Mantis shrimps arecarnivorousmarinecrustaceans of theorderStomatopoda (fromAncient Greekστόμα (stóma) 'mouth' andποδός (podós) 'foot'). Stomatopodsbranched off from other members of the classMalacostraca around 400 million years ago,^[2] with more than 520extant species of mantis shrimp known. All living species are in thesuborderUnipeltata, which arose around 250 million years ago.^[2][3] They are among the most importantpredators in manyshallow,tropical andsubtropical marinehabitats. However, despite being common in their habitats, they are poorly understood, as many species spend most of their lives sheltering in burrows and holes.^[4]

Mantis shrimp Temporal range:Carboniferous–Recent PreꞒ Ꞓ O S D C P T J K Pg N
Odontodactylus scyllarus (Red mantis shrimp)
Lysiosquillina maculata (Zebra mantis shrimp)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Malacostraca
Subclass:	Hoplocarida
Order:	Stomatopoda Latreille, 1817
Subdivisions[1]
Superfamilies and families Bathysquilloidea Bathysquillidae Indosquillidae Gonodactyloidea Alainosquillidae Hemisquillidae Gonodactylidae Odontodactylidae Protosquillidae Pseudosquillidae Takuidae Erythrosquilloidea Erythrosquillidae Lysiosquilloidea Coronididae Lysiosquillidae Nannosquillidae Tetrasquillidae Squilloidea Squillidae Eurysquilloidea Eurysquillidae Parasquilloidea Parasquillidae

Dubbed "sea locusts" byancientAssyrians, "prawn killers" inAustralia,^[5] and now sometimes referred to as "thumb splitters" due to their ability to inflict painful wounds if handled incautiously,^[6] mantis shrimp possess powerfulraptorial appendages that are used to attack and kill prey either by spearing, stunning, ordismembering; the shape of these appendages are often used to classify them into groups: extant mantis shrimp either have appendages which form heavilymineralized "clubs" that can strike with great power, or they have sharp, grasping forelimbs used to swiftly seize prey (similar to those ofpraying mantis, hence theircommon name).

Mantis shrimp typically grow to around 10 cm (3.9 in) in length, while a few species such as the zebra mantis shrimp can reach up to 38 cm (15 in).^[7] A mantis shrimp'scarapace covers only the rear part of the head and the first four segments of thethorax. Mantis shrimp widely range in colour, with species mostly being shades of brown to having multiple contrasting, vivid colours.

The mantis shrimp's second pair of thoracic appendages is adapted for powerful close-range combat. These claws can accelerate at a rate comparable to that of a.22 caliber bullet when fired, having around 1500 newtons of force with each swing/attack.^[8] The appendage differences divide mantis shrimp into two main types: those that hunt by impaling their prey with spear-like structures and those that smash prey with a powerful blow from a heavily mineralised club-like appendage. A considerable amount of damage can be inflicted after impact with these robust, hammer-like claws. This club is further divided into three subregions: the impact region, the periodic region, and the striated region. Mantis shrimp are commonly separated into distinct groups (most are categorized as either spearers or smashers but there are some outliers)^[9] as determined by the type of claws they possess:

• Spearers are armed with spiny appendages - the spines having barbed tips - used to stab and snag prey. These raptorial appendages resemble those ofpraying mantids, hence the common name of these crustaceans. This is the type found in most mantis shrimp.^[10]
• Smashers possess a much more developed club and a more rudimentary spear (which is nevertheless quite sharp and still used in fights between their own kind); the club is used to bludgeon and smash their prey apart. The inner aspect of the terminal portion of the appendage can also possess a sharp edge, used to cut prey while the mantis shrimp swims. This is found in the families Gonodactylidae, Odontodactylidae, Protosquillidae, and Takuidae.^[10]
• Spike Smashers (hammers or primitive smashers): An unspecialized form, found only in the basal family Hemisquillidae. The last segment lacks spines except at the tip, so it is not as effective at spearing but can also be used for smashing.^{[10][11][12][13]}
• Hatchet: An unusual, highly derived appendage that only a few species have. This body plan is largely unresearched.^[11][12][14]

Both types strike by rapidly unfolding and swinging theirraptorial claws at the prey, and can inflict serious damage on victims significantly greater in size than themselves. In smashers, these two weapons are employed with blinding quickness, with an acceleration of 10,400 g (102,000 m/s² or 335,000 ft/s²) and speeds of 23 m/s (83 km/h; 51 mph) from a standing start.^[15] Because they strike so rapidly, they generate vapor-filled bubbles in the water between the appendage and the striking surface—known ascavitation bubbles.^[15] The collapse of these cavitation bubbles produces measurable forces on their prey in addition to the instantaneous forces of 1,500 newtons that are caused by the impact of the appendage against the striking surface, which means that the prey is hit twice by a single strike; first by the claw and then by the collapsing cavitation bubbles that immediately follow.^[16] Even if the initial strike misses the prey, the resultingshock wave can be enough to stun or kill.

Smashers use this ability to attackcrabs,snails,rock oysters, and othermolluscs, their blunt clubs enabling them to crack the shells of their prey into pieces. Spearers, however, prefer the meat of softer animals, such asfish andcephalopods, which their barbed claws can more easily slice and snag.

The appendages are being studied as a microscale analogue for new macroscale material structures.^{[17][clarification needed]}

The eyes of the mantis shrimp are mounted on mobilestalks and can move independently of each other. The extreme mobility allows them to be rotated in all three dimensions, yet the position of their eyes has shown to have no effect on the perception of their surroundings.^[18] They are thought to have the most complex eyes in the animal kingdom and have the most complex front-end for any visual system ever discovered.^[19][20][21]

Eachcompound eye is made up of tens of thousands ofommatidia, clusters of photoreceptor cells.^[20] Each eye consists of two flattened hemispheres separated by parallel rows of specialised ommatidia, collectively called the midband. The number of omatidial rows in the midband ranges from two to six.^[19][20] This divides the eye into three regions. This configuration enables mantis shrimp to see objects that are near the mid-plane of an eye with three parts of the same eye (as can be seen in some photos showing threepseudopupils in one eye). In other words, each eye possessestrinocular vision, and thereforedepth perception, for objects near its mid-plane. The upper and lower hemispheres are used primarily for recognition of form and motion, like the eyes of many other crustaceans.^[19]

Compared with the three types ofphotoreceptor cell that humans possess in their eyes, the eyes of a mantis shrimp have between 12 and 16 types of photoreceptor cells. Furthermore, some of these stomatopods can tune the sensitivity of their long wavelength colour vision to adapt to their environment.^[22] This phenomenon, called "spectral tuning", is species-specific.^[23] Cheroske et al. did not observe spectral tuning inNeogonodactylus oerstedii, the species with the most monotonous natural photic environment. InN. bredini, a species with a variety of habitats ranging from a depth of 5 to 10 m (although it can be found down to 20 m below the surface), spectral tuning was observed, but the ability to alter wavelengths of maximum absorbance was not as pronounced as inN. wennerae, a species with much higher ecological/photic habitat diversity. The diversity of spectral tuning in Stomatopoda is also hypothesised to be directly linked to mutations in theretinal binding pocket of theopsin.^[24]

The huge diversity seen in mantis shrimp photoreceptors likely comes from ancientgene duplication events.^[25][26] One consequence of this duplication is the lack of correlation between opsin transcript number and physiologically expressed photoreceptors.^[25] One species may have six different opsin genes, but only express one spectrally distinct photoreceptor. Over the years, some mantis shrimp species have lost the ancestral phenotype, although some still maintain 16 distinct photoreceptors and four light filters. Species that live in a variety of photic environments have high selective pressure for photoreceptor diversity, and maintain ancestral phenotypes better than species that live in murky waters or are primarily nocturnal.^[25][27]

Mantis shrimp can perceive wavelengths of light ranging fromdeep ultraviolet (300 nm) tofar-red (720 nm) andpolarised light.^[20][28] In mantis shrimp in the superfamilies Gonodactyloidea, Lysiosquilloidea, and Hemisquilloidea, the midband is made up of six ommatidial rows. Rows 1 to 4 process colours, while rows 5 and 6 detectcircularly orlinearly polarised light. Twelve types of photoreceptor cells are in rows 1 to 4, four of which detect ultraviolet light.^{[19][20][28][29]} Despite the impressive range of wavelengths that mantis shrimp have the ability to see, they do not have the ability to discriminate wavelengths less than 25 nm apart.^{[clarification needed]} It is suggested that not discriminating between closely positioned wavelengths allows these organisms to make determinations of its surroundings with little processing delay. Having little delay in evaluating surroundings is important for mantis shrimp, since they are territorial and frequently in combat.^[28] However, some mantis shrimp have been found capable of distinguishing between high-saturation and low-saturation colors.^[30]

Rows 1 to 4 of the midband are specialised for colour vision, from deep ultraviolet to far red. Their UV vision can detect five different frequency bands in the deep ultraviolet. To do this, they use two photoreceptors in combination with four different colour filters.^[31][32] They are currently believed insensitive to infrared light.^[33] The optical elements in these rows have eight different classes of visual pigments and therhabdom (area of eye that absorbs light from a single direction) is divided into three differentpigmented layers (tiers), each for different wavelengths. The three tiers in rows 2 and 3 are separated by colour filters (intrarhabdomal filters) that can be divided into four distinct classes, two classes in each row. Each consists of a tier, a colour filter of one class, a tier again, a colour filter of another class, and then a last tier. These colour filters allow the mantis shrimp to see with diverse colour vision. Without the filters, the pigments themselves range only a small segment of the visual spectrum, about 490 to 550 nm.^[25] Rows 5 and 6 are also segregated into different tiers, but have only one class of visual pigment, the ninth class, and are specialised for polarisation vision. Depending upon the species, they can detect circularly polarised light, linearly polarised light, or both. A tenth class of visual pigment is found in the upper and lower hemispheres of the eye.^[19]

Some species have at least 16 photoreceptor types, which are divided into four classes (their spectral sensitivity is further tuned by colour filters in the retinas), 12 for colour analysis in the different wavelengths (including six which are sensitive to ultraviolet light^[31][34]) and four for analysing polarised light. By comparison, most humans have only four visual pigments, of which three are dedicated to see colour, and human lenses block ultraviolet light. The visual information leaving theretina seems to be processed into numerous paralleldata streams leading into thebrain, greatly reducing the analytical requirements at higher levels.^[35]

The midband covers only about 5 to 10° of the visual field at any given instant, but like most crustaceans, mantis shrimps' eyes are mounted on stalks. In mantis shrimps, the movement of the stalked eye is unusually free, and can be driven up to 70° in all possible axes of movement by eight eyecup muscles divided into six functional groups. By using these muscles to scan the surroundings with the midband, they can add information about forms, shapes, and landscape, which cannot be detected by the upper and lower hemispheres of the eyes. They can also track moving objects using large, rapid eye movements where the two eyes move independently. By combining different techniques, including movements in the same direction, the midband can cover a very wide range of the visual field.^{[citation needed]}

Six species of mantis shrimp have been reported to be able to detect circularly polarised light, which has not been documented in any other animal, and whether it is present across all species is unknown.^[36][37][38] They perform this feat by converting circularly polarized light into linearly polarized light via quarter-waveplates formed from stacks ofmicrovilli. Some of their biological quarter-waveplates perform more uniformly over the visual spectrum than any current man-made polarising optics, and this could inspire new types of optical media that would outperform early 21st centuryBlu-ray Disc technology.^[39][40]

The speciesGonodactylus smithii is the only organism known to simultaneously detect the four linear and two circular polarisation components required to measure all fourStokes parameters, which yield a full description of polarisation. It is thus believed to have optimal polarisation vision.^[37][41] It is the only animal known to have dynamic polarisation vision. This is achieved by rotational eye movements to maximise the polarisation contrast between the object in focus and its background.^[42] Since each eye moves independently from the other, it creates two separate streams of visual information.^[43]

What advantage sensitivity to polarisation confers is unclear; however, polarisation vision is used by other animals for sexual signaling and secret communication that avoids the attention of predators.^[44] This mechanism could provide an evolutionary advantage; it only requires small changes to the cell in the eye and could easily lead tonatural selection.^[45]

The eyes of mantis shrimps may enable them to recognise different types of coral, prey species (which are often transparent or semitransparent), or predators, such asbarracuda, which have shimmering scales. Alternatively, the manner in which they hunt (very rapid movements of the claws) may require very accurate ranging information, which would require accurate depth perception. The capacity to see UV light may enable observation of otherwise hard-to-detect prey on coral reefs.^[34]

During mating rituals, mantis shrimps activelyfluoresce, and the wavelength of this fluorescence matches the wavelengths detected by their eye pigments.^[46] Females are only fertile during certain phases of thetidal cycle; the ability to perceive thephase of the moon may, therefore, help prevent wasted mating efforts. It may also give these shrimps information about the size of the tide, which is important to species living in shallow water near the shore.^{[citation needed]}

Researchers suspect that the broader variety of photoreceptors in the eyes of mantis shrimps allows visual information to be preprocessed by the eyes instead of the brain, which would otherwise have to be larger to deal with the complex task ofopponent process colour perception used by other species, thus requiring more time and energy. While the eyes themselves are complex and not yet fully understood, the principle of the system appears to be simple.^[47] It has a similar set of sensitivities to the human visual system, but works in the opposite manner. In the human brain, the inferior temporal cortex has a huge number of colour-specific neurons, which process visual impulses from the eyes to extract colour information. The mantis shrimp instead uses the different types of photoreceptors in its eyes to perform the same function as the human brain neurons, resulting in a hardwired and more efficient system for an animal that requires rapid colour identification. Humans have fewer types of photoreceptors, but more colour-tuned neurons, while mantis shrimp appear to have fewer colour neurons and more classes of photoreceptors.^[48]

However, a study from 2022 failed to find unequivocal evidence for a solely "barcode"-like visual system as described above. Stomatopods of the speciesHaptosquilla trispinosa were able to distinguish high and low-saturation colors from grey, contravening Thoen and colleagues.^[30][28] It may be that some combination ofcolor opponency and photoreceptor activation comparison/barcode analysis is present.^[30]

The shrimps use a form of reflector of polarised light not seen in nature or human technology before. It allows the manipulation of light across the structure rather than through its depth, the typical way polarisers work. This allows the structure to be both small and microscopically thin, and still be able to produce big, bright, colourful polarised signals.^[49]

Mantis shrimp are long-lived and exhibit complex behaviour, such asritualised fighting, or by the use offluorescent patterns on their bodies for signalling with their own and perhaps even other species. Many have developed complex social behaviours to defend their space from rivals; mantis shrimp are typically solitary sea creatures that may aggressively defend their burrows, either rock formations orself-dug intricateburrows in theseabed. They are rarely seen outside their homes except to feed and relocate. They canlearn and remember well,^{[citation needed]} and are able to recognise neighbouring mantis shrimp with which they frequently interact. They can recognise them by visual signs and even by individual smell.^{[citation needed]}

Mantis shrimp can bediurnal,nocturnal, orcrepuscular (active at twilight), depending on the species. Unlike most crustaceans,^{[clarification needed]} they sometimes hunt, chase, and kill prey. Although some live in temperate seas, most species live in tropical and subtropical waters in theIndian andPacific Oceans, encompassing the seas betweeneastern Africa andHawaii.

Mantis shrimp live in burrows where they spend the majority of their time.^[50] The spearing species build their habitat in softsediments and the smashing species make burrows in hard substrata, such as cavities in coral. These two habitats are crucial for their ecology since they use burrows as sites for retreat and as locations for consuming their prey.^[50] Burrows and coral cavities are also used as sites for mating and for keeping their eggs safe. Stomatopod body size undergoes periodic growth which necessitates finding a new cavity or burrow that will fit the animal's new diameter. Some spearing species can modify their pre-established habitat if the burrow is made of silt or mud, which can be expanded.^[50]

Stomatopods can have as many as 20 or 30breeding episodes over their lifespan. Depending on the species, the eggs are either laid and kept in a burrow, or are carried around under the female's tail until they hatch, as in a number of other crustaceans. Also depending on the species, males and females may come together only to mate, or they may bond inmonogamous, long-term relationships.^[51]

In the monogamous species, the mantis shrimps remain with the same partner up to 20 years. They share the same burrow and may be able to coordinate their activities. Both sexes often take care of the eggs (bi-parental care). InPullosquilla and some species inNannosquilla, the female lays two clutches of eggs – one that the male tends and one that the female tends. In other species, the female looks after the eggs while the male hunts for both of them. After the eggs hatch, the offspring may spend up to three months asplankton.

Although stomatopods typically display the standard types of movement seen intrue shrimp andlobsters, one species,Nannosquilla decemspinosa, has been observed rolling itself into a crude wheel (somewhat resemblingvolvation). The species lives in shallow, sandy areas. At low tides,N. decemspinosa is often stranded by its short rear legs, which are sufficient for movement when the body is supported by water, but not on dry land. The mantis shrimp thus performs a forward flip in an attempt to roll towards the nearesttide pool.N. has been observed to roll repeatedly for 2 m (6.6 ft), but specimens typically travel less than 1 m (3.3 ft).^[52]

Although theDevonianEopteridae have been suggested to be early stomatopods, their fragmentary known remains make the referral uncertain.^[53] The oldest unambiguousstem-group mantis shrimp date to theCarboniferous (359–300 million years ago).^[53][54] Stem-group mantis shrimp are assigned to two major groups thePalaeostomatopodea and theArchaeostomatopodea, the latter of which are more closely related to modern mantis shrimp, which are assigned to the clade Unipeltata.^[53] The oldest members of Unipeltata date to theTriassic.^[54]

• FamilyGonodactylidae
  • Gonodactylus smithii
• FamilyHemisquillidae
  • Hemisquilla ensigera
  • Hemisquilla australiensis
  • Hemisquilla braziliensis
  • Hemisquilla californiensis
• FamilyLysiosquillidae
  • Lysiosquillina maculata, zebra mantis shrimp or striped mantis shrimp
• FamilyNannosquillidae
  • Nannosquilla decemspinosa
  • Platysquilla eusebia
• FamilyOdontodactylidae
  • Odontodactylus scyllarus, peacock mantis shrimp
  • Odontodactylus latirostris, pink-eared mantis shrimp
• FamilyPseudosquillidae
  • Pseudosquilla ciliata, common mantis shrimp
• FamilySquillidae
  • Oratosquilla oratoria (蝦蛄,shako)
  • Rissoides desmaresti
  • Squilla empusa
  • Squilla mantis
• FamilyTetrasquillidae
  • Heterosquilla tricarinata, New Zealand

•  
  Austrosquilla osculans
•  
  Bathysquilla crassispinosa,museum specimen
•  
  Gonodactylus platysoma
•  
  Kasim philippinensis, museum specimen
•  
  Lysiosquillina maculata
•  
  Lysiosquillina maculata outside its burrow
•  
  Pullosquilla thomassini
•  
  Squilla mantis
•  
  Vossquilla kempi

A large number of mantis shrimp species were first scientifically described by onecarcinologist,Raymond B. Manning; the collection of stomatopods he amassed is the largest in the world, covering 90% of the known species whilst 10% are still unknown.^[55]

The mantis shrimp is eaten by a variety of cultures. InJapanese cuisine, the mantis shrimp speciesOratosquilla oratoria, calledshako (蝦蛄), is eaten boiled as asushi topping, and occasionally raw assashimi.

Mantis shrimps are also abundant along Vietnam's coast, known in Vietnamese asbề bề,tôm tích ortôm tít. In regions such as Nha Trang, they are calledbàn chải, named for its resemblance to a scrub brush. The shrimp can be steamed, boiled, grilled, or dried, used withpepper, salt andlime,fish sauce andtamarind, orfennel.^[56]

InCantonese cuisine, the mantis shrimp is known as "urinating shrimp" (Chinese:瀨尿蝦;pinyin:lài niào xiā;Jyutping:laai6 niu6 haa1) because of their tendency to shoot a jet of water when picked up. After cooking, their flesh is closer to that oflobsters than that ofshrimp, and like lobsters, their shells are quite hard and require some pressure to crack. One common preparation is first deep-frying, then stir-frying with garlic and chili peppers. They may also be boiled or steamed.^{[citation needed]}

In theMediterranean countries, the mantis shrimpSquilla mantis is a common seafood, especially on theAdriatic coasts (canocchia) and theGulf of Cádiz (galera).^{[citation needed]}

In thePhilippines, the mantis shrimp is known astatampal, hipong-dapa,pitik-pitik, oralupihang-dagat, and is cooked and eaten like any other shrimp.^{[citation needed]}

InKiribati, mantis shrimp calledte waro inGilbertese are abundant and are eaten boiled.InHawaii, some mantis shrimp have grown unusually large in the contaminated water of the GrandAla Wai Canal inWaikiki. The dangers normally associated with consuming seafood caught in contaminated waters are present in these mantis shrimp.^[7]

Some saltwater aquarists keep stomatopods in captivity.^[57] Thepeacock mantis is especially colourful and desired in the trade.

While some aquarists value mantis shrimps, others consider them harmful pests, because they are voracious predators, eating other desirable inhabitants of the tank. Additionally, some rock-burrowing species can do more damage tolive rock than the fishkeeper would prefer.

The live rock with mantis shrimp burrows is considered useful by some in themarine aquarium trade and is often collected. A piece of live rock not uncommonly conveys a live mantis shrimp into an aquarium. Once inside the tank, it may feed on fish and other inhabitants, and is notoriously difficult to catch when established in a well-stocked tank.^[58] While there are accounts of this shrimp breaking glass tanks, they are rare and are usually the result of the shrimp being kept in too small a tank. While stomatopods do not eat coral, smashers can damage it if they try to make a home within it.^[59]

•  Crustaceans portal

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• The Lurker's Guide to Stomatopods—mantis shrimp
• Mantis shrimp—colourful and aggressive
• Research on Stomatopods at the University of MarylandArchived 2010-05-30 at theWayback Machine
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• TED talk
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