Temporal range: 145–0 Ma Cretaceous–Recent
|Adult female Sphodromantis viridis|
Mantodea is an order of insects that contains over 2,400 species and about 430 genera of mantises in 15 families, by far the largest family being Mantidae. Mantises are distributed worldwide in temperate and tropical habitats. They are mostly ambush predators, but a few ground-dwelling species actively pursue their prey. Their forelegs are greatly enlarged and are adapted for catching and gripping prey; their upright posture, while remaining stationary with forearms folded, has led to the name praying mantises. Their lifespan is normally about a year and in cooler climates, the adults die in winter, the autumn-laid eggs, protected by their capsule of hardened foam, hatching in the spring. Females sometimes practice sexual cannibalism, eating their mate after mating, or occasionally decapitating the male just before or during mating.
The closest relatives of mantises are the termites and cockroaches (Blattodea), the three groups being included in the superorder Dictyoptera. Mantises are sometimes confused with stick insects (Phasmatodea) and other elongated insects such as grasshoppers (Orthoptera), or other insects with raptorial forelegs such as mantisflies (Mantispidae).
Mantises were considered to have supernatural powers by early civilizations including Ancient Greece, Ancient Egypt and Assyria. A cultural trope popular in cartoons imagines the female mantis as a femme fatale. Mantises are among the insects most commonly kept as pets.
- 1 Etymology
- 2 Taxonomy and evolution
- 3 Biology
- 4 In human culture
- 5 References
- 6 External links
The name mantodea is formed from the Ancient Greek words μάντις (mantis) meaning "prophet", and εἶδος (eidos) meaning "form" or "type". It was coined in 1838 by the German entomologist Hermann Burmeister. The order is occasionally called the mantes, using a Latinized plural of Greek mantis. The name mantid properly refers only to members of the family Mantidae, which was, historically, the only family in the order, but with 14 additional families recognized in recent decades, this term is now potentially misleading. The other common name, praying mantis, applied to any species in the order, but in Europe mainly to Mantis religiosa, comes from the typical "prayer-like" posture with folded fore-limbs.
Taxonomy and evolution
There are over 2400 recognized species of mantis in about 430 genera. 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 rock crawlers (now Grylloblattodea). Kristensen (1991) combined Mantodea with the cockroaches and termites into the order Dictyoptera, suborder Mantodea.
The classification most commonly adopted is that proposed by Beier in 1968. He divided the order into eight families. Klass, in 1997, studied the external male genitalia and postulated that the families Chaeteessidae and Metallyticidae diverged from the other families at an early date. However, the Mantidae and Thespidae are both polyphyletic, so the Mantoidea will have to be revised.
The earliest mantis fossils are about 135 million years old, from Siberia. Fossils of the group are rare: by 2007 there were only about 25 fossil species. Fossil mantises, including one from Japan with spines on the front legs as in modern mantids, have been found in Cretaceous amber. Most fossils in amber are nymphs; compression fossils (in rock) include adults. Fossil mantids from the Crato Formation in Brazil include the 10mm long Santanmantis axelrodi, described in 2003; as in modern mantids, the front legs were adapted for catching prey. Well-preserved specimens yield details as small as 5 μmetre through X-ray computed tomography.
Because of the similar raptorial forelegs, mantidflies may be confused with Mantises. This similarity is an example of convergent evolution. Mantidflies do not have the leathery forewings of mantises.
Mantises have large triangular heads with a beak-like snout and biting and chewing mouthparts. 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 the head nearly 180 degrees. Mantises locate their prey by sight and have a visual range of up to twenty meters. Their compound eyes contain up to ten thousand ommatidia. A small area at the front called the "fovea" can produce the high resolution necessary to examine potential prey. The peripheral ommatidia are concerned with perceiving motion; when a moving object is perceived, the head is rapidly rotated to bring the object into the field of view of the fovea. Further movements of the prey are then tracked by movements of the mantis' head so as to keep the image centred on the fovea. The eyes are widely spaced and laterally situated, affording a wide binocular field of vision and, at close range, precise stereoscopic vision. The dark spot on each eye that moves as it rotates its head is a pseudopupil. This occurs because the ommatidia 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 then may be attracted to artificial lights. Nocturnal flight is especially important to males in search of less-mobile females that they locate 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 to avoid bats by detecting their echolocation calls and responding evasively.
The mantis thorax consists of a prothorax, a mesothorax, and a metathorax. In all species apart from the genus Mantoida, the prothorax, which bears the head and forelegs, is much longer than the other two thoracic segments. The prothorax is also flexibly articulated, allowing for a wide range of movement of the head and forelimbs while the remainder of the body remains more or less immobile.
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 are a set of discoidal spines, usually four in number, but ranging from zero 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 made of between four and five segments and ending in a two-toed claw with no arolium and used as a walking appendage.
Mantises can be loosely categorized as being macropterous (long-winged), brachypterous (short-winged), micropterous (vestigial-winged), or apterous (wingless). If not wingless, a mantis has two sets of wings: the outer wings, or tegmina, are usually narrow, opaque, and leathery. They function as camouflage and as a shield for the hind wings. The hind wings are much broader, more delicate, and transparent. In species able to fly, they are the main organs of flight. Brachypterous species are at most minimally capable of flight, other species not at all. The wings are mostly erected in these mantises for alarming enemies and attracting females. Even in many macropterous species the female is much heavier than the male, has much shorter wings, and rarely takes flight.
The abdomen of all mantises consist of ten tergites with a corresponding set of nine sternites visible in males and seven visible in females. The slim abdomen of most males allows them to take flight more easily while the thicker abdomen of the females houses the reproductive machinery for generating the ootheca. The abdomen of both sexes ends in a pair of cerci.
Diet and predation
Mantises are generalist predators of arthropods. Insects are their primary prey, but the diet of a mantis changes as it grows larger. In its first instar a mantis eats small insects. In the final instar the diet still includes more insects than anything else, but large species sometimes tackle increasingly large prey. Mantises have been observed catching and eating hummingbirds, mice and snakes among other vertebrates.
The majority of mantises are ambush predators that only feed upon live prey within their reach. They either camouflage themselves and remain stationary, waiting for prey to approach, or stalk their prey with slow, stealthy movements. Larger mantises will eat smaller individuals of their own species. Most mantises chase tempting prey if it strays close enough, and will go further when they are especially hungry. Once prey is 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 mantids, 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.
Mantises are themselves predated by vertebrates such as frogs, lizards and birds, and by invertebrates such as spiders and ants.
Generally, mantises protect themselves by camouflage. When directly threatened, many mantis species stand tall and spread their forelegs, with their wings fanning out wide. The fanning of the wings makes the mantis seem larger and more threatening, with some species having bright colors and patterns on their hind wings and inner surfaces of their front legs for this purpose. If harassment persists, a mantis may strike with its forelegs and attempt to pinch or bite. As part of the bluffing (deimatic) threat display, some species also may produce a hissing sound by expelling air from the abdominal spiracles. When flying at night, at least some mantises are able to detect the echolocation sounds produced by bats, and when the frequency begins to increase rapidly, indicating an approaching bat, they stop flying horizontally and begin a descending spiral toward the safety of the ground, often preceded by an aerial loop or spin.
Mantises, like stick insects, show rocking behavior in which the insect makes rhythmic, repetitive side-to-side movements. Functions proposed for this behaviour 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.
Mantises are camouflaged, most species being cryptically colored to resemble foliage 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 dorso-ventrally flattened so as to eliminate shadows that night reveal their presence. The species from different families called flower mantises are aggressive mimics: they resemble flowers convincingly enough actually to attract their prey which 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).
Ants are among the major insect predators of mantises. In turn, mantises such as Loxomantis, Orthodera and Statilia, like many other arthropods, avoid attacking ants. Exploiting this behavior, a variety of arthropods including some early instar mantises mimic ants to evade their predators.
While mantises can bite, they have no venom. On perceiving an attack, mantises may give a deimatic display, rearing back and spreading their front legs and wings, and in some species revealing vivid colors and eyespots to startle the predator. If caught, they may slash captors with their raptorial legs. Mantises lack chemical protection, so their displays are largely bluff.
Reproduction and life history
The mating season in temperate climates typically begins in autumn. To mate following courtship, the male usually leaps onto the female’s back, and clasps 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 then lays between 10 and 400 eggs, depending on the species. Eggs are typically deposited in a frothy mass that is produced by glands in the abdomen. This froth then 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. Despite the versatility and durability of the eggs, they are often preyed on, especially by several species of parasitoid wasps. In a few species, mostly ground and bark mantids in the family Tarachodidae, the mother guards the eggs. 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. A unique reproductive strategy is adopted by Brunner's stick mantis from the southern United States. No males have ever been found in this species, and the females breed parthenogenetically.
As in closely-related insect groups in the superorder Dictyoptera, mantises go through three life stages: egg, nymph, and adult (mantises are among the hemimetabolous insects). The nymphs and adult insects are structurally quite similar, with the nymphs being smaller and having no wings or functional genitalia. The nymphs are also sometimes colored differently from the adult, and the early stages are often mimics of ants. A mantis nymph has a sturdy, flexible exoskeleton and increases in size (often changing its diet as it does so) by molting when necessary. Molting can happen from five to ten times before the adult stage is reached, depending on the species. After the final molt, most species have wings, though some species are wingless or brachypterous ("short-winged"), particularly in the female sex.
The natural lifespan of a tropical mantis in the wild is about 10–12 months, but some species kept in captivity have been sustained for 14 months. In colder areas, females die during the winter (as well as any surviving males).
Mantis religiosa mating (brown male, green female)
Sexual cannibalism is common among most predatory species of mantises in captivity. It has sometimes been observed in the field, where about a quarter of male-female encounters result in the male's being eaten by the female. 90% of the predatory species of mantis participate in sexual cannibalism. Adult males typically outnumber females at first, but their numbers may be fairly equivalent later in the adult stage, possibly because females selectively eat the smaller males. In Tenodera sinensis, 83% of males escaped cannibalism after an encounter with a female, but since there are multiple matings, the probability of a male's being cannibalised increases cumulatively.
The female may begin feeding by biting off the male’s head (as they do with regular prey), and if mating has begun, the male’s movements may become even more vigorous in its delivery of sperm. Early researchers thought that because copulatory movement is controlled by a ganglion in the abdomen, not the head, removal of the male’s head was a reproductive strategy by females to enhance fertilization while obtaining sustenance. Later, this behavior appeared to be an artifact of intrusive laboratory observation. Whether the behavior in the field is natural, or also the result of distractions caused by the human observer, remains controversial. Mantises are highly visual organisms, and notice any disturbance occurring in the laboratory or field such as bright lights or moving scientists. Research by Liske and Davis (1984) and others found (e.g. using video recorders in vacant rooms) that Chinese mantises that had been fed ad libitum (so that they were not hungry) actually displayed elaborate courtship behavior when left undisturbed. The male engages the female in a courtship dance, to change her interest from feeding to mating. 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; some consider that submissive males gain a selective advantage by producing offspring. This theory 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 which 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 one of the most dangerous times for males, for it is at this 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 when the female is hungry and therefore likely to cannibalize her mate.
In human culture
In literature and art
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 Jingshi Zhenglei Daguan Bencao 經史證類大觀本草 ("Great History of Medical Material Annotated and Arranged by Types, Based upon the Classics and Historical Works") 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 5th century B.C. silver coins, including didrachms, from Metapontum in Sicily. In the 10th century Byzantine era Adages, Suidas describes an insect like a slow-moving green locust with long front legs; translating Zenobius 2.94 with the words seriphos (maybe a mantis) and graus, an old woman, implying a thin, dried-up stick of a body.
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 behaviour in the Insekten-Belustigungen (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 denouement, Durrell narrates:
he [Geronimo the gecko] crashed into the mantis and made her reel, and grabbed the underside of her thorax in his jaws. Cicely [the mantis] retaliated by snapping both her front legs shut on Geronimo's hind legs. They rustled and staggered across the ceiling and down the wall, each seeking to gain some advantage.
A cultural trope imagines the female mantis as a femme fatale. The idea is propagated in cartoons by Cable, Guy & Rodd, LeLievre, T. McCracken, and Mark Parisi among others. It ends Isabella Rossellini's short film about the life of a praying mantis, one of her 2008 Green Porno season for The Sundance Channel.
Two martial arts separately developed in China have movements and fighting strategies based on those of the Mantis. As one of these arts was developed in northern China, and the other in southern parts of the country, the arts are nowadays referred to (both in English and Chinese) as 'Northern Praying Mantis' and 'Southern Praying Mantis'. Both are very popular in China, and have also been imported to the West in recent decades.
In mythology and religion
The mantis was considered a god in southern African Khoi and San tradition, for its praying posture; the word for the mantis in Afrikaans is Hottentotsgod ("god of the Khoi"). Several ancient civilisations considered the insect to have supernatural powers: for the Greeks, it had the ability to show lost travellers the way home; in the Ancient Egyptian Book of the Dead the "bird-fly" is a minor god who 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").
Mantises are among the insects most widely kept as pets. Because their lifespan is only about a year, mantis enthusiasts often breed the insects. At least 31 species are kept for breeding in the United Kingdom, the Netherlands and the USA.
For 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.
Two species, the Chinese mantis and the European mantis, were deliberately introduced to North America in the hope that they would serve as pest controls for agriculture; they have spread widely in both the United States and Canada.
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|Wikisource has the text of the 1911 Encyclopædia Britannica article Mantis.|
- Mantis Study Group Information on mantids, scientific article phylogenetics and Evolution.