caridea are more closely related to lobsters and crabs than they are to the members of the sub-order Dendrobranchiata (prawns).[24] Biologists distinguish these two groups based on differences in their gill structures. The gill structure is lamellar in carideans but branching in dendrobranchiates. The easiest practical way to separate true shrimp from dendrobranchiates is to examine the second abdominal segment. The second segment of a carideans overlaps both the first and the third segment, while the second segment of a dendrobranchiate overlaps only the third segment.[25] They also differ in that carideans typically have two pairs of chelae (claws), while dendrobranchiates have three.[26] A third group, the Stenopodidea, contains around 70 species and differs from the other groups in that the third pairs of legs is greatly enlarged.[26]

Procarididea are the sister group to the Caridea, comprising only eleven species.[8][9]

Fossil record

The fossil record of the Caridean is sparse, with only 57 exclusively fossil species known.[8] The earliest of these cannot be assigned to any family, but date from the Lower Jurassic and Cretaceous.[27] A number of extinct genera cannot be placed in any superfamily:[8]

References

  1. 1 2 S. De Grave; Y. Cai; A. Anker (2008). Estelle Virginia Balian; C. Lévêque; H. Segers; K. Martens, eds. "Global diversity of shrimps (Crustacea: Decapoda: Caridea) in freshwater". Hydrobiologia. Springer. 595 (1: Freshwater Animal Diversity Assessment): 287–293. doi:10.1007/s10750-007-9024-2. ISBN 978-1-4020-8258-0.
  2. 1 2 3 4 5 6 Fenner A. Chace, Jr. & Donald P. Abbott (1980). "Caridea: the shrimps". In Robert Hugh Morris, Donald Putnam Abbott & Eugene Clinton Haderlie. Intertidal Invertebrates of California. Stanford University Press. pp. 567–576. ISBN 978-0-8047-1045-9.
  3. ↑ Gary C. B. Poore; Shane T. Ahyong (2004). "Caridea – shrimps". Marine Decapod Crustacea of Southern Australia: a Guide to Identification. CSIRO Publishing. pp. 53–57. ISBN 9780643069060.
  4. ↑ "A bouillabaisse of fascinating facts about fish". NOAA: National Marine Fisheries Service. Retrieved October 22, 2009.
  5. 1 2 3 4 Based on data sourced from the FishStat database, FAO.
  6. ↑ Pandalus borealis (Krøyer, 1838) FAO, Species Fact Sheet. Retrieved September 2012.
  7. ↑ Crangon crangon (Linnaeus, 1758) FAO, Species Fact Sheet. Retrieved September 2012.
  8. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 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.
  9. 1 2 3 S. De Grave & C. H. J. M. Fransen (2011). "Carideorum Catalogus: the Recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda)". Zoologische Mededelingen. 85 (9): 195–589, figs. 1–59. ISBN 978-90-6519-200-4.
  10. ↑ Michael Türkay (2012). "Alpheoidea". World Register of Marine Species. Retrieved February 8, 2012.
  11. ↑ Joel W. Martin; George E. Davis (2001). An Updated Classification of the Recent Crustacea (PDF). Natural History Museum of Los Angeles County. pp. 132 pp.
  12. ↑ Raymond T. Bauer (2004). "Evolutionary history of the Caridea". Remarkable Shrimps: Adaptations and Natural History of the Carideans. Animal natural history series. 7. University of Oklahoma Press. pp. 204–219. ISBN 978-0-8061-3555-7.
  13. 1 2 Heather D. Bracken; Sammy De Grave; Darryl L. Felder (2009). "Phylogeny of the infraorder Caridea based on mitochondrial and nuclear genes (Crustacea: Decapoda)". In Joel W. Martin; Keith A. Crandall; Darryl L. Felder. Decapod Crustacean Phylogenetics. Crustacean issues. 18. CRC Press. pp. 281–305. ISBN 978-1-4200-9258-5.
  14. ↑ "Crangon crangon". ARKive. Retrieved June 24, 2011.
  15. ↑ Alexander L. Vereshchaka (1997). "New family and superfamily for a deep-sea caridean shrimp from the Galathea collections". Journal of Crustacean Biology. 17 (2): 361–373. doi:10.2307/1549285. JSTOR 1549285.
  16. ↑ Sammy DeGrave; Ka Hou Chu; Tin-Yam Y. Chan (2010). "On the systematic position of Galatheacaris abyssalis (Decapoda: Galatheacaridoidea)". Journal of Crustacean Biology. 30 (3): 521–527. doi:10.1651/10-3278.1.
  17. ↑ Sammy De Grave; Michael Türkay (2011). "Nematocarcinoidea". World Register of Marine Species. Retrieved September 29, 2011.
  18. ↑ Gary C. B. Poore (2004). "Superfamily Nematocarcinoidea Smith, 1884". Marine decapod Crustacea of Southern Australia: a Guide to Identification. CSIRO Publishing. pp. 115–122. ISBN 978-0-643-06906-0.
  19. 1 2 "Rhynchocinetidae". Australian Faunal Directory. Department of the Environment, Water, Heritage and the Arts. October 9, 2008. Retrieved August 13, 2010.
  20. ↑ Raymond T. Bauer (2004). "Physetocarididae". Remarkable Shrimps: Adaptations and Natural History of the Carideans. Animal natural history series. 7. University of Oklahoma Press. pp. 65–66. ISBN 978-0-8061-3555-7.
  21. ↑ Raymond B. Manning & Fenner A. Chace, Jr. (1971). "Shrimps of the family Processidae from the Northwestern Atlantic Ocean (Crustacea: Decapoda: Caridea)" (PDF). Smithsonian Contributions to Zoology. 89.
  22. ↑ Fenner A. Chace, Jr. & Lipke Holthuis (1978). "Psalidopus: the scissor-foot shrimps (Crustacea: Decapoda: Caridea)" (PDF). Smithsonian Contributions to Zoology. 277: 22 pp. doi:10.5479/si.00810282.277.
  23. ↑ Masahiro Toriyama; Hiroshi Horikawa (1993). "A new caridean shrimp, Psalidopus tosaensis, from Tosa Bay, Japan (Decapoda: Caridea, Psalidopodidae)" (PDF). Bulletin of the Nansei National Fisheries Research Institute. 26: 1–8.
  24. ↑ "Biology of Shrimps". Museum Victoria Australia. Retrieved January 9, 2010.
  25. ↑ Charles Raabe; Linda Raabe (2008). "The Caridean shrimp: Shrimp Anatomy - Illustrations and Glossary".
  26. 1 2 Raymond T. Bauer (2004). "What is a caridean shrimp?". Remarkable Shrimps: Adaptations and Natural History of the Carideans. Animal Natural History Series. 7. University of Oklahoma Press. pp. 3–14. ISBN 978-0-8061-3555-7.
  27. ↑ Fenner A. Chace, Jr. & Raymond B. Manning (1972). "Two new caridean shrimps, one representing a new family, from marine pools on Ascension Island (Crustacea: Decapoda: Natantia)" (PDF). Smithsonian Contributions to Zoology. 131: 18 pp. doi:10.5479/si.00810282.131.
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