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Mantises are an order (Mantodea) of insects that contains over 2,400 species in about 460 genera in 33 families. The largest family is the Mantidae ("mantids"). Mantises are distributed worldwide in temperate and tropical habitats. They have triangular heads with bulging eyes supported on flexible necks. Their elongated bodies may or may not have wings, but all mantodeans have forelegs that are greatly enlarged and adapted for catching and gripping prey; their upright posture, while remaining stationary with forearms folded, resembling a praying posture, has led to the common name praying mantis.

Etymology
The name mantodea is formed from the Ancient Greek words (mántis) meaning "prophet", and (eîdos) meaning "form" or "type". It was coined in 1838 by the German entomologist Hermann Burmeister. The name "mantid" properly refers only to members of the family Mantidae, which was, historically, the only family in the order. The other common name, praying mantis, applied to any species in the order (though in Europe mainly to Mantis religiosa), comes from the typical prayer-like posture these mantises adopt when their forelegs are folded. The vernacular form "mantises" (used in this article, singular "Mantis") was originally confined largely to the US, with "mantids" (singular "mantid") predominantly used for the group in the UK and elsewhere, until the family Mantidae was further split in 2002; at present, only some 75 out of 430 known genera are mantids, the rest are in 28 other families, and therefore no longer "mantids". == Taxonomy and evolution ==
Taxonomy and evolution
'' Over 2,400 species of mantises in about 430 genera are recognized. They are predominantly found in tropical regions, but some live in temperate areas. The systematics of mantises have long been disputed. Mantises, along with stick insects (Phasmatodea), were once placed in the order Orthoptera with the cockroaches (now Blattodea) and ice crawlers (now Grylloblattodea). Kristensen (1991) combined the Mantodea with the cockroaches and termites into the order Dictyoptera, suborder Mantodea. Phylogeny External Evolutionary relationships based on Evangelista et al. 2019 are shown in the cladogram: }} Internal One of the earliest classifications splitting an all-inclusive Mantidae into multiple families was that proposed by Beier in 1968, recognizing eight families, though it was not until Ehrmann's reclassification into 15 families in 2002 However, as previously configured, the Mantidae and Thespidae especially were considered polyphyletic, so the Mantodea have been revised substantially as of 2019 and now includes 29 families. {{cladogram |1=† Extinct Genera |label2=Eumantodea |2={{clade |label1=Chaeteessoidea |1=Chaeteessidae |label2=Spinomantodea |2={{clade |label1=Mantoidoidea |1=Mantoididae |label2=Schizomantodea |2={{clade |label1=Metallyticoidea |1=Metallyticidae |label2=Artimantodea |2={{clade |label1=Amerimantodea |1= |label2=Cernomantodea |2={{clade |label1=Nanomantodea |1= |label2=Metamantodea |2={SUBCLADE_Metamantodea} }} }} }} }} }} }} |targetA={SUBCLADE_Metamantodea} |subcladeA= }} Fossil mantises '', a primitive fossil mantis known from the Early Cretaceous of Brazil, and one of the oldest members of the group Mantises are thought to have evolved from cockroach-like ancestors. Some of the earliest confidently identified mantis fossils date to the Early Cretaceous, Fossil mantises, including one from Japan with spines on the front legs as in modern mantises, have been found in Cretaceous amber. Most fossils in amber are nymphs; compression fossils (in rock) include adults. Fossil mantises from the Crato Formation in Brazil include the long Santanmantis axelrodi, described in 2003; as in modern mantises, the front legs were adapted for catching prey. Well-preserved specimens yield details as small as 5 μm through X-ray computed tomography. == Biology ==
Biology
Anatomy Mantises have large, triangular heads with a beak-like snout and mandibles. They have two bulbous compound eyes, three small simple eyes, and a pair of antennae. The articulation of the neck is also remarkably flexible; some species of mantis can rotate their heads nearly 180°. Mantises also are unique to the Dictyoptera in that they have tympanate hearing, with two tympana in an auditory chamber in their metathorax. Most mantises can only hear ultrasound. Mantises have two spiked, grasping forelegs ("raptorial legs") in which prey items are caught and held securely. In most insect legs, including the posterior four legs of a mantis, the coxa and trochanter combine as an inconspicuous base of the leg; in the raptorial legs, however, the coxa and trochanter combine to form a segment about as long as the femur, which is a spiky part of the grasping apparatus (see illustration). Located at the base of the femur is a set of discoidal spines, usually four in number, but ranging from none to as many as five depending on the species. These spines are preceded by a number of tooth-like tubercles, which, along with a similar series of tubercles along the tibia and the apical claw near its tip, give the foreleg of the mantis its grasp on its prey. The foreleg ends in a delicate tarsus used as a walking appendage, made of four or five segments and ending in a two-toed claw with no arolium. The abdomen of all mantises consists of 10 tergites, with a corresponding set of nine sternites visible in males and seven visible in females. The abdomen tends to be slimmer in males than females, but ends in a pair of cerci in both sexes. They locate their prey by sight; their compound eyes contain up to 10,000 ommatidia. A small area at the front called the fovea has greater visual acuity than the rest of the eye, and can produce the high resolution necessary to examine potential prey. The peripheral ommatidia are concerned with perceiving motion; when a moving object is noticed, the head is rapidly rotated to bring the object into the visual field of the fovea. Further motions of the prey are then tracked by movements of the mantis's head so as to keep the image centered on the fovea. The use of stereoscopic vision differs from humans or primates because they specifically utilize this vision for capturing and spotting prey. The eyes are widely spaced and laterally situated, affording a wide binocular field of vision and precise stereoscopic vision at close range. The dark spot on each eye that moves as it rotates its head is a pseudopupil. This occurs because the ommatidia that are viewed "head-on" absorb the incident light, while those to the side reflect it. As their hunting relies heavily on vision, mantises are primarily diurnal. Many species, however, fly at night and may then be attracted to artificial lights. They have good night vision. Male mantises in the family Liturgusidae are more frequently collected at night, suggesting greater nocturnal activity or attraction to light sources. This pattern likely extends to other mantis families, where males are also more commonly observed during nighttime surveys. Nocturnal flight is especially important to males in locating less-mobile females by detecting their pheromones. Flying at night exposes mantises to fewer bird predators than diurnal flight would. Many mantises also have an auditory thoracic organ that helps them avoid bats by detecting their echolocation calls and responding evasively. Diet and hunting Mantises are generalist predators of arthropods. Larger mantises sometimes eat smaller individuals of their own species, as well as small vertebrates such as lizards, frogs, fish, and particularly small birds. Most mantises stalk tempting prey if it strays close enough, and will go farther when they are especially hungry. Once within reach, mantises strike rapidly to grasp the prey with their spiked raptorial forelegs. Some ground and bark species pursue their prey in a more active way. For example, members of a few genera such as the ground mantises Entella, Ligaria, and Ligariella run over dry ground seeking prey, much as tiger beetles do. The foregut of some species extends the whole length of the insect and can be used to store prey for digestion later. This may be advantageous in an insect that feeds intermittently. Chinese mantises live longer, grow faster, and produce more young when they are able to eat pollen. File:Tenodera sinensis 2 Luc Viatour.jpg|Tenodera sinensis feeding on a cricket File:Pm eating pm.webm|thumbtime=46|Tenodera sinensis consuming a smaller immature mantis File:Mantis catches grasshopper.webm|thumbtime=3|A chinese mantis catches a differential grasshopper Antipredator adaptations Mantises are preyed on by vertebrates such as frogs, lizards, and birds, and by invertebrates such as spiders, large species of hornets, and ants. Some hunting wasps, such as some species of Tachytes, also paralyze some species of mantis to feed their young. Generally, mantises protect themselves by camouflage, most species being cryptically colored to resemble leaves or other backgrounds, both to avoid predators and to better snare their prey. Those that live on uniformly colored surfaces such as bare earth or tree bark are dorsoventrally flattened so as to eliminate shadows that might reveal their presence. The species from different families called flower mantises are aggressive mimics: they resemble flowers convincingly enough to attract prey that come to collect pollen and nectar. Some species in Africa and Australia are able to turn black after a molt towards the end of the dry season; at this time of year, bush fires occur and this coloration enables them to blend in with the fire-ravaged landscape (fire melanism). Mantises, like stick insects, show rocking behavior in which the insect makes rhythmic, repetitive side-to-side movements. Functions proposed for this behavior include the enhancement of crypsis by means of the resemblance to vegetation moving in the wind. However, the repetitive swaying movements may be most important in allowing the insects to discriminate objects from the background by their relative movement, a visual mechanism typical of animals with simpler sight systems. Rocking movements by these generally sedentary insects may replace flying or running as a source of relative motion of objects in the visual field. As ants may be predators of mantises, genera such as Loxomantis, Orthodera, and Statilia, like many other arthropods, avoid attacking them. A variety of arthropods, including some early-instar mantises, exploit this behavior and mimic ants to evade their predators. File:Choeradodis stalii MHNT dos.jpg|Leaf mimicry: Choeradodis has leaf-like forewings and a widened green thorax. File:Gottesanbeterin Abwehr.JPG|Adult female Iris oratoria performs a bluffing threat display, rearing back with the forelegs and wings spread and mouth opened. File:Gemmatus.jpg|The jeweled flower mantis, Creobroter gemmatus: the brightly colored wings are opened suddenly in a deimatic display to startle predators. File:Ant-Mantis.jpg|Some mantis nymphs mimic ants to avoid predators. File:Schizocephala bicornis, Grass Mantis, Udaipur.jpg|The grass mantis Schizocephala bicornis mimics dry grass both to avoid predators and to approach unwitting prey. File:Mantis Hymenopus coronatus 2 Luc Viatour.jpg|Aggressive mimicry: Malaysian orchid mantises are camouflaged pink or yellow, matching the coloration of local orchids. Reproduction and life history The mating season in temperate climates typically takes place in autumn, To mate following courtship, the male usually leaps onto the female's back, clasping her thorax and wing bases with his forelegs. He then arches his abdomen to deposit and store sperm in a special chamber near the tip of the female's abdomen. The female lays between 10 and 400 eggs, depending on the species. Eggs are typically deposited in a froth mass-produced by glands in the abdomen. This froth hardens, creating a protective capsule, which together with the egg mass is called an ootheca. Depending on the species, the ootheca can be attached to a flat surface, wrapped around a plant, or even deposited in the ground. The cryptic Tarachodes maurus positions herself on bark with her abdomen covering her egg capsule, ambushing passing prey and moving very little until the eggs hatch. In temperate climates, adults do not survive the winter and the eggs undergo a diapause, hatching in the spring. Around 90% of the predatory species of mantises exhibit sexual cannibalism. Adult males typically outnumber females at first, but their numbers may be fairly equivalent later in the adult stage, In Tenodera sinensis, 83% of males escape cannibalism after an encounter with a female, but since multiple matings occur, the probability of a male's being eaten increases cumulatively. Under such circumstances, the female has been known to respond with a defensive deimatic display by flashing the colored eyespots on the inside of her front legs. The reason for sexual cannibalism has been debated; experiments show that females on poor diets are likelier to engage in sexual cannibalism than those on good diets. Some hypothesize that submissive males gain a selective advantage by producing offspring; this is supported by a quantifiable increase in the duration of copulation among males which are cannibalized, in some cases doubling both the duration and the chance of fertilization. This is contrasted by a study where males were seen to approach hungry females with more caution, and were shown to remain mounted on hungry females for a longer time, indicating that males that actively avoid cannibalism may mate with multiple females. The same study also found that hungry females generally attracted fewer males than those that were well fed. The act of dismounting after copulation is dangerous for males, for it is the time that females most frequently cannibalize their mates. An increase in mounting duration appears to indicate that males wait for an opportune time to dismount a hungry female, who would be likely to cannibalize her mate. == Relationship with humans ==
Relationship with humans
In culture, literature and art rest in the shape of a mantis, Edo period, Japan, c. 1800 One of the earliest mantis references is in the ancient Chinese dictionary Erya, which gives its attributes in poetry, where it represents courage and fearlessness, and a brief description. A later text, the () from 1108, gives accurate details of the construction of the egg packages, the development cycle, anatomy, and the function of the antennae. Although mantises are rarely mentioned in Ancient Greek sources, a female mantis in threat posture is accurately illustrated on a series of fifth-century BC silver coins, including didrachms, from Metapontum in Lucania. The 10th-century Byzantine encyclopedia Suda described the insect called "mantis" (μάντις) as a pale green, clumsy, and slow-moving locust, adding that some people observed its movements for the purpose of augury. In addition, the Suda mentions the phrase "arouraia mantis" (Ἀρουραία μάντις), explaining that it was a proverbial expression used to mock people who were sluggish and ineffectual but still treated as if they had wisdom or insight. He translates Zenobius 2.94 with the words seriphos (maybe a mantis) and graus, an old woman, implying a thin, dried-up stick of a body. Mantises are a common motif in Luna Polychrome ceramics of pre-Columbian Nicaragua, and are believed to represent a deity or spirit called "Madre Culebra". Western descriptions of the biology and morphology of the mantises became more accurate in the 18th century. Roesel von Rosenhof illustrated and described mantises and their cannibalistic behavior in the '' (Insect Entertainments''). Aldous Huxley made philosophical observations about the nature of death while two mantises mated in the sight of two characters in his 1962 novel Island (the species was Gongylus gongylodes). The naturalist Gerald Durrell's humorously autobiographical 1956 book My Family and Other Animals includes a four-page account of an almost evenly matched battle between a mantis and a gecko. Shortly before the fatal dénouement, Durrell narrates: M. C. Escher's woodcut Dream depicts a human-sized mantis standing on a sleeping bishop. A cultural trope imagines the female mantis as a femme fatale. The idea is propagated in cartoons by Cable, Guy and Rodd, LeLievre, T. McCracken, and Mark Parisi, among others. It ends Isabella Rossellini's short film about the life of a praying mantis in her 2008 Green Porno season for the Sundance Channel. The Deadly Mantis is a 1957 American science fiction monster film, with a giant mantis threatening mankind. Martial arts , Shi DeRu and Shi DeYang, demonstrating the Southern Praying Mantis style of martial art Two martial arts separately developed in China have movements and fighting strategies based on those of the mantis. and 'Southern Praying Mantis'. Both are very popular in China, and have also been exported to the West in recent decades. In mythology and religion According to local beliefs in Africa, this insect brings good luck. The mantis was revered by the southern African Khoi and San in whose cultures man and nature were intertwined; for its praying posture, the mantis was even named '' ("god of the Hottentots") in the Afrikaans language that had developed among the first European settlers. However, at least for the San, the mantis was only one of the manifestations of a trickster-deity, ǀKaggen, who could assume many other forms, such as a snake, hare or vulture. Several ancient civilizations did consider the insect to have supernatural powers; for the Greeks, it had the ability to show lost travelers the way home; in the Ancient Egyptian Book of the Dead, the "bird-fly" is a minor god that leads the souls of the dead to the underworld; in a list of 9th-century BC Nineveh grasshoppers (buru), the mantis is named necromancer (buru-enmeli) and soothsayer (buru-enmeli-ashaga''). Some pre-Columbian cultures in western Nicaragua have preserved oral traditions of the mantis as "Madre Culebra", a powerful predator and symbol of female symbolic authority. Because the lifespan of a mantis is only about a year, people who want to keep mantises often breed them. In 2013 at least 31 species were kept and bred in the United Kingdom, the Netherlands, and the United States. In 1996 at least 50 species were known to be kept in captivity by members of the Mantis Study Group. For pest control Naturally occurring mantis populations provide plant pest control. Gardeners who prefer to avoid pesticides may encourage mantises in the hope of controlling insect pests. However, mantises do not have key attributes of biological pest control agents; they do not specialize in a single pest insect, and do not multiply rapidly in response to an increase in such a prey species, but are general predators. They therefore have "negligible value" in biological control. Robotics In 2016, the Association for the Advancement of Artificial Intelligence had produced a prototype robot inspired by the forelegs of the praying mantis, with front legs that allow the robot to walk, climb steps, and grasp objects. The multi-jointed leg provides dexterity via a rotatable joint. Future models may include a more spiked foreleg to improve the grip and ability to support more weight. ==See also==
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