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Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Crustacea
Superclassis: Multicrustacea
Classis: Malacostraca
Subclassis: Eumalacostraca
Superordo: Eucarida
Ordo: Decapoda
Subordo: Pleocyemata
Infraordo: Caridea
Superfamilia: Alpheoidea

Familia: Alpheidae
Genera: AlpheopsisAlpheus – Amphibetaeus – Arete – Aretopsis – AthanasAthanopsis – Automate – Bannereus – Batella – Betaeopsis – BetaeusBruceopsis – Caligoneus – Fenneralpheus – Hamalpheus – HarperalpheusJengalpheops – Leptalpheus – LeptathanasLeslibetaeus – Hamalpheus – Metalpheus – Mohocaris – Nennalpheus – Neoalpheopsis – Notalpheus – Parabetaeus – Pomagnathus – Potamalpheops – Prionalpheus – Pseudathanas – Pterocaris – Racilius – Richalpheus – SalmoneusSynalpheusTriacanthoneus – Vexillipar

Name

Alpheidae Rafinesque, 1815

Type genus: Alpheus Fabricius, 1798
References

Rafinesque, C.S. 1815. Analyse de la nature, ou tableau de l'univers et des corps organisés. Palerme: L'Imprimerie de Jean Barravecchia. 224 pp. BHL Reference page.

Almeida, A.O., Terossi, M., Buranelli, R.C., Castilho, A.L., Costa, R.C., Zara, F.J. & Mantelatto, F.L. 2018. Checklist of decapods (Crustacea) from the coast of São Paulo State (Brazil) supported by integrative molecular and morphological data: II. Infraorder Caridea: family Alpheidae. Zootaxa 4450(3): 331–358. DOI: 10.11646/zootaxa.4450.3.2 Paywall Reference page.
Anker, A. 2010: Description of a new genus and two new species of alpheid shrimps from Guam (Crustacea, Decapoda). Pp. 389-404 in De Grave, S.; Fransen, C.H.J.M. (eds.): Contributions to shrimp taxonomy. Zootaxa, 2372: 1–414. Preview
Anker, A. 2010: New findings of rare or little-known alpheid shrimp genera (Crustacea, Decapoda) in Moorea, French Polynesia. Zootaxa, 2403: 23–41. Preview
Anker, A. 2010: A new genus and three new species of alpheid shrimps (Crustacea, Decapoda, Caridea) from the tropical American coasts. Zootaxa, 2652: 47–63. Preview.
Anker, A. 2015: Description of a new distinctive species of Parabetaeus Coutière, 1897 (Decapoda: Caridea: Alpheidae) from the Indo-West Pacific. Zootaxa 3957(5): 585–595. DOI: 10.11646/zootaxa.3957.5.7. Preview (PDF) Reference page.
Anker, A. 2015: Alpheopsis balaeniceps sp. nov., an unusual alpheid shrimp (Decapoda: Caridea) from French Polynesia. Zootaxa 3972(1): 85–92. DOI: 10.11646/zootaxa.3972.1.6. Preview (PDF) Reference page.
Anker, A. 2017. A new deep-water species of Alpheopsis Coutière, 1897 from the Indo-West Pacific (Decapoda: Alpheidae). Zootaxa 4226(4): 571–580. DOI: 10.11646/zootaxa.4226.4.7. Full article (PDF) Reference page.
Anker, A. 2017. Strongly carinate species of Alpheopsis Coutière, 1897 of the tropical Atlantic and eastern Pacific, with redescription of A. trigona (Rathbun, 1901) and description of three new species (Malacostraca: Decapoda: Alpheidae). Zootaxa 4277(2): 199–227. DOI: 10.11646/zootaxa.4277.2.2. Reference page.
Anker, A. & M.-S. Jeng, 2006: RICHALPHEUS PALMERI, N. GEN., N. SP., AN INFAUNAL ALPHEID SHRIMP FROM THE PHILIPPINES, WITH REDESCRIPTION OF AMPHIBETAEUS JOUSSEAUMEI (COUTIÈRE, 1896) (DECAPODA: CARIDEA). Journal of Crustacean Biology 26(3): 379–391. Abstract: DOI: 10.1651/S-2647.1.
Anker, A.; Poddoubtchenko, D.; Wehrtmann, I.S. 2006: Leslibetaeus coibita, n. gen., n. sp., a new alpheid shrimp from the Pacific coast of Panama (Crustacea: Decapoda). Zootaxa, 1183: 27–41. Abstract & excerpt
Komai, T. & Fujita, Y. 2018. A new genus and new species of alpheid shrimp from a marine cave in the Ryukyu Islands, Japan, with additional record of Salmoneus antricola Komai, Yamada & Yunokawa, 2015 (Crustacea: Decapoda: Caridea). Zootaxa 4369(4): 575–586. DOI: 10.11646/zootaxa.4369.4.7 Reference page.
Wang, Y-R. & Sha, Z-L. 2017. Description of two new species of the genus Automate De Man, 1888 (Crustacea: Decapoda: Caridea) from the South China Sea. Zootaxa 4238(1): 30–42. DOI: 10.11646/zootaxa.4238.1.2. Reference page.

Links

Analyse Nature: 98
Alpheidae Rafinesque, 1815 – Taxon details on Integrated Taxonomic Information System (ITIS).

Vernacular names
English: Snapping shrimp, Pistol shrimp, Alpheid shrimp
日本語: テッポウエビ科
русский: Раки-щелкуны
ไทย: กุ้งดีดขัน, กุ้งกระเตาะ
中文: 鼓虾科

Alpheidae is a family of caridean snapping shrimp, characterized by having asymmetrical claws, the larger of which is typically capable of producing a loud snapping sound. Other common names for animals in the group are pistol shrimp or alpheid shrimp.[citation needed]

The family is diverse and worldwide in distribution, consisting of about 1,119[citation needed] species within 38 or more genera.[1] The two most prominent genera are Alpheus and Synalpheus, with species numbering well over 250 and 100, respectively.[2][3] Most snapping shrimp dig burrows and are common inhabitants of coral reefs, submerged seagrass flats, and oyster reefs. While most genera and species are found in tropical and temperate coastal and marine waters, Betaeus inhabits cold seas and Potamalpheops is found only in freshwater caves.

When in colonies, the snapping shrimp can interfere with sonar and underwater communication. The shrimp are considered a major source of noise in the ocean.[4]

Description

The snapping shrimp grows to only 3–5 cm (1.2–2.0 in) long. It is distinctive for its disproportionately large claw, larger than half the shrimp's body. The claw can be on either arm of the body, and, unlike most shrimp claws, does not have typical pincers at the end. Rather, it has a pistol-like feature made of two parts. A joint allows the "hammer" part to move backward into a right-angled position. When released, it snaps into the other part of the claw, emitting an enormously powerful wave of bubbles capable of stunning larger fish and breaking small glass jars.[5]
Ecology
Alpheus randalli with a goby of the genus Amblyeleotris

Some pistol shrimp species share burrows with goby fish in a mutualistic symbiotic relationship. The burrow is built and tended by the pistol shrimp, and the goby provides protection by watching out for danger. When both are out of the burrow, the shrimp maintains contact with the goby using its antennae. The goby, having better vision, alerts the shrimp of danger using a characteristic tail movement, and then both retreat into the safety of the shared burrow.[6] This association has been observed in species that inhabit coral reef habitats.

Eusocial behavior has been discovered in the genus Synalpheus. The species Synalpheus regalis lives inside sponges in colonies that can number over 300.[7] All of them are the offspring of a single large female, the queen, and possibly a single male. The offspring are divided into workers who care for the young and predominantly male soldiers who protect the colony with their huge claws.[7]

Pistol shrimp have the ability to reverse claws. When the snapping claw is lost, the missing limb will regenerate into a smaller claw and the original smaller appendage will grow into a new snapping claw. Laboratory research has shown that severing the nerve of the snapping claw induces the conversion of the smaller limb into a second snapping claw. The reversal of claw asymmetry in snapping shrimp is thought to be unique in nature.[8]

The claw of the snapping shrimp is a dimorphic addition to the arsenal of the shrimp. The snapping shrimp species will retain the same mate after copulation, making them monogamous. Most females of the Alpheidae species are susceptible to mating. Young females become receptive to males either just before (premolt stage) or after the puberty molt, making them physiologically mature and morphologically able to carry the egg mass. Male presence during the molt is beneficial for the female, as searching for a male during her soft‐bodied receptive phase would put her at mortal risk. Mates have more success with partners having greater body mass. The larger shrimp are most successful. These animals practice mate guarding, leading to a decline in mate competition, as well as bonding of partners. The male and female will defend their shelter to protect both territory and young. Larva develop in three stages: The nauplius larvae, zoea, and post larval stages.
Snapping effect
Snapping shrimp claw action. 1. closed pistol shrimp claw with hidden plunger (P). 2. open claw with exposed (P) and chamber (C). 3. open claw with water (W) entering (C). 4. claw with (P) pushed into chamber (C), forcing jet stream (J) out of (C).

The snapping shrimp competes with much larger animals such as the sperm whale and beluga whale for the title of loudest animal in the sea. The animal snaps a specialized claw shut to create a cavitation bubble that generates acoustic pressures of up to 80 kilopascals (12 psi) at a distance of 4 cm from the claw. As it ejects from the claw, the bubble reaches speeds of 100 km/h (62 mph).[citation needed] The pressure is high enough to kill small fish.[9] It corresponds to a peak pressure level of 218 decibels relative to one micropascal (dB re 1 μPa), equivalent to a zero to peak source level of 190 dB re 1 μPa m. Au and Banks measured peak to peak source levels between 185 and 190 dB re 1 μPa m, depending on the size of the claw.[10] Similar values are reported by Ferguson and Cleary.[11] The duration of the click is less than 1 millisecond.

The snap can also produce sonoluminescence from the collapsing cavitation bubble. As it collapses, the cavitation bubble reaches temperatures of over 5,000 K (4,700 °C).[12] In comparison, the surface temperature of the sun is estimated to be around 5,772 K (5,500 °C).[13] The light is of lower intensity than the light produced by typical sonoluminescence and is not visible to the naked eye. It is most likely a by-product of the shock wave with no biological significance. However, it was the first known instance of an animal producing light by this effect. It has subsequently been discovered that another group of crustaceans, the mantis shrimp, contains species whose club-like forelimbs can strike so quickly and with such force as to induce sonoluminescent cavitation bubbles upon impact.[14]

The snapping is used for hunting (hence the alternative name "pistol shrimp"), as well as for communication. When hunting, the shrimp usually lies in an obscured spot, such as a burrow. The shrimp then extends its antennae outwards to determine if any fish are passing by. Once it feels movement, the shrimp inches out of its hiding place, pulls back its claw, and releases a "shot" which stuns the prey; the shrimp then pulls it to the burrow and feeds on it.[citation needed]

When in colonies, the snapping shrimp can interfere with sonar and underwater communication.[4][15][16] The shrimp are a major source of noise in the ocean[4] and can interfere with anti-submarine warfare.[17][18]
Genera
Betaeopsis aequimanus
Synalpheus fritzmuelleri

More than 620 species are currently recognised in the family Alpheidae, distributed among 45 genera. The largest of these are Alpheus, with 283 species, and Synalpheus, with 146 species.[19]

Acanthanas Anker, Poddoubtchenko & Jeng, 2006
Alpheopsis Coutière, 1896
Alpheus Fabricius, 1798
Amphibetaeus Coutière, 1896
Arete Stimpson, 1860
Aretopsis De Man, 1910
Athanas Leach, 1814
Athanopsis Coutière, 1897
Automate De Man, 1888
Bannereus Bruce, 1988
Batella Holthuis, 1955
Bermudacaris Anker & Iliffe, 2000
Betaeopsis Yaldwyn, 1971
Betaeus Dana, 1852
Bruceopsis Anker, 2010
Coronalpheus Wicksten, 1999
Coutieralpheus Anker & Felder, 2005
Deioneus Dworschak, Anker & Abed-Navandi, 2000
Fenneralpheus Felder & Manning, 1986
Harperalpheus Felder & Anker, 2007
Jengalpheops Anker & Dworschak, 2007
Leptalpheus Williams, 1965
Leptathanas De Grave & Anker, 2008
Leslibetaeus Anker, Poddoubtchenko & Wehrtmann, 2006
Metabetaeus Borradaile, 1899
Metalpheus Coutière, 1908
Mohocaris Holthuis, 1973
Nennalpheus Banner & Banner, 1981
Notalpheus G. Méndez & Wicksten, 1982
Orygmalpheus De Grave & Anker, 2000
Parabetaeus Coutière, 1896
Pomagnathus Chace, 1937
Potamalpheops Powell, 1979
Prionalpheus Banner & Banner, 1960
Pseudalpheopsis Anker, 2007
Pseudathanas Bruce, 1983
Pterocaris Heller, 1862
Racilius Paul’son, 1875
Richalpheus Anker & Jeng, 2006
Rugathanas Anker & Jeng, 2007
Salmoneus Holthuis, 1955
Stenalpheops Miya, 1997
Synalpheus Bate, 1888
Thuylamea Nguyên, 2001
Triacanthoneus Anker, 2010
Vexillipar Chace, 1988
Yagerocaris Kensley, 1988

References

A. Anker; S. T. Ahyong; P. Y. Noel; A. R. Palmer (2006). "Morphological phylogeny of alpheid shrimps: parallel preadaptation and the origin of a key morphological innovation, the snapping claw". Evolution. 60 (12): 2507–2528. doi:10.1554/05-486.1. PMID 17263113. S2CID 18414340.
W. Kim; L. G. Abele (1988). "The snapping shrimp genus Alpheus from the Eastern Pacific (Decapoda: Caridea: Alpheidae)". Smithsonian Contributions to Zoology. 454 (454): 1–119. doi:10.5479/si.00810282.454.
Fenner A. Chace Jr. (1988). "The Caridean Shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907–1910, Part 5: Family Alpheidae" (PDF). Smithsonian Contributions to Zoology. 466: 1–99.
"Shrimp, bubble and pop". BBC News. September 21, 2000. Retrieved July 2, 2011.
Maurice Burton; Robert Burton (1970). The International Wildlife Encyclopedia, Volume 1. Marshall Cavendish. p. 2366.
I. Karplus (1987). "The association between gobiid fishes and burrowing alpheid shrimps". Oceanography and Marine Biology: An Annual Review. 25: 507–562.
J. E. Duffy (1996). "Eusociality in a coral-reef shrimp". Nature. 381 (6582): 512–514. doi:10.1038/381512a0. S2CID 33166806.
M. R. McClure (1996). "Symmetry of large claws in snapping shrimp in nature (Crustacea: Decapoda: Alpheidae)". Crustaceana. 69 (7): 920–921. doi:10.1163/156854096X00321.
M. Versluis; B. Schmitz; A. von der Heydt; D. Lohse (2000). "How snapping shrimp snap: through cavitating bubbles". Science. 289 (5487): 2114–2117. doi:10.1126/science.289.5487.2114. PMID 11000111.
W. W. L. Au; K. Banks (1998). "The acoustics of the snapping shrimp Synalpheus parneomeris in Kaneohe Bay". Journal of the Acoustical Society of America. 103 (1): 41–47. doi:10.1121/1.423234.
B. G. Ferguson; J. L. Cleary (2001). "In situ source level and source position estimates of biological transient signals produced by snapping shrimp in an underwater environment". Journal of the Acoustical Society of America. 109 (6): 3031–3037. doi:10.1121/1.1339823. PMID 11425145.
D. Lohse; B. Schmitz; M. Versluis (2001). "Snapping shrimp make flashing bubbles". Nature. 413 (6855): 477–478. doi:10.1038/35097152. PMID 11586346. S2CID 4429684.
Williams, D.R. (1 July 2013). "Sun Fact Sheet". NASA Goddard Space Flight Center. Archived from the original on 15 July 2010. Retrieved 12 August 2013.
S. N. Patek; R. L. Caldwell (2005). "Extreme impact and cavitation forces of a biological hammer: strike forces of the peacock mantis shrimp" (PDF). The Journal of Experimental Biology. 208 (19): 3655–3664. doi:10.1242/jeb.01831. PMID 16169943. S2CID 312009.
Kenneth Chang (September 26, 2000). "Sleuths solve case of bubble mistaken for a snapping shrimp". The New York Times. p. 5. Retrieved July 2, 2011.
"Sea creatures trouble sonar operators – new enzyme". The New York Times. February 2, 1947. Retrieved July 2, 2011.
Stuart Rock. "Submarine hunting in Somerset" (PDF). thalesgroup.com. Archived from the original (PDF) on 27 March 2018. Retrieved 26 March 2018.]
"Underwater Drones Join Microphones to Listen for Chinese Nuclear Submarines - AUVAC". auvac.org. Archived from the original on 23 July 2018. Retrieved 26 March 2018.
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.

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