mantis shrimp

File:MantisShrimpLyd.jpg. The folded raptorial claws are flanking the carapace]]

Mantis shrimp typically grow to around {{cvt|10|cm}} in length, while a few species such as the zebra mantis shrimp can reach up to {{cvt|38|cm}}.{{cite news |url=http://the.honoluluadvertiser.com/article/2003/Feb/14/ln/ln01a.html |author=James Gonser |work=The Honolulu Advertiser |date=February 15, 2003 |title=Large shrimp thriving in Ala Wai Canal muck |access-date=July 20, 2006 |archive-date=November 11, 2020 |archive-url=https://web.archive.org/web/20201111193014/http://the.honoluluadvertiser.com/article/2003/Feb/14/ln/ln01a.html |url-status=live }} A mantis shrimp's carapace covers only the rear part of the head and the first four segments of the thorax. 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.{{cite web | url=https://sites.nd.edu/biomechanics-in-the-wild/2019/03/05/how-the-mantis-shrimp-packs-its-punch/#:~:text=The%20mantis%20shrimp%2C%20a%20six%20inch%20long%20crustacean,around%201500%20newtons%20of%20force%20with%20each%20blow | title=How the Mantis Shrimp Packs its Punch | Biomechanics in the Wild }} 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){{Cite web |title=Why are Mantis Shrimp so Awesome? |url=https://www.calacademy.org/explore-science/why-are-mantis-shrimp-so-awesome |access-date=2022-07-21 |website=California Academy of Sciences |language=en |archive-date=2022-08-10 |archive-url=https://web.archive.org/web/20220810083818/https://calacademy.org/explore-science/why-are-mantis-shrimp-so-awesome |url-status=live }} 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 of praying mantids, hence the common name of these crustaceans. This is the type found in most mantis shrimp.{{cite journal |last1=Anderson |first1=Philip S.L. |last2=Claverie |first2=Thomas |last3=Patek |first3=S.N. |date=2014-07-01 |title=Levers and linkages: mechanical trade-offs in a power-amplified system |url=https://academic.oup.com/evolut/article/68/7/1919/6852628 |journal=Evolution |volume=68 |issue=7 |pages=1919–1933 |doi=10.1111/evo.12407 |pmid=24635148 |access-date=2025-01-08}}
• 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.
• 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.{{Cite web |title=h_californiensis |url=https://ucmp.berkeley.edu/arthropoda/crustacea/malacostraca/eumalacostraca/royslist/species.php?name=h_californiensis |access-date=2022-07-21 |website=ucmp.berkeley.edu |archive-date=2023-04-18 |archive-url=https://web.archive.org/web/20230418023755/https://ucmp.berkeley.edu/arthropoda/crustacea/malacostraca/eumalacostraca/royslist/species.php?name=h_californiensis |url-status=live }}
• Hatchet: An unusual, highly derived appendage that only a few species have. This body plan is largely unresearched.{{Cite web |title=How mantis shrimp evolved many shapes with same powerful punch |url=https://phys.org/news/2015-02-mantis-shrimp-evolved-powerful.html |access-date=2022-07-21 |website=phys.org |language=en |archive-date=2022-07-21 |archive-url=https://web.archive.org/web/20220721161949/https://phys.org/news/2015-02-mantis-shrimp-evolved-powerful.html |url-status=live }}{{Cite web |title=Roy's List of Stomatopods for the Aquarium |url=https://ucmp.berkeley.edu/arthropoda/crustacea/malacostraca/eumalacostraca/royslist/ |access-date=2022-07-21 |website=ucmp.berkeley.edu |archive-date=2022-08-23 |archive-url=https://web.archive.org/web/20220823173035/https://ucmp.berkeley.edu/arthropoda/crustacea/malacostraca/eumalacostraca/royslist/ |url-status=live }}{{Cite web |title=a_derijardi |url=https://ucmp.berkeley.edu/arthropoda/crustacea/malacostraca/eumalacostraca/royslist/species.php?name=a_derijardi |access-date=2022-07-21 |website=ucmp.berkeley.edu |archive-date=2022-01-31 |archive-url=https://web.archive.org/web/20220131024528/https://ucmp.berkeley.edu/arthropoda/crustacea/malacostraca/eumalacostraca/royslist/species.php?name=a_derijardi |url-status=live }}

File:20220123 stomatopod strike mechanics spearing en.gif (raptorial claw, ballistic claw) of mantis shrimp]]

Both types strike by rapidly unfolding and swinging their raptorial 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 {{cvt|23|m/s|km/h mph|lk=on}} from a standing start.{{cite journal |author=S. N. Patek, W. L. Korff & R. L. Caldwell |year=2004 |journal=Nature |volume=428 |pages=819–820 |title=Deadly strike mechanism of a mantis shrimp |doi=10.1038/428819a |pmid=15103366 |issue=6985 |bibcode=2004Natur.428..819P |s2cid=4324997 |url=https://pateklab.biology.duke.edu/sites/pateklab.biology.duke.edu/files/Pateketal2004Nature.pdf |access-date=2017-05-02 |archive-date=2021-01-26 |archive-url=https://web.archive.org/web/20210126130108/https://pateklab.biology.duke.edu/sites/pateklab.biology.duke.edu/files/Pateketal2004Nature.pdf |url-status=dead}} Because they strike so rapidly, they generate vapor-filled bubbles in the water between the appendage and the striking surface—known as cavitation bubbles. 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.{{cite journal |author1=S. N. Patek |author2=R. L. Caldwell |name-list-style=amp |year=2005 |journal=Journal of Experimental Biology |volume=208 |pages=3655–3664 |title=Extreme impact and cavitation forces of a biological hammer: strike forces of the peacock mantis shrimp |doi=10.1242/jeb.01831 |pmid=16169943 |issue=19 |doi-access=free}} Even if the initial strike misses the prey, the resulting shock wave can be enough to stun or kill.

Smashers use this ability to attack crabs, snails, rock oysters, and other molluscs, their blunt clubs enabling them to crack the shells of their prey into pieces. Spearers, however, prefer the meat of softer animals, such as fish and cephalopods, which their barbed claws can more easily slice and snag.

The appendages are being studied as a microscale analogue for new macroscale material structures.{{cite news |title=Mantis shrimp inspires next generation of ultra-strong materials |url=https://www.spacedaily.com/reports/Mantis_shrimp_inspires_next_generation_of_ultra_strong_materials_999.html |work=Space Daily |date=June 1, 2016 |access-date=May 13, 2020 |archive-date=May 24, 2021 |archive-url=https://web.archive.org/web/20210524171606/https://www.spacedaily.com/reports/Mantis_shrimp_inspires_next_generation_of_ultra_strong_materials_999.html |url-status=live }}{{Clarify|date=January 2025}}

{{Cleanup|section|reason=Section is quite long (partially due to repetitive sections) and its subsections may not be well delineated, and many sections may be too technical/lacking in clarity for the general audience. Might be more appropriate to have separate article on Stomatopod vision|date=January 2025}}

File:Odontodactylus scyllarus eyes.jpgs, indicating the ommatidia that are pointing towards the camera]]

File:Mantis shrimp eyes.jpg eyes]]

The eyes of the mantis shrimp are mounted on mobile stalks 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.{{Cite journal |title=Complex gaze stabilization in mantis shrimp |first1=Ilse M. |last1=Daly |first2=Martin J. |last2=How |first3=Julian C. |last3=Partridge |first4=Nicholas W. |last4=Roberts |date=May 16, 2018 |journal=Proceedings of the Royal Society B: Biological Sciences |volume=285 |issue=1878 |pages=20180594 |doi=10.1098/rspb.2018.0594 |pmid=29720419 |pmc=5966611}} 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.{{cite journal |last1=Cronin |first1=Thomas W. |last2=Bok |first2=Michael J. |last3=Marshall |first3=N. Justin |last4=Caldwell |first4=Roy L. |title=Filtering and polychromatic vision in mantis shrimps: themes in visible and ultraviolet vision |journal=Philosophical Transactions of the Royal Society B: Biological Sciences |date=19 February 2014 |volume=369 |issue=1636 |pages=20130032 |doi=10.1098/rstb.2013.0032 |pmid=24395960 |pmc=3886321}}{{cite web |last=Franklin |first=Amanda M. |title=Mantis shrimp have the world's best eyes – but why? |publisher=The Conversation |date=September 4, 2013 |access-date=July 5, 2018 |url=https://theconversation.com/mantis-shrimp-have-the-worlds-best-eyes-but-why-17577 |archive-date=July 5, 2018 |archive-url=https://web.archive.org/web/20180705204125/https://theconversation.com/amp/mantis-shrimp-have-the-worlds-best-eyes-but-why-17577 |url-status=live }}{{cite journal |last=Milius |first=Susan |title=Mantis shrimp flub color vision test |journal=Science News |year=2012 |volume=182 |issue=6 |pages=11 |jstor=23351000 |doi=10.1002/scin.5591820609}}

Each compound eye is made up of tens of thousands of ommatidia, clusters of photoreceptor cells. 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. 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 three pseudopupils in one eye). In other words, each eye possesses trinocular vision, and therefore depth 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.

Compared with the three types of photoreceptor 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.{{cite journal |last=Cronin |first=Thomas W. |title=Sensory adaptation: Tunable colour vision in a mantis shrimp |journal=Nature |volume=411 |issue=6837 |pages=547–8 |doi=10.1038/35079184 |pmid=11385560 |year=2001 |bibcode=2001Natur.411..547C |s2cid=205017718}} This phenomenon, called "spectral tuning", is species-specific.{{cite journal |title=Evolutionary variation in the expression of phenotypically plastic color vision in Caribbean mantis shrimps, genus Neogonodactylus. |journal=Marine Biology |volume=150 |issue=2 |pages=213–220 |doi=10.1007/s00227-006-0313-5 |year=2006 |last1=Cheroske |first1=Alexander G. |last2=Barber |first2=Paul H. |last3=Cronin |first3=Thomas W. |bibcode=2006MarBi.150..213C |url=http://darchive.mblwhoilibrary.org/bitstream/1912/1391/1/CheroskeetalMB2006reprint.pdf |hdl=1912/1391 |s2cid=40203342 |hdl-access=free |access-date=2019-09-02 |archive-date=2024-01-04 |archive-url=https://web.archive.org/web/20240104013921/https://darchive.mblwhoilibrary.org/server/api/core/bitstreams/cb05f806-6948-5234-a382-fa09c4afb279/content |url-status=live }} Cheroske et al. did not observe spectral tuning in Neogonodactylus oerstedii, the species with the most monotonous natural photic environment. In N. 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 in N. 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 the retinal binding pocket of the opsin.{{cite journal |last1=Porter |first1=Megan L. |last2=Bok |first2=Michael J. |last3=Robinson |first3=Phyllis R. |last4=Cronin |first4=Thomas W. |title=Molecular diversity of visual pigments in Stomatopoda (Crustacea) |journal=Visual Neuroscience |date=1 May 2009 |volume=26 |issue=3 |pages=255–265 |doi=10.1017/S0952523809090129 |pmid=19534844 |s2cid=6516816}}

The huge diversity seen in mantis shrimp photoreceptors likely comes from ancient gene duplication events.{{cite journal |last1=Porter |first1=Megan L. |last2=Speiser |first2=Daniel I. |last3=Zaharoff |first3=Alexander K. |last4=Caldwell |first4=Roy L. |last5=Cronin |first5=Thomas W. |last6=Oakley |first6=Todd H. |year=2013 |title=The Evolution of Complexity in the Visual Systems of Stomatopods: Insights from Transcriptomics. |journal=Integrative and Comparative Biology |volume=53 |issue=1 |pages=39–49 |doi=10.1093/icb/ict060 |pmid=23727979 |doi-access=free}} One consequence of this duplication is the lack of correlation between opsin transcript number and physiologically expressed photoreceptors. 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.{{cite journal |title=Evolution of anatomical and physiological specialisation in the compound eyes of stomatopod crustaceans. |journal=Journal of Experimental Biology |volume=213}}

Mantis shrimp can perceive wavelengths of light ranging from deep ultraviolet (300 nm) to far-red (720 nm) and polarised light.{{cite journal |last1=Thoen |first1=Hanne H. |last2=How |first2=Martin J. |last3=Chiou |first3=Tsyr-Huei |last4=Marshall |first4=Nicholas Justin |date=January 24, 2014 |title=A Different Form of Color Vision in Mantis Shrimp |journal=Science |volume=334 |issue=6169 |pages=411–413 |bibcode=2014Sci...343..411T |doi=10.1126/science.1245824 |pmid=24458639 |s2cid=31784941}} 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 detect circularly or linearly polarised light. Twelve types of photoreceptor cells are in rows 1 to 4, four of which detect ultraviolet light.{{cite journal |last1=Marshall |first1=Nicholas Justin |last2=Oberwinkler |first2=Johannes |title=Ultraviolet vision: the colourful world of the mantis shrimp |journal=Nature |date=October 28, 1999 |volume=401 |pages=873–874 |doi=10.1038/44751 |pmid=10553902 |issue=6756 |bibcode=1999Natur.401..873M |s2cid=4360184}} 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|reason=How much is this?|date=July 2024}} 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. However, some mantis shrimp have been found capable of distinguishing between high-saturation and low-saturation colors.{{Cite journal |last1=Streets |first1=Amy |last2=England |first2=Hayley |last3=Marshall |first3=Justin |date=2022-03-15 |title=Colour vision in stomatopod crustaceans: more questions than answers |journal=Journal of Experimental Biology |language=en |volume=225 |issue=6 |doi=10.1242/jeb.243699 |issn=0022-0949 |pmc=9001920 |pmid=35224643|bibcode=2022JExpB.225B3699S }}

File:Mantis Shrimp at the National Aquarium (Baltimore) - July 2017.jpg]]

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.{{cite journal |author1=Michael Bok |author2=Megan Porter |author3=Allen Place |author4=Thomas Cronin |title=Biological Sunscreens Tune Polychromatic Ultraviolet Vision in Mantis Shrimp |journal=Current Biology |year=2014 |volume=24 |issue=14 |pages=1636–42 |doi=10.1016/j.cub.2014.05.071 |pmid=24998530 |doi-access=free|bibcode=2014CBio...24.1636B }}[http://www.latimes.com/science/sciencenow/la-sci-sn-mantis-shrimp-20140703-story.html Mantis shrimp wear tinted shades to see UV light] {{Webarchive|url=https://web.archive.org/web/20141122181512/http://www.latimes.com/science/sciencenow/la-sci-sn-mantis-shrimp-20140703-story.html |date=2014-11-22 }}. Latimes.com (2014-07-05). Retrieved on 2015-10-21. They are currently believed insensitive to infrared light.{{cite journal |author1=David Cowles |author2=Jaclyn R. Van Dolson |author3=Lisa R. Hainey |author4=Dallas M. Dick |year=2006 |title=The use of different eye regions in the mantis shrimp Hemisquilla californiensis Stephenson, 1967 (Crustacea: Stomatopoda) for detecting objects |journal=Journal of Experimental Marine Biology and Ecology |volume=330 |issue=2 |pages=528–534 |doi=10.1016/j.jembe.2005.09.016|bibcode=2006JEMBE.330..528C }} The optical elements in these rows have eight different classes of visual pigments and the rhabdom (area of eye that absorbs light from a single direction) is divided into three different pigmented 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.{{cite journal |title=The molecular genetics and evolution of colour and polarization vision in stomatopod crustaceans. |journal=Ophthalmic Physiology |volume=30}} 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.

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{{cite web |last=DuRant |first=Hassan |date=3 July 2014 |title=Mantis shrimp use 'nature's sunblock' to see UV |url=https://www.science.org/content/article/mantis-shrimp-use-natures-sunblock-see-uv |work=sciencemag.org |access-date=5 July 2014 |archive-date=25 April 2023 |archive-url=https://web.archive.org/web/20230425094609/https://www.science.org/content/article/mantis-shrimp-use-natures-sunblock-see-uv |url-status=live }}) 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 the retina seems to be processed into numerous parallel data streams leading into the brain, greatly reducing the analytical requirements at higher levels.{{cite journal |last1=Cronin |first1=Thomas W. |last2=Marshall |first2=Justin |title=Parallel processing and image analysis in the eyes of mantis shrimps |journal=The Biological Bulletin |volume=200 |issue=2 |pages=177–183 |year=2001 |pmid=11341580 |doi=10.2307/1543312 |jstor=1543312 |s2cid=12381929 |url=https://www.biodiversitylibrary.org/part/11007 |access-date=2021-05-24 |archive-date=2020-06-19 |archive-url=https://web.archive.org/web/20200619133225/https://www.biodiversitylibrary.org/part/11007 |url-status=live }}

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|date=July 2024}}

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.{{cite journal |last1=Chiou |first1=Tsyr-Huei |last2=Kleinlogel |first2=Sanja |last3=Cronin |first3=Tom |last4=Caldwell |first4=Roy |last5=Loeffler |first5=Birte |last6=Siddiqi |first6=Afsheen |last7=Goldzien |first7=Alan |last8=Marshall |first8=Justin |title=Circular polarization vision in a stomatopod crustacean |journal=Current Biology |volume=18 |issue=6 |pages=429–434 |date=March 25, 2008 |doi=10.1016/j.cub.2008.02.066 |pmid=18356053 |s2cid=6925705 |doi-access=free|bibcode=2008CBio...18..429C }}{{cite journal |last1=Kleinlogel |first1=Sonja |last2=White |first2=Andrew |title=The secret world of shrimps: polarisation vision at its best |journal=PLoS ONE |volume=3 |issue=5 |pages=e2190 |year=2009 |doi=10.1371/journal.pone.0002190 |pmid=18478095 |pmc=2377063 |bibcode=2008PLoSO...3.2190K |arxiv=0804.2162 |doi-access=free}}{{cite journal |last1=Templin |first1=Rachel M. |last2=How |first2=Martin J. |last3=Roberts |first3=Nicholas W. |last4=Chiou |first4=Tsyr-Huei |last5=Marshall |first5=Justin |title=Circularly polarized light detection in stomatopod crustaceans: a comparison of photoreceptors and possible function in six species |journal=The Journal of Experimental Biology |date=15 September 2017 |volume=220 |issue=18 |pages=3222–3230 |doi=10.1242/jeb.162941 |pmid=28667244 |doi-access=free|hdl=1983/1f1c982f-9a88-4184-b59a-2cebd73ec818 |hdl-access=free }} They perform this feat by converting circularly polarized light into linearly polarized light via quarter-waveplates formed from stacks of microvilli. 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 century Blu-ray Disc technology.{{cite journal |title=A biological quarter-wave retarder with excellent achromaticity in the visible wavelength region |last1=Roberts |first1=Nicholas W. |last2=Chiou |first2=Tsyr-Huei |last3=Marshall |first3=Nicholas Justin |last4=Cronin |first4=Thomas W. |journal=Nature Photonics |volume=3 |issue=11 |pages=641–644 |year=2009 |doi=10.1038/nphoton.2009.189 |bibcode=2009NaPho...3..641R}}{{cite web |last=Lee |first=Chris |title=A crustacean eye that rivals the best optical equipment |publisher=Ars Technica |work=Nobel Intent |date=November 1, 2009 |url=https://arstechnica.com/science/news/2009/11/a-crusty-eye-sees-curly-light.ars |access-date=June 14, 2017 |archive-date=April 5, 2012 |archive-url=https://web.archive.org/web/20120405130942/http://arstechnica.com/science/news/2009/11/a-crusty-eye-sees-curly-light.ars |url-status=live }}

The species Gonodactylus smithii is the only organism known to simultaneously detect the four linear and two circular polarisation components required to measure all four Stokes parameters, which yield a full description of polarisation. It is thus believed to have optimal polarisation vision.{{cite web |last=Minard |first=Anne |title="Weird beastie" shrimp have super-vision |publisher=National Geographic Society |date=May 19, 2008 |url=http://news.nationalgeographic.com/news/2008/05/080519-shrimp-colors.html |archive-url=https://web.archive.org/web/20080527213305/http://news.nationalgeographic.com/news/2008/05/080519-shrimp-colors.html |url-status=dead |archive-date=May 27, 2008}} 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.{{Cite journal |last1=Daly |first1=Ilse M. |last2=How |first2=Martin J. |last3=Partridge |first3=Julian C. |last4=Roberts |first4=Nicholas W. |date=2018-05-16 |title=Complex gaze stabilization in mantis shrimp |journal=Proceedings of the Royal Society B: Biological Sciences |volume=285 |issue=1878 |pages=20180594 |doi=10.1098/rspb.2018.0594 |pmc=5966611 |pmid=29720419}} Since each eye moves independently from the other, it creates two separate streams of visual information.{{cite web |url=http://www.newsweek.com/mantis-shrimp-have-perfected-eye-roll-better-see-things-we-cant-even-imagine-480162 |title=Mantis shrimp have perfected the eye roll to see things we can't imagine |website=Newsweek |date=14 July 2016 |access-date=6 February 2017 |archive-date=6 February 2017 |archive-url=https://web.archive.org/web/20170206210222/http://www.newsweek.com/mantis-shrimp-have-perfected-eye-roll-better-see-things-we-cant-even-imagine-480162 |url-status=live }}

File:Pseudosquilla.JPG]]

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.{{cite journal |last1=How |first1=M. J. |last2=Porter |first2=M. L. |last3=Radford |first3=A. N. |last4=Feller |first4=K. D. |last5=Temple |first5=S. E. |last6=Caldwell |first6=R. L. |last7=Marshall |first7=N. J. |last8=Cronin |first8=T. W. |last9=Roberts |first9=N. W. |title=Out of the blue: the evolution of horizontally polarized signals in Haptosquilla (Crustacea, Stomatopoda, Protosquillidae) |journal=Journal of Experimental Biology |date=7 August 2014 |volume=217 |issue=19 |pages=3425–3431 |doi=10.1242/jeb.107581 |pmid=25104760 |doi-access=free|hdl=11603/13393 |hdl-access=free }} This mechanism could provide an evolutionary advantage; it only requires small changes to the cell in the eye and could easily lead to natural selection.{{cite press release |title=Mantis shrimps could show us the way to a better DVD |publisher=University of Bristol |date=25 October 2009 |url=http://www.bristol.ac.uk/news/2009/6591.html |access-date=May 13, 2020 |archive-date=31 October 2020 |archive-url=https://web.archive.org/web/20201031051957/http://www.bristol.ac.uk/news/2009/6591.html |url-status=live }}

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 as barracuda, 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.

During mating rituals, mantis shrimps actively fluoresce, and the wavelength of this fluorescence matches the wavelengths detected by their eye pigments.{{cite journal |author1=C. H. Mazel |author2=T. W. Cronin |author3=R. L. Caldwell |author4=N. J. Marshall |year=2004 |title=Fluorescent enhancement of signaling in a mantis shrimp |journal=Science |volume=303 |issue=5654 |page=51 |doi=10.1126/science.1089803 |pmid=14615546 |s2cid=35009047}} Females are only fertile during certain phases of the tidal cycle; the ability to perceive the phase 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|date=December 2024}}

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 of opponent 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.{{cite journal |last1=Morrison |first1=Jessica |title=Mantis shrimp's super colour vision debunked |journal=Nature |date=23 January 2014 |doi=10.1038/nature.2014.14578 |s2cid=191386729}} 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.{{cite web |last1=Macknik |first1=Stephen L. |title=Parallels between Shrimp and Human Color Vision |url=https://blogs.scientificamerican.com/illusion-chasers/parallels-between-shrimp-and-human-color-vision/ |website=Scientific American Blog Network |date=March 20, 2014 |access-date=May 13, 2020 |archive-date=May 25, 2020 |archive-url=https://web.archive.org/web/20200525140407/https://blogs.scientificamerican.com/illusion-chasers/parallels-between-shrimp-and-human-color-vision/ |url-status=live }}

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

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.[http://www.bristol.ac.uk/news/2016/february/mantis-shrimp.html New type of optical material discovered in the secret language of the mantis shrimp] {{Webarchive|url=https://web.archive.org/web/20160307200843/http://bristol.ac.uk/news/2016/february/mantis-shrimp.html |date=2016-03-07 }}. Bristol University (17 February 2016)

File:Careful now... (29866247452).jpg at Wakatobi National Park Sulawesi, partially out of its burrow]]

Mantis shrimp are long-lived and exhibit complex behaviour, such as ritualised fighting, or by the use of fluorescent 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 or self-dug intricate burrows in the seabed. They are rarely seen outside their homes except to feed and relocate. They can learn and remember well,{{cn|date=January 2025}} 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|date=January 2025}}

Mantis shrimp can be diurnal, nocturnal, or crepuscular (active at twilight), depending on the species. Unlike most crustaceans,{{Clarify|reason=Such as? Is pursuit predation really that rare in aquatic crustaceans?|date=January 2025}} they sometimes hunt, chase, and kill prey. Although some live in temperate seas, most species live in tropical and subtropical waters in the Indian and Pacific Oceans, encompassing the seas between eastern Africa and Hawaii.

Mantis shrimp live in burrows where they spend the majority of their time.{{Cite book |last1=Mead |last2=Caldwell |first1=K. |first2=R. |year=2001 |chapter=Mantis Shrimp: Olfactory Apparatus and Chemosensory Behavior |page=219 |editor=Breithaupt, T. |editor2=Thiel, M. |title=Chemical Communication in Crustaceans |publisher=Springer |location=Chile |isbn=9780387771014}} The spearing species build their habitat in soft sediments 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. 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.

File:Stomatopoda (10.1590-2358-2936e2018005) Figure 4.jpg

Stomatopods can have as many as 20 or 30 breeding 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 in monogamous, long-term relationships.{{cite web |title=Sharing the job: monogamy and parental care |url=http://www.ucmp.berkeley.edu/aquarius/monogamy.html |publisher=University of California, Berkeley |access-date=2009-11-13 |archive-date=2009-09-14 |archive-url=https://web.archive.org/web/20090914125517/http://www.ucmp.berkeley.edu/aquarius/monogamy.html |url-status=live }}

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). In Pullosquilla and some species in Nannosquilla, 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 as plankton.

Although stomatopods typically display the standard types of movement seen in true shrimp and lobsters, one species, Nannosquilla decemspinosa, has been observed rolling itself into a crude wheel (somewhat resembling volvation). 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 nearest tide pool. N. has been observed to roll repeatedly for {{cvt|2|m|ft}}, but specimens typically travel less than {{cvt|1|m|ft}}.{{cite journal |author=Caldwell, Roy L. |journal=Nature |volume=282 |pages=71–73 |year=1979 |title=A unique form of locomotion in a stomatopod – backward somersaulting |doi=10.1038/282071a0 |issue=5734 |bibcode=1979Natur.282...71C |s2cid=4311328}}

file:Daidal.png, a primitive Carboniferous mantis shrimp]]

Although the Devonian Eopteridae have been suggested to be early stomatopods, their fragmentary known remains make the referral uncertain. The oldest unambiguous stem-group mantis shrimp date to the Carboniferous (359–300 million years ago). Stem-group mantis shrimp are assigned to two major groups the Palaeostomatopodea and the Archaeostomatopodea, the latter of which are more closely related to modern mantis shrimp, which are assigned to the clade Unipeltata.{{Cite journal |last1=Van Der Wal |first1=Cara |last2=Ahyong |first2=Shane T. |last3=Ho |first3=Simon Y.W. |last4=Lo |first4=Nathan |date=2017-09-21 |title=The evolutionary history of Stomatopoda (Crustacea: Malacostraca) inferred from molecular data |journal=PeerJ |language=en |volume=5 |pages=e3844 |doi=10.7717/peerj.3844 |issn=2167-8359 |pmc=5610894 |pmid=28948111 |doi-access=free}} The oldest members of Unipeltata date to the Triassic.{{Cite journal |last1=Smith |first1=C.P.A. |last2=Aubier |first2=P. |last3=Charbonnier |first3=S. |last4=Laville |first4=T. |last5=Olivier |first5=N. |last6=Escarguel |first6=G. |last7=Jenks |first7=J.F. |last8=Bylund |first8=K.G. |last9=Fara |first9=E. |last10=Brayard |first10=A. |date=2023-03-31 |title=Closing a major gap in mantis shrimp evolution - first fossils of Stomatopoda from the Triassic |url=http://www.geology.cz/bulletin/contents/art1864 |journal=Bulletin of Geosciences |language=en |pages=95–110 |doi=10.3140/bull.geosci.1864 |issn=1802-8225 |doi-access=free |access-date=2023-05-23 |archive-date=2023-06-06 |archive-url=https://web.archive.org/web/20230606082726/http://www.geology.cz/bulletin/contents/art1864 |url-status=live }}

• Family Gonodactylidae
• Gonodactylus smithii
• Family Hemisquillidae
• Hemisquilla ensigera
• Hemisquilla australiensis
• Hemisquilla braziliensis
• Hemisquilla californiensis
• Family Lysiosquillidae
• Lysiosquillina maculata, zebra mantis shrimp or striped mantis shrimp
• Family Nannosquillidae
• Nannosquilla decemspinosa
• Platysquilla eusebia
• Family Odontodactylidae
• Odontodactylus scyllarus, peacock mantis shrimp
• Odontodactylus latirostris, pink-eared mantis shrimp
• Family Pseudosquillidae
• Pseudosquilla ciliata, common mantis shrimp
• Family Squillidae
• {{nihongo|Oratosquilla oratoria|蝦蛄|shako}}
• Rissoides desmaresti
• Squilla empusa
• Squilla mantis
• Family Tetrasquillidae
• Heterosquilla tricarinata, New Zealand

File:Austrosquilla osculans 61399682.jpg|Austrosquilla osculans

File:Specimen of Bathysquilla crassispinosa.JPG|Bathysquilla crassispinosa, museum specimen

File:Gonodactylus platysoma.jpg|Gonodactylus platysoma

File:Kasim philippinensis (MNHN-IU-2014-23089) 001.jpeg|Kasim philippinensis, museum specimen

File:Mantis shrimp.jpg|Lysiosquillina maculata

File:Mantis Shrimp Sole and Eel - Lysiosquillina maculata (cropped).jpg|Lysiosquillina maculata outside its burrow

File:Pullosquilla thomassini (10.3897-zookeys.721.20588) Figure 4.jpg|Pullosquilla thomassini

File:Squilla mantis.jpg|Squilla mantis

File:Vossquilla kempi (10.1590-2358-2936e2017012) Figure 3 (cropped).jpg|Vossquilla kempi

A large number of mantis shrimp species were first scientifically described by one carcinologist, 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.{{cite journal |title=Raymond B. Manning: an appreciation |author1=Paul F. Clark |author2=Frederick R. Schram |name-list-style=amp |journal=Journal of Crustacean Biology |volume=29 |issue=4 |pages=431–457 |year=2009 |doi=10.1651/09-3158.1 |s2cid=85803151 |doi-access=|bibcode=2009JCBio..29..431C }}

{{missing information|more specifics on species, better if combined with some list of fisheries. For example, Harpiosquilla harpax and Oratosquilla anomala are commonly mentioned in SE Asian contexts|date=October 2021}}

File:Hai san Hau Loc05.JPG, Thanh Hóa, Vietnam]]

The mantis shrimp is eaten by a variety of cultures. In Japanese cuisine, the mantis shrimp species Oratosquilla oratoria, called {{nihongo|shako|蝦蛄}}, is eaten boiled as a sushi topping, and occasionally raw as sashimi.

Mantis shrimps are also abundant along Vietnam's coast, known in Vietnamese as bề bề, tôm tích or tôm tít. In regions such as Nha Trang, they are called bàn chải, named for its resemblance to a scrub brush. The shrimp can be steamed, boiled, grilled, or dried, used with pepper, salt and lime, fish sauce and tamarind, or fennel.{{cite web |url=http://www.dinhduong.com.vn/story/tom-tit-ac-san-mien-song-nuoc |title=Tôm tít – Đặc sản miền sông nước |date=October 1, 2009 |access-date=January 8, 2011 |language=vi |publisher=Dinh dưỡng |archive-date=August 16, 2012 |archive-url=https://web.archive.org/web/20120816004130/http://dinhduong.com.vn/story/tom-tit-ac-san-mien-song-nuoc |url-status=dead}}

File:Banhkhotomtit.JPG, Việt Nam]]

In Cantonese cuisine, the mantis shrimp is known as "urinating shrimp" ({{zh|c=瀨尿蝦|p=lài niào xiā|j=laai6 niu6 haa1}}) because of their tendency to shoot a jet of water when picked up. After cooking, their flesh is closer to that of lobsters than that of shrimp, 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|date=July 2024}}

In the Mediterranean countries, the mantis shrimp Squilla mantis is a common seafood, especially on the Adriatic coasts (canocchia) and the Gulf of Cádiz (galera).{{Citation needed|date=July 2024}}

In the Philippines, the mantis shrimp is known as tatampal, hipong-dapa, pitik-pitik, or alupihang-dagat, and is cooked and eaten like any other shrimp.{{Citation needed|date=July 2024}}

In Kiribati, mantis shrimp called te waro in Gilbertese are abundant and are eaten boiled.

In Hawaii, some mantis shrimp have grown unusually large in the contaminated water of the Grand Ala Wai Canal in Waikiki. The dangers normally associated with consuming seafood caught in contaminated waters are present in these mantis shrimp.

File:Harpiosquilla harpax SeaDonuts.jpg in aquaria]]

Some saltwater aquarists keep stomatopods in captivity.[http://reefkeeping.com/issues/2004-03/jf/feature/index.php A Load of Learnin' About Mantis Shrimps] {{Webarchive|url=https://web.archive.org/web/20110715170218/http://reefkeeping.com/issues/2004-03/jf/feature/index.php |date=2011-07-15 }}, by James Fatherree, in [http://www.reefkeeping.com/joomla/index.php/reefkeeping-blog ReefKeeping] {{Webarchive|url=https://web.archive.org/web/20140219203654/http://reefkeeping.com/joomla/index.php/reefkeeping-blog |date=2014-02-19 }} online magazine. The peacock 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 to live rock than the fishkeeper would prefer.

The live rock with mantis shrimp burrows is considered useful by some in the marine 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.{{cite book |author=Nick Dakin |year=2004 |title=The Marine Aquarium |publisher=London: Andromeda |isbn=978-1-902389-67-7}} 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.{{cite news |url=https://www.usatoday.com/tech/columnist/aprilholladay/2006-01-09-shrimp_x.htm |work=USA Today |date=September 1, 2006 |title=Shrimp spring into shattering action |author=April Holladay |access-date=2017-09-04 |archive-date=2012-06-30 |archive-url=https://web.archive.org/web/20120630105918/http://www.usatoday.com/tech/columnist/aprilholladay/2006-01-09-shrimp_x.htm |url-status=live }}

• {{Portal-inline|Crustaceans}}

{{Reflist|30em}}

{{Commons}}

• {{cite web | last1=Rayne | first1=Elizabeth | title=The seemingly indestructible fists of the mantis shrimp can take a punch | website=Ars Technica | date=February 22, 2025 | url=https://arstechnica.com/science/2025/02/the-seemingly-indestructible-fists-of-the-mantis-shrimp-can-take-a-punch/ | access-date=2025-02-23}}
• [https://web.archive.org/web/20081207035945/http://www.tafi.org.au/zooplankton/imagekey/malacostraca/hoplocarida/ Hoplocarida: Stomatopoda fact sheet—Guide to the marine zooplankton of south eastern Australia]
• [https://www.keyapa.com/mantis/ The Lurker's Guide to Stomatopods—mantis shrimp]
• [https://australian.museum/learn/animals/crustaceans/mantis-shrimp/ Mantis shrimp—colourful and aggressive]
• [http://www.life.umd.edu/biology/faculty/mreaka/index.html Research on Stomatopods at the University of Maryland] {{Webarchive |url=https://web.archive.org/web/20100530020152/http://www.life.umd.edu/biology/faculty/mreaka/index.html |date=2010-05-30 }}
• {{cite web |url=http://www.theaquariumwiki.com/Mantis_shrimp |title=Mantis shrimp |publisher=The Aquarium Wiki |date=April 7, 2006 |access-date=October 21, 2006 |archive-date=September 30, 2011 |archive-url=https://web.archive.org/web/20110930222515/http://www.theaquariumwiki.com/Mantis_shrimp |url-status=dead }}
• [http://ib.berkeley.edu/labs/caldwell/ Caldwell Lab at the University of California, Berkeley]
• [https://web.archive.org/web/20100522170927/http://www2.bio.umass.edu/biology/pateklab/home Patek Lab at the University of Massachusetts, Amherst]
• {{cite web |url=http://scienceblogs.com/pharyngula/2008/05/the_superior_eyes_of_shrimp.php |title=The superior eyes of shrimp |work=Pharyngula |date=May 24, 2008 |last1=Myers |first1=P. Z. |author-link=PZ Myers |url-status=dead |archive-url=https://web.archive.org/web/20090912115946/http://scienceblogs.com/pharyngula/2008/05/the_superior_eyes_of_shrimp.php |archive-date=September 12, 2009}}
• {{cite web |url=http://blogs.nature.com/news/2008/05/shrimps_super_sight.html |first1=Daniel |last1=Cressey |title=Shrimp's super sight |work=The Great Beyond |date=May 14, 2008 |publisher=nature.com |access-date=May 13, 2020 |archive-date=September 22, 2020 |archive-url=https://web.archive.org/web/20200922091231/http://blogs.nature.com/news/2008/05/shrimps_super_sight.html |url-status=dead}}
• [http://www.danapointfishcompany.com/mantis-shrimp/ Dana Point Fish Company—Top and Bottom Views of Mantis Shrimp]
• [http://www.ted.com/talks/sheila_patek_clocks_the_fastest_animals TED talk]
• [https://www.youtube.com/watch?v=Lm1ChtK9QDU Deep Look (PBS)]

{{Malacostraca}}

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