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mantis
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{{Short description|Order of insects}}{{Other uses}}{{Redirect|Praying mantis}}{{Featured article}}{{Use dmy dates|date=February 2022}}{{Use American English|date=August 2015}}{{Automatic taxobox| taxon = Mantodea
Early Cretaceous | Recent}}| image = European praying mantis (Mantis religiosa) green female Dobruja.jpg| image_caption = Mantis religiosa, RomaniaHermann Burmeister>Burmeister, 1838| subdivision_ranks = Families#Taxonomy and evolution>text| synonyms = * Manteodea Burmeister, 1829
  • Mantearia
  • Mantoptera
}}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 Mantodea have forelegs that are greatly enlarged and adapted for catching and gripping prey; their upright posture, while remaining stationary with forearms folded, has led to the common name praying mantis.The closest relatives of mantises are termites and cockroaches (Blattodea), which are all within the superorder Dictyoptera. Mantises are sometimes confused with stick insects (Phasmatodea), other elongated insects such as grasshoppers (Orthoptera), or other more distantly related insects with raptorial forelegs such as mantisflies (Mantispidae). 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.

Taxonomy and evolution

Over 2,400 species of mantis in about 430 genera are recognized.WEB,weblink Mantodea Species File Online, Daniel, Otte, Lauren, Spearman, 17 July 2012, They are predominantly found in tropical regions, but some live in temperate areas.BOOK, Prete, Fredrick R., Wells, Harrington, Wells, Patrick H., Hurd, Lawrence E., The Praying Mantids, Mantid in Ecological Research, Hurd, I. E., 1999, Johns Hopkins University Press, 978-0-8018-6174-1, 231,weblink 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.BOOK, Costa, James, The Other Insect Societies, Harvard University Press, 135–136, 978-0-674-02163-1,weblink 2006, BOOK, Capinera, John L., Encyclopedia of Entomology,weblink 2008, 4, Springer, 978-1-4020-6242-1, 3033–3037, The name mantodea is formed from the Ancient Greek words (mantis) meaning "prophet", and (eidos) meaning "form" or "type". It was coined in 1838 by the German entomologist Hermann Burmeister.BOOK, College entomology, Essig, Edward Oliver, 1947, Macmillan Company, 809878, 124, 900,weblink {{OEtymD|mantis}} 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. The other common name, praying mantis, applied to any species in the orderBOOK, Bullock, William, 1812,weblink A companion to the London Museum and Pantherion, 12th, (though in Europe mainly to Mantis religiosa), comes from the typical "prayer-like" posture with folded forelimbs.BOOK, Partington, Charles Frederick, 1837,weblink The British Cyclopædia of Natural History, W. S. Orr, 1, WEB, Praying Mantis, 10 September 2010, National Geographic Society,weblinkweblink" title="web.archive.org/web/20100111130055weblink">weblink dead, 11 January 2010, 28 August 2015, The vernacular plural "mantises" (used in this article) was 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.Bragg, P. E. (1996). "Mantis, Mantid, Mantids, Mantises". Mantis Study Group Newsletter, 1:4.Ehrmann, R. 2002. Mantodea: Gottesanbeterinnen der Welt. Natur und Tier, Münster.

Phylogeny

External

Evolutionary relationships based on Evangelista et al. 2019 are shown in the cladogram:JOURNAL, Evangelista, D.A., Wipfler, B., O., Bethoux, Donath, A., Fujita, M., Kohli, M.K., Legendre, F., Liu, Machida, Misof, Peters, Podsiadlowski, Rust, Schuette, Tollenaar, Ware, Wappler, Zhou, Meusemann, Simon, An integrative phylogenomic approach illuminates the evolutionary history of cockroaches and termites (Blattodea), Proc. R. Soc. B, 23 January 2019, 286, 1895, 10.1098/rspb.2018.2076, 30963947, 6364590, free, {hide}clade|label1=Dictyoptera|1={{clade
|label1=Mantodea
|1= (Mantises)
|label2=Blattodea
|2= (Cockroaches and termites)
{edih}
}}

Internal

One of the earliest classifications splitting an all-inclusive Mantidae into multiple families was that proposed by Beier in 1968, recognizing eight families,JOURNAL, Beier, M., 1968, Ordnung Mantodea (Fangheuschrecken), Handbuch der Zoologie, 4, 2, 3–12, 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.BOOK, Klass, Klaus-Dieter, The external male genitalia and phylogeny of Blattaria and Mantodea,weblink 1997, Zoologisches Forschungsinstitut, 978-3-925382-45-1, However, as previously configured, the Mantidae and Thespidae especially were considered polyphyletic,BOOK, Martill, David M., Bechly, Günter, Loveridge, Robert F., The Crato Fossil Beds of Brazil: Window into an Ancient World,weblink 2007, Cambridge University Press, 978-1-139-46776-6, 236–238, so the Mantodea have been revised substantially as of 2019 and now includes 29 families.Schwarz CJ, Roy R (2019) The systematics of Mantodea revisited: an updated classification incorporating multiple data sources (Insecta: Dictyoptera) Annales de la Société entomologique de France (N.S.) International Journal of Entomology 55 [2]: 101–196.{{cladogram|align=left
Extant taxon>extant Mantodea familiesGRIMALDI >FIRST1=DAVID TITLE=A REVISION OF CRETACEOUS MANTISES AND THEIR RELATIONSHIPS, INCLUDING NEW TAXA (INSECTA: DICTYOPTERA: MANTODEA) VOLUME= HDL=2246/2838 DOI=10.1206/0003-0082(2003)4122.0.CO;2 URL=HTTPS://WWW.BIODIVERSITYLIBRARY.ORG/BIBLIOGRAPHY/178207, style=font-size:80%;|label1=Mantodea|1={{clade
|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={{clade
|label1=Thespoidea
|1=Thespidae
|label2=Acanthopoidea
|2={{clade
|1=Angelidae
|2={{clade
|1=Coptopterygidae
|2={{clade
|1=Liturgusidae
|2={{clade
|1=Photinaidae
|2=Acanthopidae
{edih}
}}
}}
}} }}
|label2=Cernomantodea
|2={{clade
|label1=Nanomantodea
|1={{clade
|label1=Chroicopteroidea
|1=Chroicopteridae
|label2=Nanomantoidea
|2={{clade
|1=Leptomantellidae
|2={{clade
|1=Amorphoscelidae
|2=Nanomantidae
}}
}} }}
|label2=Metamantodea
|2={SUBCLADE_Metamantodea}
}}
}}
}}
}}
}} }}
|targetA={SUBCLADE_Metamantodea}
|subcladeA={{Clade
|1={{clade
|label1=Gonypetoidea
|1=Gonypetidae
|label2=Lobipedia
|2={{clade
|label1=Epaphroditoidea
|1={{clade
|1=Epaphroditidae
|2=Majangidae
}}
|label2=Mantimorpha
|2={{clade
|label1=Haanioidea
|1=Haaniidae
|label2=Heteromantodea
|2={{clade
|label1=Eremiaphiloidea
|1={{clade
|1=Rivetinidae
|2={{clade
|1=Amelidae
|2={{clade
|1=Eremiaphilidae
|2=Toxoderidae
}}
}} }}
|label2=Pareumantodea
|2={{clade
|label1=Hoplocoryphoidea
|1=Hoplocoryphidae
|label2=Calomantodea
|2={{clade
|label1=Miomantoidea
|1=Miomantidae
|label2=Promantidea
|2={{clade
|label1=Galinthiadoidea
|1=Galinthiadidae
|label2=Mantidea
|2={{clade
|label1=Hymenopodoidea
|1={{clade
|1=Empusidae
|2=Hymenopodidae
}}
|label2=Mantoidea
|2={{clade
|1=Dactylopterygidae
|2={{clade
|1=Deroplatyidae
|2=Mantidae
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}} }}
}{{See also|List of mantis genera and species}}

Fossil mantises

File:Santanmantis reconstruction.png|thumb|Life restoration of SantanmantisSantanmantisMantises are thought to have evolved from cockroach-like ancestors.JOURNAL, Li, Xinran, 2019-08-30, Disambiguating the scientific names of cockroaches,weblink Palaeoentomology, 2, 4, 390–402, 10.11646/palaeoentomology.2.4.13, 202789239, 2624-2834, The earliest confidently identified mantis fossils date to the Early Cretaceous. Fossils of the group are rare: by 2022, 37 fossil species are known.JOURNAL, Terríquez-Beltrán, J., Riquelme, F., Varela-Hernández, F., 2022-10-31, A new species of mantis (Insecta: Mantodea: Amelidae) from the Miocene Amber-Lagerstätte in Mexico,weblink Historical Biology, en, 1–8, 10.1080/08912963.2022.2134782, 253280592, 0891-2963, Fossil mantises, including one from Japan with spines on the front legs as in modern mantises, have been found in Cretaceous amber.WEB, Ryall, Julian, Ancient Praying Mantis Found in Amber,weblinkweblink" title="web.archive.org/web/20080429193405weblink">weblink dead, 29 April 2008, National Geographic Society, 30 July 2015, 25 April 2008, Most fossils in amber are nymphs; compression fossils (in rock) include adults. Fossil mantises from the Crato Formation in Brazil include the {{convert|10|mm|in|adj=on|abbr=on}} 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:{{div col|colwidth=22em}} {{div col end}}

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 (leathery forewings) 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.JOURNAL, Boyden, Thomas C., Mimicry, predation and potential pollination by the mantispid, Climaciella brunnea var. instabilis (Say) (Mantispidae: Neuroptera), Journal of the New York Entomological Society, 1983, 91, 4, 508–511, 25009393,

Biology

Anatomy

{{Multiple image|total_width=330|image1=Raptrix perspicua MHNT male.jpg|alt1=Mantis wings, the forewing leathery, the hindwing triangular|caption1=Wing arrangement of a typical mantis, adult male Raptrix perspicua|image2=MantisLegGBMNH.jpg|alt2=Raptorial foreleg of a mantis, armed with long spinesAnatomical terms of location>proximal segment of the grasping part of the leg.}}(File:Mantis - Kanagawa - 2023 Sept 11.webm|thumb|Mantis moving on a wall.)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.BOOK, Siwanowicz, Igor, 2009, Animals Up Close,weblink Dorling Kindersley, 38–39, -978-1-405-33731-1, 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.JOURNAL, Yager, David, Svenson, Gavin, Patterns of praying mantis auditory system evolution based on morphological, molecular, neurophysiological, and behavioural data., Biological Journal of the Linnean Society, 1 July 2008, 94, 3, 541–568, 10.1111/j.1095-8312.2008.00996.x, Oxford University Press, en, free, 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.BOOK, The Praying Mantids, Prete, Fredrick R., Wells, Harrington, Wells, Patrick H., Hurd, Lawrence E., Morphology and Taxonomy, Roy, Roger, 1999, 21–33, Johns Hopkins University Press, 978-0-8018-6174-1,weblink 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 and leathery. They function as camouflage and as a shield for the hindwings, which are clearer and more delicate.WEB, Kemper, William T., Insect Order ID: Mantodea (Praying Mantises, Mantids),weblink Missouri Botanical Garden, 21 September 2015, 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

File:Large brown mantid close up nohair.jpg|thumb|left|Head of Archimantis latistyla, showing the compound eyes and labrumlabrumMantises have stereo vision.JOURNAL, 10.1038/srep18718, 26740144, 4703989, Insect stereopsis demonstrated using a 3D insect cinema, Scientific Reports, 6, 18718, 2016, Nityananda, Vivek, Tarawneh, Ghaith, Rosner, Ronny, Nicolas, Judith, Crichton, Stuart, Read, Jenny, 2016NatSR...618718N, JOURNAL, Rossel, S, 1983, Binocular stereopsis in an insect, Nature, 302, 5911, 821–822, 10.1038/302821a0, 1983Natur.302..821R, 4264254, JOURNAL, 13229–13232, 24075, 1996, Rossel, S, Binocular vision in insects: How mantids solve the correspondence problem, Proceedings of the National Academy of Sciences of the United States of America, 93, 23, 11038523, 10.1073/pnas.93.23.13229, 1996PNAS...9313229R, free, 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.JOURNAL, Corrette, Brian J., Prey capture in the praying mantis Tenodera aridifolia sinensis: coordination of the capture sequence and strike movements, Journal of Experimental Biology, 1990, 148, 147–180, 10.1242/jeb.148.1.147,weblink 2407798, BOOK, Simmons, Peter J., Young, David, Nerve Cells and Animal Behaviour,weblink 1999, Cambridge University Press, 978-0-521-62726-9, 8990, The eyes are widely spaced and laterally situated, affording a wide binocular field of vision and precise stereoscopic vision at close range.BOOK, Howard, Ian P., Rogers, Brian J., Binocular Vision and Stereopsis,weblink 1995, Oxford University Press, 978-0-19-508476-4, 646, 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.JOURNAL, Zeil, Jochen, Al-Mutairi, Maha M., 1996, Variations in the optical properties of the compound eyes of Uca lactea annulipes, The Journal of Experimental Biology, 199, 7, 1569–1577, 10.1242/jeb.199.7.1569,weblink 9319471, 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. They have good night vision.WEB, Robert, 2023-07-05, Can Praying Mantis See in the Dark?,weblink 2024-02-16, Insects Authority, en-US, Mantises in the family Liturgusidae collected at night have been shown to be predominately males;JOURNAL, Bragg, P.E., 2010, A review of the Liturgusidae of Borneo (Insecta: Mantodea), Sepilok Bulletin, 12, 21–36, this is probably true for most mantises. 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.BOOK, The Praying Mantids, Prete, Fredrick R., Wells, Harrington, Wells, Patrick H., Hurd, Lawrence E., 1999, Hearing, Yager, David, 101–103, Johns Hopkins University Press, 978-0-8018-6174-1,weblink BOOK, Grimaldi, David, Engel, Michael S., 2005, Evolution of the Insects,weblink limited, Cambridge University Press, 257, 978-0-521-82149-0,

Diet and hunting

File:Tenodera sinensis 2 Luc Viatour.jpg|thumb|right|Tenodera sinensis feeding on a cricketcricketFile:Pm eating pm.webm|thumb|right|thumbtime=46|Tenodera sinensisTenodera sinensisMantises 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.JOURNAL, Mantids – The Praying Predators, National Geographic (magazine), National Geographic, Edward S., Ross, 268–280, 165, 2, February 1984, 0027-9358, 643483454, Larger mantises sometimes eat smaller individuals of their own species,BOOK, Capinera, John L., Encyclopedia of Entomology,weblink 2008, Springer, 978-1-4020-6242-1, 1509, as well as small vertebrates such as lizards, frogs, fish, and particularly small birds.BOOK, Grimaldi, David, Engel, Michael S., Evolution of the Insects,weblink 2005, Cambridge University Press, 978-0-521-82149-0, 254, JOURNAL, Nyffeler, Martin, Maxwell, Michael R., Remsen, J. V. Jr., Bird predation by praying mantises: a global perspective, The Wilson Journal of Ornithology, 2017, 129, 2, 331–344, 10.1676/16-100.1, 90832425, JOURNAL, Battison, Roberto, etal, The fishing mantid: predation on fish as a new adaptive strategy for praying mantids (Insecta: Mantodea), Journal of Orthoptera Research, 27, 2, 155–158, 10.3897/jor.27.28067,weblink 2018, free, Most mantises stalk tempting prey if it strays close enough, and will go further when they are especially hungry.JOURNAL, Gelperin, Alan, Feeding behaviour of the praying mantis: a learned modification, Nature, July 1968, 219, 5152, 399–400, 10.1038/219399a0, 5667084, 1968Natur.219..399G, 10951932, Once within reach, mantises strike rapidly to grasp the prey with their spiked raptorial forelegs.JOURNAL, Prete, F. R., Cleal, K. S., The predatory strike of free ranging praying mantises, Sphodromantis lineola (Burmeister). I: Strikes in the mid-sagittal plane, Brain, Behavior and Evolution, 1996, 48, 4, 173–190, 8886389, 10.1159/000113196, 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 fore gut 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.BOOK, Capinera, John L., Encyclopedia of Entomology,weblink 2008, Springer, 978-1-4020-6242-1, 116, Chinese mantises live longer, grow faster, and produce more young when they are able to eat pollen.JOURNAL, 10.1603/0046-225X-32.4.881, Pollen feeding and fitness in praying mantids: the vegetarian side of a tritrophic predator, Environmental Entomology, 32, 4, 881–885, 2003, Beckman, Noelle, Hurd, Lawrence E., free,

Antipredator adaptations

{{further|Flower mantis}}Mantises are preyed on by vertebrates such as frogs, lizards, and birds, and by invertebrates such as spiders, large species of hornets, and ants.JOURNAL, Iyer, Geetha, Down to earth,weblink Frontline Magazine, 28, 17, 13 August 2011, Some hunting wasps, such as some species of Tachytes also paralyze some species of mantis to feed their young.BOOK, Jean-Henri Fabre, More Hunting Wasps,weblink 1921, Dodd, Mead, Generally, mantises protect themselves by camouflage, most species being cryptically colored to resemble foliage or other backgrounds, both to avoid predators and to better snare their prey.BOOK, Gullan, P. J., Cranston, P. S., 2010, The Insects: An Outline of Entomology,weblink limited, Wiley, 4th, 978-1-118-84616-2, 370, 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.BOOK, The Praying Mantids, Prete, Fredrick R., Wells, Harrington, Wells, Patrick H., Hurd, Lawrence E., 1999, 282–293, Edmunds, Malcolm, Brunner, Dani, Ethology of Defenses against Predators, Johns Hopkins University Press, 978-0-8018-6174-1,weblink 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.BOOK, Cott, Hugh, Adaptive Coloration in Animals, Methuen, 1940, 392–393, Hugh Cott, Adaptive Coloration in Animals, JOURNAL, Annandale, Nelson, Observations on the habits and natural surroundings of insects made during the 'Skeat Expedition' to the Malay Peninsula, 1899–1900, Proceedings of the Zoological Society of London, 69, 1900, 862–865, JOURNAL, O'Hanlon, James C., Holwell, Gregory I., Herberstein, Marie E., Pollinator deception in the orchid mantis, The American Naturalist, 2014, 183, 1, 126–132, 10.1086/673858, 24334741, 2228423, free, 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).File:Mantis Hymenopus coronatus 2 Luc Viatour.jpg|thumb|upright|Aggressive mimicry: Malaysian orchid mantises are camouflaged pink or yellow, matching the coloration of local orchidorchidFile:Motion Camouflage.webm|thumb|right|thumbtime=2|Praying mantises exhibiting motion camouflagemotion camouflageWhen 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 enhancing this effect with bright colors and patterns on their hindwings and inner surfaces of their front legs. 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 may also produce a hissing sound by expelling air from the abdominal spiracles. Mantises lack chemical protection, so their displays are largely bluff. When flying at night, at least some mantises are able to detect the echolocation sounds produced by bats; 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. If caught, they may slash captors with their raptorial legs.JOURNAL, Yager, D., May, M., 1993, Coming in on a wing and an ear, Natural History, 102, 1, 28–33, WEB,weblinkweblink" title="web.archive.org/web/20060521084215weblink">weblink dead, 21 May 2006, Praying Mantis Uses Ultrasonic Hearing to Dodge Bats, 19 November 2002, National Geographic Society, 17 August 2012, 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.JOURNAL, O'Dea, J. D., Eine zusatzliche oder alternative Funktion der 'kryptischen' Schaukelbewegung bei Gottesanbeterinnen und Stabschrecken (Mantodea, Phasmatodea), Entomologische Zeitschrift, 101, 1–2, 1991, 25–27, 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.JOURNAL, Nelson, Ximena J., Jackson, Robert R., 2006, Innate aversion to ants (Hymenoptera: Formicidae) and ant mimics: experimental findings from mantises (Mantodea), Biological Journal of the Linnean Society, 88, 1, 23–32, 10.1111/j.1095-8312.2006.00598.x, free, Choeradodis stalii MHNT dos.jpg|Leaf mimicry: Choeradodis has leaf-like forewings and a widened green thorax.Gottesanbeterin Abwehr.JPG|Adult female Iris oratoria performs a bluffing threat display, rearing back with the forelegs and wings spread and mouth opened.Gemmatus.jpg|The jeweled flower mantis, Creobroter gemmatus: the brightly colored wings are opened suddenly in a deimatic display to startle predators.Ant-Mantis.jpg|Some mantis nymphs mimic ants to avoid predators.

Reproduction and life history

(File:Mantases hatching.webm|thumb|right|thumbtime=24|Praying mantis nymphs emerging from their ootheca.)The mating season in temperate climates typically takes place in autumn, while in tropical areas, mating can occur at any time of the year.BOOK, Prete, Fredrick R., Wells, Harrington, Wells, Patrick H., Hurd, Lawrence E., The Praying Mantids, Mating Behavior, Michael, Maxwell. R., 1999, Johns Hopkins University Press, 978-0-8018-6174-1, 70,weblink 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. 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 mantises in the family Tarachodidae, the mother guards the eggs.JOURNAL, 10.1080/00222936408651488, Ene, J. C., 1964, The distribution and post-embryonic development of Tarachodes afzelli (Stal) (Mantodea : Eremiaphilidae), Journal of Natural History, 7, 80, 493–511, 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. An unusual 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. The ability to reproduce by parthenogenesis has been recorded in at least two other species, Sphodromantis viridis and Miomantis sp., although these species usually reproduce sexually.Bragg, P.E. (1987) A case of parthenogenesis in a mantis. Bulletin of the Amateur Entomologists' Society, 46 (356): 160.JOURNAL, Bragg, P. E., Parthenogenetic mantid named, Bulletin of the Amateur Entomologists' Society,weblink Mantis Study Group, 8 January 2017, December 1989, 48, 367, 242, Dickie, S. (1996) Parthenogenesis in mantids. Mantis Study Group Newsletter, 1: 5. In temperate climates, adults do not survive the winter and the eggs undergo a diapause, hatching in the spring.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). For smaller species, the eggs may hatch in 3–4 weeks as opposed to 4–6 weeks for larger species. The nymphs may be colored differently from the adult, and the early stages are often mimics of ants. A mantis nymph grows bigger as it molts its exoskeleton. Molting can happen five to 10 times before the adult stage is reached, depending on the species. After the final molt, most species have wings, though some species remain wingless or brachypterous ("short-winged"), particularly in the female sex. The lifespan of a mantis depends on the species; smaller ones may live 4–8 weeks, while larger species may live 4–6 months.BOOK, Prete, Fredrick R., Wells, Harrington, Wells, Patrick H., Hurd, Lawrence E., The Praying Mantids, Ecology of Praying Mantids, Hurd, I. E.,weblink 1999, Johns Hopkins University Press, 978-0-8018-6174-1, 43–49, File:Mantis religiosa couple.JPG|Mantis religiosa mating (brown male, green female)File:Mantis laying ootheca.jpg|Stagmomantis carolina laying oothecaFile:Mantis religiosa (egg case).jpg|Recently laid M. religiosa oothecaFile:PrayingMantisesJustBorn (cropped).jpg|Hatching from the oothecaFile:Sphodromantis Lineola molting.jpg|Sphodromantis lineola molting

Sexual cannibalism

{{further|Sexual cannibalism}}(File:Praying Mantis Sexual Cannibalism European-37.jpg|thumb|Sexual cannibalism in Mantis religiosa)Sexual cannibalism is common among most predatory species of mantises in captivity. It has sometimes been observed in natural populations, where about a quarter of male–female encounters result in the male being eaten by the female.JOURNAL, Lawrence, S. E., Sexual cannibalism in the praying mantid, Mantis religiosa: a field study, Animal Behaviour, 1992, 43, 4, 569–583, 10.1016/S0003-3472(05)81017-6, 53199427, JOURNAL, Hurd, L. E., Eisenberg, R. M., Fagan, W. F., Tilmon, K. J., Snyder, W. E., Vandersall, K. S., Datz, S. G., Welch, J. D., Cannibalism reverses male-biased sex ratio in adult mantids: female strategy against food limitation?, Oikos, 1994, 69, 2, 193–198, 10.2307/3546137, 3546137, JOURNAL, Maxwell, Michael R., Lifetime mating opportunities and male mating behaviour in sexually cannibalistic praying mantids, Animal Behaviour, 55, 4, 1011–1028, 1998, 10.1006/anbe.1997.0671, 9632486, 26887566, Around 90% of the predatory species of mantises exhibit sexual cannibalism.JOURNAL, 10.1146/annurev.ecolsys.110308.120238, The importance of ecological and phylogenetic conditions for the occurrence and frequency of sexual cannibalism, Annual Review of Ecology, Evolution, and Systematics, 40, 21–39, 2009, Wilder, Shawn M., Rypstra, Ann L., Elgar, Mark A., 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.WEB, Do Female Praying Mantises Always Eat the Males?,weblink Entomology Today, 31 July 2015, 22 December 2013, 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.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 is natural in the field or also the result of distractions caused by the human observer remains controversial. Mantises are highly visual organisms and notice any disturbance in the laboratory or field, such as bright lights or moving scientists. 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.JOURNAL, 10.1016/S0003-3472(84)80170-0, Sexual behaviour of the Chinese praying mantis, Animal Behaviour, 32, 3, 916–918, 1984, Liske, E., Davis, W. J., 53144893, 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.JOURNAL, Lelito, Jonathan P., Brown, William D., 2006, Complicity or conflict over sexual cannibalism? Male risk taking in the praying mantis Tenodera aridifolia sinensis, The American Naturalist, 168, 2, 263–269, 10.1086/505757, 16874635, 32185318, 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.JOURNAL, 10.1016/s0003-3472(87)80024-6, Courtship and mating behaviour of the Chinese praying mantis, Tenodera aridifolia sinenesis, Animal Behaviour, 35, 5, 1524–1537, 1987, Liske, E., Davis, W. J., 53191218, 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.JOURNAL, 10.1111/j.1365-2311.2010.01239.x, Effects of female feeding regime in a sexually cannibalistic mantid: fecundity, cannibalism, and male response in Stagmomantis limbata (Mantodea), Ecological Entomology, 35, 6, 775–787, 2010, Maxwell, Michael R., Gallego, Kevin M., Barry, Katherine L., 85923330, 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. Experiments have revealed that the sex ratio in an environment determines male copulatory behavior of Mantis religiosa which in turn affects the cannibalistic tendencies of the female and support the sperm competition hypothesis because the polyandrous treatment recorded the highest copulation duration time and lowest cannibalism. This further suggests that dismounting the female can make males susceptible to cannibalism.JOURNAL, Prokop, P, Václav, R, Males Respond to the Risk of Sperm Competition in the Sexually Cannibalistic Praying Mantis, Mantis religiosa, Ethology, 2005, 111, 9, 836–848, 10.1111/j.1439-0310.2005.01113.x,

Relationship with humans

In culture, literature and art

File:Murata Seimin - Brush Rest in the Shape of a Praying Mantis - Walters 541323.jpg|thumb|left|Bronze ink brush rest in the shape of a mantis, Edo periodEdo periodOne 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 {{ill|Jingshi Zhenglei Daguan Bencao|zh|证类本草|italic=1}} ({{translation|"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 fifth-century BC silver coins, including didrachms, from Metapontum in Lucania.BOOK, Bodson, Liliane, Campbell, Gordon Lindsay, The Oxford Handbook of Animals in Classical Thought and Life,weblink 2014, Oxford University Press, 978-0-19-958942-5, 557–558, In the 10th century AD, Byzantine era Adages, Suidas describes an insect resembling a slow-moving green locust with long front legs.WEB, Entomomancy,weblink Occultopedia, California Astrology Association, 30 July 2015, 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.BOOK, Erasmus, Desiderius, Fantazzi, Charles, Adages: Iivii1 to Iiiiii100,weblink 1992, University of Toronto Press, 978-0-8020-2831-0, 334–335, 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".JOURNAL, McCafferty, Geoffrey, McCafferty, Sharisse, 1 April 2020, Praying to the Predator: Symbols of Insect Animism on Luna/El Menco Polychrome from Pre-Columbian Pacific Nicaragua, 10.1525/lavc.2020.220004, Latin American and Latinx Visual Culture, 2, 2, 28–46, 225959388, 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).File:Wandering Violin Mantis (Gongylus gongylodes) Photograph By Shantanu Kuveskar.jpg|thumb|upright|In Island, Aldous Huxley reflected on death as a pair of alt=The thin-legged mantis Gongylus gongylodesIn the early 1900s, people in the United States Ozarks region referred to them as Devil's horses.BOOK, Randolph, Vance,weblink Ozark Magic and Folklore, 2012-07-31, Courier Corporation, 978-0-486-12296-0, en, 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:{{blockquote|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 hindlegs. They rustled and staggered across the ceiling and down the wall, each seeking to gain some advantage.BOOK, Durrell, Gerald, My Family and Other Animals,weblink 2006, 1956, Penguin Books, 978-0-14-193609-3, 204–208, }}M. C. Escher's woodcut Dream depicts a human-sized mantis standing on a sleeping bishop.BOOK, Emmer, Michele, Schattschneider, Doris, M.C. Escher's Legacy: A Centennial Celebration,weblink 2007, Springer, 978-3-540-28849-7, 66, 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.WEB, Wade, Lisa, Shoddy Research and Cultural Tropes: The Praying Mantis,weblink The Society Pages, 9 October 2015, 29 November 2010, WEB, Praying Mantises Cartoons and Comics,weblink CartoonStock, 30 July 2015, WEB, Parisi, Mark,weblinkkey/praying+mantis/, Praying Mantis Cartoons, Off the Mark, 30 July 2015, dead, https:web.archive.org/web/20150925144024weblink 25 September 2015, WEB, McCracken, T., Praying Mantis Should Be Gay,weblink McHumor Cartoons, 30 July 2015, 5 March 2016,weblink dead, It ends Isabella Rossellini's short film about the life of a praying mantis in her 2008 Green Porno season for the Sundance Channel.WEB, Rossellini, Isabella, Isabella Rossellini, Green Porno,weblinkweblink 15 December 2021, live, Sundance Channel, 1 September 2015, 2008, {{cbignore}}WEB, Rossellini, Isabella, Green Porno: a series of short films by Isabella Rossellini, Sundance Channel, 2009,weblink Press Kit, 1 September 2015, The Deadly Mantis is a 1957 American science fiction monster film, with a giant mantis threatening mankind.Imdb

Martial arts

File:Shi DeRu and Shi DeYang.jpg|thumb|Grandmasters of the Shaolin Temple, Shi DeRu and Shi DeYang, demonstrating the Southern Praying MantisSouthern Praying MantisTwo martial arts separately developed in China have movements and fighting strategies based on those of the mantis.WEB, History of Praying Mantis Kung Fu,weblinkweblink" title="web.archive.org/web/20151001110604weblink">weblink 1 October 2015, Praying Mantis Kung Fu Academy, 30 September 2015, As one of these arts was developed in northern China, and the other in southern parts of the country, the arts are today referred to (both in English and Chinese) as 'Northern Praying Mantis'WEB, Funk, Jon, Praying Mantis Kung Fu,weblink Praying Mantis Kung Fu, 31 July 2015, and 'Southern Praying Mantis'.BOOK, Hagood, Roger D., 18 Buddha Hands: Southern Praying Mantis Kung Fu, 2012, Southern Mantis Press, 978-0-9857240-1-6, Both are very popular in China, and have also been exported to the West in recent decades.WEB, Barnes, Bryan, Northern Praying Mantis Kung Fu,weblink Shantung Northern Praying Mantis Kung Fu, 30 September 2015, WEB, Orum, Pete, Chow Gar,weblink Southern Mantis Kung Fu, 30 September 2015,

In mythology and religion

According to local beliefs in Africa, this insect brings good luck.Is a Praying Mantis Good Luck?, Reference.com, 11 April 2020 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.WEB, Insek-kaleidoskoop: Die 'skynheilige' hottentotsgot,weblink Mieliestronk.com, 9 October 2015, 24 May 2019,weblink" title="web.archive.org/web/20190524230352weblink">weblink dead, 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.WEB, Siyabona Africa: Kruger National Park: San,weblink Siyabona Africa, 30 June 2017, 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).BOOK, Prete, Fredrick R., Wells, Harrington, Wells, Patrick H., Hurd, Lawrence E., The Praying Mantids, The Predatory Behavior of Mantids: Historical Attitudes and Contemporary Questions, Prete, Frederick R., Wells, Harrington, Wells, Patrick H., amp,weblink 1999, Johns Hopkins University Press, 978-0-8018-6174-1, 3–8, ENCYCLOPEDIA, Mantid,weblink Encyclopædia Britannica, 30 July 2015, 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.

As pets

File:Gray adult female Carolina Mantis.JPG|thumb|Gray adult female Carolina mantisCarolina mantisMantises are among the insects most widely kept as pets.WEB, Pet bugs: Kentucky arthropods,weblink University of Kentucky, 31 July 2015, WEB, Van Zomeren, Linda, European Mantis, 16 March 2011,weblink Keeping Insects, 31 July 2015, 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.REPORT, Breeding Reports: 1st July 2013–1st October 2013, Newsletter,weblink UK Mantis Forums, UK Mantis Forums Newsletter, 12, 31 July 2015, October 2013, dead,weblink" title="web.archive.org/web/20150923103205weblink">weblink 23 September 2015, In 1996 at least 50 species were known to be kept in captivity by members of the Mantis Study Group.Bragg, P.E. [editor] (1996) Species in culture. Mantis Study Group Newsletter, 1: 2–3.

For pest control

Naturally occurring mantis populations provide plant pest control.JOURNAL, Zhang, Wei, Ricketts, Taylor H., Kremen, Claire, Carney, Karen, Swinton, Scott M., Ecosystem services and dis-services to agriculture, Ecological Economics (journal), Ecological Economics, International Society for Ecological Economics (Elsevier), 64, 2, 2007, 0921-8009, 10.1016/j.ecolecon.2007.02.024, 253–260, 32112906,weblink Gardeners who prefer to avoid pesticides may encourage mantises in the hope of controlling insect pests.WEB, Doutt, R. L., The Praying Mantis (Leaflet 21019),weblink University of California Division of Agricultural Sciences, 31 July 2015, 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.WEB, Praying and Chinese Mantises,weblink New York State Department of Environmental Conservation, 26 August 2015,

Robotics

{{further|Biomimicry}}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.WEB, Cardona, Ramon, Touretzky, David, Leg Design for a Praying Mantis Robot,weblink Association for the Advancement of Artificial Intelligence, Florida Artificial Intelligence Research Society, 12 April 2016, 24 June 2016,weblink dead,

See also

{{Clear}}

References

{{Reflist|28em}}

External links

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