Mantis


Mantises are an order of insects that contains over 2,400 species in about 460 genera in 33 families. The largest family is the Mantidae. 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.
The closest relatives of mantises are termites and cockroaches, which are all within the superorder Dictyoptera. Mantises are sometimes confused with stick insects, other elongated insects such as grasshoppers, or other more distantly related insects with raptorial forelegs such as mantisflies. Mantises are mostly ambush predators, but a few ground-dwelling species are found actively pursuing their prey. They normally live for about a year. In cooler climates, the adults lay eggs in autumn, then die. The eggs are protected by their hard capsules and hatch in the spring. Females sometimes practice sexual cannibalism, eating their mates after copulation.
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.

Etymology

The name mantodea is formed from the Ancient Greek words meaning "prophet", and 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, comes from the typical prayer-like posture these mantises adopt when their forelegs are folded. The vernacular plural "mantises" was originally confined largely to the US, with "mantids" predominantly used as the plural 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

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, were once placed in the order Orthoptera with the cockroaches and ice crawlers. Kristensen 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 that a multiple-family classification became universally adopted. 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, 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.

Fossil mantises

Mantises are thought to have evolved from cockroach-like ancestors. Some of the earliest confidently identified mantis fossils date to the Early Cretaceous, although the Jurassic taxon Lovec was identified in 2024 from the Karabastau Formation. Fossils of the group are rare: by 2022, 37 fossil species are known. 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 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. Extinct families and genera include:
  • †Juramantidae
  • †Lovecidae
  • †Baissomantidae
  • †Gryllomantidae
  • †Cretomantidae
  • †Santanmantidae
  • †Amelidae
  • Incertae sedis:
  • * †Jersimantis
  • * †Chaeteessites
  • * †Cretophotina
  • * †''Ambermantis''

    Similar insects in the [Neuroptera]

Because of the superficially similar raptorial forelegs, mantidflies may be confused with mantises, though they are unrelated. Their similarity is an example of convergent evolution; mantidflies do not have tegmina like mantises, their antennae are shorter and less thread-like, and the raptorial tibia is more muscular than that of a similar-sized mantis and bends back farther in preparation for shooting out to grasp prey.

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°.
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 movements of the head and fore limbs while the remainder of the body remains more or less immobile. 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 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. 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.
Mantises can be loosely categorized as being macropterous, brachypterous, micropterous, or apterous. If not wingless, a mantis has two sets of wings: the outer wings, or tegmina, are usually narrow and leathery. They function as camouflage and as a shield for the hindwings, which are clearer and more delicate.
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.

Vision

Mantises have stereo vision. 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. 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 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 further 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. Some mantis species such as Euantissa pulchra can discriminate between different types of prey, and approach spiders mimicking non-aggressive ant species much more than spiders that mimick aggressive ant species.
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.