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Alpheidae is a
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
of
caridea The Caridea, commonly known as caridean shrimp or true shrimp, are an infraorder of shrimp within the order Decapoda. This infraorder contains all species of true shrimp. They are found widely around the world in both fresh and salt water. Many ...
n 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. The family is diverse and worldwide in distribution, consisting of about 1,119 species within 38 or more genera. The two most prominent genera are ''Alpheus'' and ''Synalpheus'', with species numbering well over 250 and 100, respectively. Most snapping shrimp dig burrows and are common inhabitants of
coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of Colony (biology), colonies of coral polyp (zoology), polyps held together by calcium carbonate. Most coral reefs are built from stony corals, wh ...
s, submerged
seagrass Seagrasses are the only flowering plants which grow in marine environments. There are about 60 species of fully marine seagrasses which belong to four families ( Posidoniaceae, Zosteraceae, Hydrocharitaceae and Cymodoceaceae), all in the ...
flats, and
oyster Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not ...
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 Sonar (sound navigation and ranging or sonic navigation and ranging) is a technique that uses sound propagation (usually underwater, as in submarine navigation) to navigate, measure distances (ranging), communicate with or detect objects on o ...
and underwater communication. The shrimp are considered a major source of sound in the ocean.


Description

The "Pistol Shrimp" grows to only 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.


Ecology

Some pistol shrimp species share burrows with goby fish in a mutualistic
symbiotic relationship Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
. 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. This association has been observed in species that inhabit
coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of Colony (biology), colonies of coral polyp (zoology), polyps held together by calcium carbonate. Most coral reefs are built from stony corals, wh ...
habitats.
Eusocial Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping gen ...
behavior has been discovered in the genus ''
Synalpheus ''Synalpheus'' is a genus of snapping shrimp of the family Alpheidae, presently containing more than 100 species; new ones are described on a regular basis, and the exact number even of described species is disputed. ''Zuzalpheus'' The genus ''Z ...
''. The species ''
Synalpheus regalis ''Synalpheus regalis'' is a species of snapping shrimp that commonly live in sponges in the coral reefs along the tropical West Atlantic. They form a prominent component of the diverse marine cryptofauna of the region. For the span of their e ...
'' lives inside
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate throu ...
s in colonies that can number over 300. 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. 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. 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 Crustaceans may pass through a number of larval and immature stages between hatching from their eggs and reaching their adult form. Each of the stages is separated by a moult, in which the hard exoskeleton is shed to allow the animal to grow. The ...
, and post larval stages.


Snapping effect

The snapping shrimp competes with much larger animals such as the
sperm whale The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the genus ''Physeter'' and one of three extant species in the sperm whale famil ...
and
beluga whale The beluga whale () (''Delphinapterus leucas'') is an Arctic and sub-Arctic cetacean. It is one of two members of the family Monodontidae, along with the narwhal, and the only member of the genus ''Delphinapterus''. It is also known as the ...
for the title of loudest animal in the sea. The animal snaps a specialized claw shut to create a
cavitation Cavitation is a phenomenon in which the static pressure of a liquid reduces to below the liquid's vapour pressure, leading to the formation of small vapor-filled cavities in the liquid. When subjected to higher pressure, these cavities, ca ...
bubble that generates acoustic pressures of up to at a distance of 4 cm from the claw. As it ejects from the claw, the bubble reaches speeds of . The pressure is high enough to kill small fish. It corresponds to a peak pressure level of 218 
decibels The decibel (symbol: dB) is a relative unit of measurement equal to one tenth of a bel (B). It expresses the ratio of two values of a power or root-power quantity on a logarithmic scale. Two signals whose levels differ by one decibel have a ...
relative to one
micropascal The pascal (symbol: Pa) is the unit of pressure in the International System of Units (SI), and is also used to quantify internal pressure, stress, Young's modulus, and ultimate tensile strength. The unit, named after Blaise Pascal, is defined as ...
(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. Similar values are reported by Ferguson and Cleary. The duration of the click is less than 1 
millisecond A millisecond (from '' milli-'' and second; symbol: ms) is a unit of time in the International System of Units (SI) equal to one thousandth (0.001 or 10−3 or 1/1000) of a second and to 1000 microseconds. A unit of 10 milliseconds may be calle ...
. The snap can also produce
sonoluminescence Sonoluminescence is the emission of light from imploding bubbles in a liquid when excited by sound. History The sonoluminescence effect was first discovered at the University of Cologne in 1934 as a result of work on sonar. Hermann Frenzel ...
from the collapsing
cavitation Cavitation is a phenomenon in which the static pressure of a liquid reduces to below the liquid's vapour pressure, leading to the formation of small vapor-filled cavities in the liquid. When subjected to higher pressure, these cavities, ca ...
bubble. As it collapses, the cavitation bubble emits a short flash of light with a broad spectrum. If the light were of thermal origin it would require a temperature of the emitter of over . In comparison, the surface temperature of the
sun The Sun is the star at the center of the Solar System. It is a nearly perfect ball of hot plasma, heated to incandescence by nuclear fusion reactions in its core. The Sun radiates this energy mainly as light, ultraviolet, and infrared radi ...
is estimated to be around . The light is of lower intensity than the light produced by typical sonoluminescence and is not visible to the
naked eye Naked eye, also called bare eye or unaided eye, is the practice of engaging in visual perception unaided by a magnifying, light-collecting optical instrument, such as a telescope or microscope, or eye protection. Vision corrected to norma ...
. 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
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapoda, decapods, ostracoda, seed shrimp, branchiopoda, branchiopods, argulidae, fish lice, krill, remipedes, isopoda, isopods, barnacles, copepods, ...
s, the
mantis shrimp Mantis shrimp, or stomatopods, are carnivorous marine crustaceans of the order Stomatopoda (). Stomatopods branched off from other members of the class Malacostraca around 340 million years ago. Mantis shrimp typically grow to around in length, ...
, contains species whose club-like forelimbs can strike so quickly and with such force as to induce sonoluminescent cavitation bubbles upon impact. 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. When in colonies, the snapping shrimp can interfere with
sonar Sonar (sound navigation and ranging or sonic navigation and ranging) is a technique that uses sound propagation (usually underwater, as in submarine navigation) to navigate, measure distances (ranging), communicate with or detect objects on o ...
and underwater communication. The shrimp are a major source of noise in the ocean and can interfere with
anti-submarine warfare Anti-submarine warfare (ASW, or in older form A/S) is a branch of underwater warfare that uses surface warships, aircraft, submarines, or other platforms, to find, track, and deter, damage, or destroy enemy submarines. Such operations are typ ...
.


Genera

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 ''Synalpheus'' is a genus of snapping shrimp of the family Alpheidae, presently containing more than 100 species; new ones are described on a regular basis, and the exact number even of described species is disputed. ''Zuzalpheus'' The genus ''Z ...
'', with 146 species. *'' Acanthanas'' Anker, Poddoubtchenko & Jeng, 2006 *'' Alpheopsis'' Coutière, 1896 *'' Alpheus'' Fabricius, 1798 *'' Amphibetaeus'' Coutière, 1896 *''
Arete ''Arete'' (Greek: ) is a concept in ancient Greek thought that, in its most basic sense, refers to 'excellence' of any kind Liddell, H.G. & Scott, R. '' A Greek–English Lexicon'', 9th ed. (Oxford, 1940), s.v.br>—especially a person or thi ...
'' Stimpson, 1860 *'' Aretopsis'' De Man, 1910 *''
Athanas ''Athanas'' is a genus of shrimp of the family Alpheidae. These are small shrimp measuring 2 cm in length. Females have smaller chelae than males. Some species, including ''A. djiboutensis'', are called "bulldozer shrimp" because of the wa ...
'' Leach, 1814 *'' Athanopsis'' Coutière, 1897 *''
Automate Automation describes a wide range of technologies that reduce human intervention in processes, namely by predetermining decision criteria, subprocess relationships, and related actions, as well as embodying those predeterminations in machines ...
'' De Man, 1888 *'' Bannereus'' Bruce, 1988 *''
Batella ''Batella'' is a genus of snapping shrimp comprising three species: *'' Batella leptocarpus'' Chace, 1988 *'' Batella parvimanus'' (Bate, 1888) (syn. The Botanical and Zoological Codes of nomenclature treat the concept of synonymy different ...
'' Holthuis, 1955 *'' Bermudacaris'' Anker & Iliffe, 2000 *'' Betaeopsis'' Yaldwyn, 1971 *'' Betaeus'' Dana, 1852 *'' Bruceopsis'' Anker, 2010 *'' Coronalpheus'' Wicksten, 1999 *'' Coutieralpheus'' Anker & Felder, 2005 *''
Deioneus In Greek mythology, Deioneus (; Ancient Greek: Δηιονεύς means "ravager") or Deion (; Ancient Greek: ) is a name attributed to the following individuals: *Deioneus, king of Phocis and son of King Aeolus of Aeolia and Enarete, daughter ...
'' 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 In Greek mythology, Salmoneus (; Ancient Greek: Σαλμωνεύς) was 'the wicked'Hesiod, '' Ehoiai'' fr. 4 as cited in Plutarch, ''Moralia'' p. 747; Scholia on Pindar, ''Pythian Ode'' 4.263 eponymous king and founder of Salmone in Pisatis. ...
'' Holthuis, 1955 *'' Stenalpheops'' Miya, 1997 *''
Synalpheus ''Synalpheus'' is a genus of snapping shrimp of the family Alpheidae, presently containing more than 100 species; new ones are described on a regular basis, and the exact number even of described species is disputed. ''Zuzalpheus'' The genus ''Z ...
'' Bate, 1888 *'' Thuylamea'' Nguyên, 2001 *'' Triacanthoneus'' Anker, 2010 *'' Vexillipar'' Chace, 1988 *'' Yagerocaris'' Kensley, 1988


References


External links


How snapping shrimp snap
University of Twente The University of Twente (Dutch: ''Universiteit Twente''; , abbr. ) is a public technical university located in Enschede, Netherlands. The university has been placed in the top 170 universities in the world by multiple central ranking tables. I ...

Article on pistol shrimp going into physical details

Radiolab episode: ''Bigger Than Bacon''
– the history and science of snapping shrimp {{Taxonbar, from=Q311534 Taxa named by Constantine Samuel Rafinesque Decapod families