|Superfamilies and families |
Mantis shrimp or stomatopods are marine crustaceans, the members of the order Stomatopoda. They commonly reach 30 centimetres (12 in) in length, though in exceptional cases have been recorded at up to 38 cm (15 in). The largest ever caught has a length of 46 cm (18 in) in the ocean near Fort Pierce, Florida of USA. The carapace of mantis shrimp covers only the rear part of the head and the first four segments of the thorax. There are more than 400 species of Mantis shrimp. They come in a variety of colours, from shades of brown to bright neon colours and are among the most important predators in many shallow, tropical and sub-tropical marine habitats. Despite being common, they are poorly understood as many species spend most of their life tucked away in burrows and holes.
Called "sea locusts" by ancient Assyrians, "prawn killers" in Australia and now sometimes referred to as "thumb splitters" – because of the animal's ability to inflict painful gashes if handled incautiously – mantis shrimp sport powerful claws that they use to attack and kill prey by spearing, stunning, or dismemberment. In captivity, some larger species are capable of breaking through aquarium glass with a single strike.
Around 400 species of mantis shrimp have currently been described worldwide; all living species are in the suborder Unipeltata.
These aggressive and typically solitary sea creatures spend most of their time hiding in rock formations or burrowing intricate passageways in the sea bed. They rarely exit their homes except to feed and relocate, and can be diurnal, nocturnal, or crepuscular, depending on the species. They either wait for prey to chance upon them or, unlike most crustaceans, sometimes hunt, chase, and kill prey. Although some live in temperate seas, most species live in tropical and subtropical seas in the Indian and Pacific Oceans between eastern Africa and Hawaii.
They are commonly separated into two distinct groups determined by the manner of claws they possess:
- Spearers are armed with spiny appendages topped with barbed tips, used to stab and snag prey.
- Smashers, on the other hand, 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 meals apart. The inner aspect of the dactyl (the terminal portion of the appendage) can also possess a sharp edge, with which the animal can cut prey while it swims.
Both types strike by rapidly unfolding and swinging their raptorial claws at the prey, and are capable of inflicting 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 from a standing start. Because they strike so rapidly, they generate cavitation bubbles between the appendage and the striking surface. 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. Even if the initial strike misses the prey, the resulting shock wave can be enough to stun or kill the prey.
The snap can also produce sonoluminescence from the collapsing bubble. This will produce a very small amount of light and high temperatures in the range of several thousand kelvins within the collapsing bubble, although both the light and high temperatures are too weak and short-lived to be detected without advanced scientific equipment. The light emission and temperature increase probably have no biological significance but are rather side-effects of the rapid snapping motion. Pistol shrimp produce this effect in a very similar manner.
Smashers use this ability to attack snails, crabs, molluscs and rock oysters, their blunt clubs enabling them to crack the shells of their prey into pieces. Spearers, on the other hand, prefer the meat of softer animals, like fish, which their barbed claws can more easily slice and snag.
The mantis shrimp has one of the most elaborate visual systems ever discovered.
The midband region of its eye is made up of six rows of specialised ommatidia. Four rows carry up to 16 different photoreceptor pigments, 12 for colour sensitivity, others for colour filtering. The vision of the mantis shrimp is so precise that it can perceive both polarised light and multispectral images. Their eyes (both mounted on mobile stalks and capable of moving independently of each other) are similarly variably coloured and are considered to be the most complex eyes in the animal kingdom.
Each compound eye is made up of up to ten thousand ommatidia of the apposition type. Each eye consists of two flattened hemispheres separated by six parallel rows of specialised ommatidia, collectively called the midband, which divides the eye into three regions. This configuration enables mantis shrimp to see objects with three parts of the same eye. In other words, each eye possesses trinocular vision and depth perception. The upper and lower hemispheres are used primarily for recognition of form and motion, like the eyes of many other crustaceans.
Rows 1–4 of the midband are specialised for colour vision, from ultra-violet to longer wavelengths. Their UV-vision can detect five different frequencies in the deep ultraviolet. To do this they use two photoreceptors in combination with four different colour filters. They are not currently believed to be sensitive to infrared light. The optical elements in these rows have eight different classes of visual pigments and the rhabdom 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. It is organised like a sandwich; a tier, a colour filter of one class, a tier again, a colour filter of another class, and then a last tier. Rows 5–6 are also segregated into different tiers, but have only one class of visual pigment (a ninth class) and are specialised for polarisation vision. They can detect different planes of polarised light. A tenth class of visual pigment is found in the dorsal and ventral hemispheres of the eye.
The midband only covers about 5°–10° of the visual field at any given instant, but like most crustaceans, mantis shrimps have their the eyes mounted on stalks. In mantis shrimps the movement of the stalked eye is unusually free, and can be driven in all possible axes of movement – up to at least 70° – by eight individual 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 hemisphere of the eye. They can also track moving objects using large, rapid eye movements where the two eyes move independently. By combining different techniques, including saccadic movements, the midband can cover a very wide range of the visual field.
Some species have at least 16 different photoreceptor types, which are divided into four classes (their spectral sensitivity is further tuned by colour filters in the retinas), 12 of them for colour analysis in the different wavelengths (including six which are sensitive to ultraviolet light) and four of them for analysing polarised light. By comparison, most humans have only four visual pigments, of which three are dedicated to see colour, and the human lenses block ultraviolet light. The visual information leaving the retina seems to be processed into numerous parallel data streams leading into the central nervous system, greatly reducing the analytical requirements at higher levels.
At least two species have been reported to be able to detect circularly polarised light. Some of their biological quarter-wave plates perform more uniformly over the visual spectrum than any current man-made polarizing optics, and it has been speculated that this could inspire a new type of optical media that would outperform the current generation of Blu-ray disc technology.
The species Gonodactylus smithii is the only organism known to simultaneously detect the four linear and two circular polarization components required to measure all four Stokes parameters, which yield a full description of polarization. It is thus believed to have optimal polarization vision.
Suggested advantages of visual system
What advantage sensitivity to polarization confers is unclear; however polarization vision is used by other animals for sexual signalling and secret communication that avoids the attention of predators. This mechanism could provide an evolutionary advantage; it only requires small changes to the cell in the eye and would be easily selected for.
The eyes of mantis shrimp may enable them to recognize different types of coral, prey species (which are often transparent or semi-transparent), or predators, such as barracuda, which have shimmering scales. Alternatively, the manner in which mantis shrimp hunt (very rapid movements of the claws) may require very accurate ranging information, which would require accurate depth perception.
During mating rituals, mantis shrimp actively fluoresce, and the wavelength of this fluorescence matches the wavelengths detected by their eye pigments. 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 mantis shrimp information about the size of the tide, which is important to species living in shallow water near the shore.
It has been suggested that the capacity to see UV light enables observation of otherwise hard to detect prey on coral reefs.
Research also shows their visual experience of colours are not that different from humans. The eyes are actually a mechanism that operates at the level of individual cones and makes the brain more efficient. This system allows the visual information to be preprocessed by the eyes instead of the brain, which would otherwise have to be larger to deal with the stream of raw data and thus require more time and energy. While the eyes themselves are complex and not yet fully understood, the principle of the system appears to be simple. It shows great similarities with that of humans, but works the other way around. In the human brain, and area called the inferior temporal cortex have a huge amount of colour-specific neurons, processing visual impulses from the eyes to create colourful experiences. The mantis shrimp use a similar strategy, except that it is the different types of photoreceptors in its eyes that perform the same function as the human brain neurons, and is thus a more hardwired and efficient system for an animal where speed and rapid colour identification is required. Humans have fewer types of photoreceptors, but more colour-tuned neurons, while mantis shrimps appears to have fewer colour neurons and more classes of photoreceptors.
An October 2014 publication by researchers from the University of Queensland stated that the compound eyes of mantis shrimp are capable of detecting cancer and the activity of neurons, since they are sensitive to detecting polarised light that reflects differently off cancerous and healthy tissue. The study claims that this ability can be replicated through a camera through the use of aluminium nanowires to replicate polarisation-filtering microvilli on top of photodiodes.
Mantis shrimp are long-lived and exhibit complex behaviour, such as ritualised fighting. Some species use fluorescent patterns on their bodies for signalling with their own and maybe even other species, expanding their range of behavioural signals. They can learn and remember well, and are able to recognise individual neighbours with whom they frequently interact. They can recognise them by visual signs and even by individual smell. Many have developed complex social behaviour to defend their space from rivals.
In a lifetime, they can have as many as 20 or 30 breeding episodes. Depending on the species, the eggs can be laid and kept in a burrow, or they can be carried around under the female's tail until they hatch. Also depending on the species, male and female may come together only to mate, or they may bond in monogamous long-term relationships.
In the monogamous species, the mantis shrimp remain with the same partner for 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 (biparental care). In Pullosquilla and some species in Nannosquilla, the female will lay two clutches of eggs: one that the male tends and one that the female tends. In other species, the female will look after the eggs while the male hunts for both of them. Once the eggs hatch, the offspring may spend up to three months as plankton.
Although stomatopods typically display the standard locomotion types as seen in true shrimp and lobsters, one species, Nannosquilla decemspinosa, has been observed flipping itself into a crude wheel. The species lives in shallow, sandy areas. At low tides, N. decemspinosa is often stranded by its short rear legs, which are sufficient for locomotion when the body is supported by water, but not on dry land. The mantis shrimp then performs a forward flip in an attempt to roll towards the next tide pool. N. decemspinosa has been observed to roll repeatedly for 2 metres (6.6 ft), but specimens typically travel less than 1 m (3.3 ft).
Mantis shrimp are abundant in the coastal regions of south Vietnam, known in Vietnamese as tôm tít or tôm tích. The shrimp can be steamed, boiled, grilled or dried; used with pepper, salt, and lime; fish sauce and tamarind; or fennel.
In Cantonese cuisine, the mantis shrimp is known as "pissing shrimp" (Chinese: 攋尿蝦; pinyin: lài niào xiā; Jyutping: laaih niu hā) 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. Usually they are deep fried with garlic and chili peppers.
In the Philippines, the mantis shrimp is known as tatampal, hipong-dapa or alupihang-dagat and is cooked and eaten like shrimp.
The usual concerns associated with consuming seafood caught in contaminated waters apply to mantis shrimp. In Hawaii, some have grown unusually large in the very dirty waters of the Grand Ala Wai Canal in Waikiki.
While some aquarists value mantis shrimp, others consider them harmful pests, because:
- They are voracious predators, eating other desirable inhabitants of the tank,
- Some of the largest species can break aquarium glass by striking it
- Some rock-burrowing species can do more damage to live rock than the fishkeeper would prefer
The live rock with mantis shrimp burrows are actually considered useful by some in the marine aquarium trade and are often collected. It is not uncommon for a piece of live rock to convey a live mantis shrimp into an aquarium. Once inside the tank, they may feed on fish, and other inhabitants. They are notoriously difficult to catch when established in a well-stocked tank, and there are accounts of them breaking glass tanks. It should be noted that while stomatopods do not eat coral, the smashers can damage it if they wish to make a home within it.
- Family Gonodactylidae
- Family Lysiosquillidae
- Lysiosquillina maculata, zebra mantis shrimp or striped mantis shrimp - the largest species
- Family Nannosquillidae
- Family Odontodactylidae
- Odontodactylus scyllarus, peacock mantis shrimp
- Family Pseudosquillidae
- Pseudosquilla ciliata, common mantis shrimp
- Family Squillidae
- Family Tetrasquillidae
- Heterosquilla tricarinata, New Zealand
A large number of the mantis shrimp species were first scientifically described by one carcinologist, Raymond B. Manning, and the collection of stomatopods he amassed is the largest in the world, covering 90% of the known species.
- Joel W. Martin & George E. Davis (2001). An Updated Classification of the Recent Crustacea (PDF). Natural History Museum of Los Angeles County. p. 132.
- James Gonser (February 14, 2003). "Large shrimp thriving in Ala Wai Canal muck". Honolulu Advertiser.
- "Huge Mantis shrimp" (in Chinese).
- Ross Piper (2007). Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals. Greenwood Press. ISBN 0-313-33922-8.
- "Mantis shrimps", Queensland Museum
- Gilbert L. Voss (2002). "Order Stomatopoda: Mantis shrimp or thumb splitters". Seashore Life of Florida and the Caribbean. Dover pictorial archive series. Courier Dover Publications. pp. 120–122. ISBN 978-0-486-42068-4.
- April Holladay (September 1, 2006). "Shrimp spring into shattering action". USA Today.
- "Stomatopoda". Tree of Life Web Project. January 1, 2002.
- S. N. Patek, W. L. Korff & R. L. Caldwell (2004). "Deadly strike mechanism of a mantis shrimp" (PDF). Nature 428 (6985): 819–820. Bibcode:2004Natur.428..819P. doi:10.1038/428819a. PMID 15103366.
- S. N. Patek & R. L. Caldwell (2005). "Extreme impact and cavitation forces of a biological hammer: strike forces of the peacock mantis shrimp". Journal of Experimental Biology 208 (19): 3655–3664. doi:10.1242/jeb.01831. PMID 16169943.
- Milius, Susan (2012). "Mantis shrimp flub color vision test". Science News 182 (6): 11. JSTOR 23351000.
- Justin Marshall & Johannes Oberwinkler (1999). "Ultraviolet vision: the colourful world of the mantis shrimp". Nature 401 (6756): 873–874. Bibcode:1999Natur.401..873M. doi:10.1038/44751. PMID 10553902.
- Patrick Kilday (September 28, 2005). "Mantis shrimp boasts most advanced eyes". The Daily Californian.
- Mantis shrimp wear tinted shades to see UV light
- David Cowles, Jaclyn R. Van Dolson, Lisa R. Hainey & Dallas M. Dick (2006). "The use of different eye regions in the mantis shrimp Hemisquilla californiensis Stephenson, 1967 (Crustacea: Stomatopoda) for detecting objects". Journal of Experimental Marine Biology and Ecology 330 (2): 528–534. doi:10.1016/j.jembe.2005.09.016.
- DuRant, Hassan (3 July 2014). "Mantis shrimp use 'nature's sunblock' to see UV". sciencemag.org. Retrieved 5 July 2014.
- Thomas W. Cronin & Justin Marshall (2001). "Parallel processing and image analysis in the eyes of mantis shrimps". The Biological Bulletin 200 (2): 177–183. PMID 11341580.
- Tsyr-Huei Chiou, Sonja Kleinlogel, Tom Cronin, Roy Caldwell, Birte Loeffler, Afsheen Siddiqi, Alan Goldizen & Justin Marshall (2008). "Circular polarization vision in a stomatopod crustacean". Current Biology 18 (6): 429–34. doi:10.1016/j.cub.2008.02.066. PMID 18356053.
- Sonja Kleinlogel & Andrew White (2008). "The secret world of shrimps: polarisation vision at its best". PLoS ONE 3 (5): e2190. arXiv:0804.2162. Bibcode:2008PLoSO...3.2190K. doi:10.1371/journal.pone.0002190. PMC 2377063. PMID 18478095.
- N. W. Roberts, T. H. Chiou, N. J. Marshall & T. W. Cronin (2009). "A biological quarter-wave retarder with excellent achromaticity in the visible wavelength region". Nature Photonics 3 (11): 641–644. Bibcode:2009NaPho...3..641R. doi:10.1038/nphoton.2009.189.
- Chris Lee (November 1, 2009). "A crustacean eye that rivals the best optical equipment". Nobel Intent. Ars Technica.
- Anne Minard (May 19, 2008). ""Weird beastie" shrimp have super-vision". National Geographic News.
- Bristol University: Mantis shrimps could show us the way to a better DVD, 25 October 2009
- C. H. Mazel, T. W. Cronin, R. L. Caldwell & N. J. Marshall (2004). "Fluorescent enhancement of signaling in a mantis shrimp". Science 303 (5654): 51. doi:10.1126/science.1089803. PMID 14615546.
- Mantis shrimp's super colour vision debunked
- Parallels Between Mantis Shrimp and Human Color Vision
- T. York, S. Powell, S. Gao, L. Kahan, T. Charanya, D. Saha, N. Roberts, T. Cronin, J. Marshall, S. Achilefu, S. Lake, B. Raman & V. Gruev (2014). "Bioinspired polarization imaging sensors: from circuits and optics to signal processing algorithms and biomedical applications". Proceedings of the IEEE 102 (10): 1450–1469. doi:10.1109/JPROC.2014.2342537.
- "Nature's elegant and efficient vision systems can detect cancer". University of Queensland. September 22, 2014. Retrieved November 21, 2014.
- "Sharing the job: monogamy and parental care". University of California, Berkeley.
- Roy L. Caldwell (1979). "A unique form of locomotion in a stomatopod – backward somersaulting". Nature 282 (5734): 71–73. Bibcode:1979Natur.282...71C. doi:10.1038/282071a0.
- "Tôm tít – Đặc sản miền sông nước" (in Vietnamese). Dinh dưỡng. October 1, 2009. Retrieved January 8, 2011.
- A Load of Learnin' About Mantis Shrimps, by James Fatherree, in ReefKeeping online magazine.
- Nick Dakin (2004). The Marine Aquarium. London: Andromeda. ISBN 1-902389-67-0.
- Paul F. Clark & Frederick R. Schram (2009). "Raymond B. Manning: an appreciation". Journal of Crustacean Biology 29 (4): 431–457. doi:10.1651/09-3158.1.
|Wikispecies has information related to: Stomatopoda|
|Wikimedia Commons has media related to Stomatopoda.|
- Hoplocarida: Stomatopoda fact sheet – Guide to the marine zooplankton of south eastern Australia
- The Lurker's Guide to Stomatopods – mantis shrimp
- Mantis shrimp – colourful and aggressive
- Research on Stomatopods at the University of Maryland
- "Mantis shrimp". The Aquarium Wiki. April 7, 2006.
- Caldwell Lab at the University of California, Berkeley
- Patek Lab at the University of Massachusetts, Amherst
- High speed video of mantis shrimp
- P. Z. Myers (May 24, 2008). "The superior eyes of shrimp". Pharyngula.
- Daniel Cressey (May 14, 2008). "Shrimp's super sight". The Great Beyond. nature.com.