Mantis shrimp or stomatopods are marine crustaceans, the members of the order Stomatopoda. They may reach 30 centimetres (12 in) in length, although exceptional cases of up to 38 cm (15 in) have been recorded. The carapace of mantis shrimp covers only the rear part of the head and the first four segments of the thorax. Mantis shrimp appear in a variety of colours, from shades of browns to bright neon colours. Although they are common animals and among the most important predators in many shallow, tropical and sub-tropical marine habitats 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. Although it happens rarely, some larger species of mantis shrimp are capable of breaking through aquarium glass with a single strike from this weapon.
Ecology
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 either wait for prey to chance upon them or, unlike most crustaceans, at times they hunt, chase, and kill prey. They rarely exit their homes except to feed and relocate, and can be diurnal, nocturnal, or crepuscular, depending on the species. Most species live in tropical and subtropical seas (Indian and Pacific Oceans between eastern Africa and Hawaii), although some live in temperate seas.
Classification and the claw
Around 400 species of mantis shrimp have currently been described worldwide; all living species are in the suborder Unipeltata. They are commonly separated into two distinct groups determined by the manner of claws they possess:
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, about the acceleration of a .22 calibre bullet. 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 kill or stun 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.
Eyes
The front of Lysiosquillina maculata, showing the stalked eyes
A colourful stomatopod, the peacock mantis shrimp, (Odontodactylus scyllarus) seen in the Andaman Sea off Thailand
The midband region of the mantis shrimp's eye is made up of six rows of specialized ommatidia. Four rows carry 16 differing sorts of photoreceptor pigments, 12 for colour sensitivity, others for colour filtering. The mantis shrimp has such good eyes it can perceive both polarized light and hyperspectral colour vision. Their eyes (both mounted on mobile stalks and constantly moving about independently of each other) are similarly variably coloured and are considered to be the most complex eyes in the animal kingdom. They permit both serial and parallel analysis of visual stimuli.
Each compound eye is made up of up to 10,000 separate ommatidia of the apposition type. Each eye consists of two flattened hemispheres separated by six parallel rows of highly specialised ommatidia, collectively called the midband, which divides the eye into three regions. This is a design which makes it possible for mantis shrimp to see objects with three different parts of the same eye. In other words, each individual eye possesses trinocular vision and depth perception. The upper and lower hemispheres are used primarily for recognition of forms and motion, not colour vision, like the eyes of many other crustaceans.
Rows 1–4 of the midband are specialised for colour vision, from ultra-violet to longer wavelengths, but aren't 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 adapted 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 segregated into different tiers too, 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 a small area of about 5°–10° of the visual field at any given instant, but like in most crustaceans, the eyes are mounted on stalks. In mantis shrimps the movement of the stalked eye is unusually free, and can be driven in all possible axes, up to at least 70°, of movement 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 four which are sensitive to ultraviolet light) and four of them for analysing polarised light. By comparison, humans have only five visual pigments, four dedicated to see colour but the 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 circular polarized light, and in some cases their biological quarter-wave plates perform more uniformly over the entire visual spectrum than any current man-made polarizing optics, the application of which it is speculated could be applied to a new type of optical media that performs even better than 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 for Stokes parameters, which yield a full description of polarization. It is thus believed to have optimal polarization vision.
Close-up of the trinocular vision of Pseudosquilla ciliata
Reasons given for powerful eyesight
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.
The fact that those with the most advanced vision also are the species with the most colourful bodies suggests the evolution of colour vision has taken the same direction as the peacock's tail.
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 for species living in shallow water near the shore.
Behavior
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).
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. Although it happens rarely, some larger species of mantis shrimp are capable of breaking through aquarium glass with a single strike from this weapon.
Ecology
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 either wait for prey to chance upon them or, unlike most crustaceans, at times they hunt, chase, and kill prey. They rarely exit their homes except to feed and relocate, and can be diurnal, nocturnal, or crepuscular, depending on the species. Most species live in tropical and subtropical seas (Indian and Pacific Oceans between eastern Africa and Hawaii), although some live in temperate seas.
Classification and the claw
Around 400 species of mantis shrimp have currently been described worldwide; all living species are in the suborder Unipeltata. 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, about the acceleration of a .22 calibre bullet. 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 kill or stun 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.
Eyes
The front of Lysiosquillina maculata, showing the stalked eyes
A colourful stomatopod, the peacock mantis shrimp, (Odontodactylus scyllarus) seen in the Andaman Sea off Thailand
The midband region of the mantis shrimp's eye is made up of six rows of specialized ommatidia. Four rows carry 16 differing sorts of photoreceptor pigments, 12 for colour sensitivity, others for colour filtering. The mantis shrimp has such good eyes it can perceive both polarized light and hyperspectral colour vision. Their eyes (both mounted on mobile stalks and constantly moving about independently of each other) are similarly variably coloured and are considered to be the most complex eyes in the animal kingdom. They permit both serial and parallel analysis of visual stimuli.
Each compound eye is made up of up to 10,000 separate ommatidia of the apposition type. Each eye consists of two flattened hemispheres separated by six parallel rows of highly specialised ommatidia, collectively called the midband, which divides the eye into three regions. This is a design which makes it possible for mantis shrimp to see objects with three different parts of the same eye. In other words, each individual eye possesses trinocular vision and depth perception. The upper and lower hemispheres are used primarily for recognition of forms and motion, not colour vision, like the eyes of many other crustaceans.
Rows 1–4 of the midband are specialised for colour vision, from ultra-violet to longer wavelengths, but aren't 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 adapted 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 segregated into different tiers too, 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 a small area of about 5°–10° of the visual field at any given instant, but like in most crustaceans, the eyes are mounted on stalks. In mantis shrimps the movement of the stalked eye is unusually free, and can be driven in all possible axes, up to at least 70°, of movement 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 four which are sensitive to ultraviolet light) and four of them for analysing polarised light. By comparison, humans have only five visual pigments, four dedicated to see colour but the 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 circular polarized light, and in some cases their biological quarter-wave plates perform more uniformly over the entire visual spectrum than any current man-made polarizing optics, the application of which it is speculated could be applied to a new type of optical media that performs even better than 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 for Stokes parameters, which yield a full description of polarization. It is thus believed to have optimal polarization vision.
Close-up of the trinocular vision of Pseudosquilla ciliata
Reasons given for powerful eyesight
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.
The fact that those with the most advanced vision also are the species with the most colourful bodies suggests the evolution of colour vision has taken the same direction as the peacock's tail.
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 for species living in shallow water near the shore.
Behavior
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).
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