Editing European mantis

{{Short description|Species of praying mantis}}
{{Speciesbox
| name = European mantis
| fossil_range = Piacenzian – Recent<ref>{{cite web | url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=306570 | title=Fossilworks: Mantis religiosa }}</ref>
| image = European praying mantis (Mantis religiosa) green female Dobruja.jpg
| image_caption = Adult female, [[Romania]]
| status = LC
| status_system = IUCN3.1
| status_ref = <ref name="iucn">{{Cite iucn| author= Battiston, R. |year= 2016 |title= ''Mantis religiosa'' |page= e.T44793247A44798476 |doi=10.2305/IUCN.UK.2016-1.RLTS.T44793247A44798476.en |access-date=16 November 2023}}</ref>
| genus = Mantis
| species = religiosa
| authority = ([[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]])
| display_parents = 4
| subdivision_ranks = Subspecies
| subdivision = [[#Subspecies|See text]]
| synonyms = *''Gryllus religiosus'' <small>[[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]</small>
}}

The '''European mantis''' ('''''Mantis religiosa''''') is a large [[Hemimetabolism|hemimetabolic]] insect in the family of the [[Mantidae]] ('mantids'), which is the largest family of the order [[Mantodea]] (mantises).<ref name="KralOrientationBehavior">{{cite journal |last=Kral |first=Karl |date=March 2014 |title=Orientation Behavior with and Without Visual Cues in Newly Hatched and Adult Praying Mantis |journal=Journal of Insect Behavior |volume=27 |issue=2 |pages=192–205 |doi=10.1007/s10905-013-9415-3|bibcode=2014JIBeh..27..192K |s2cid=17222572 }}</ref> Their common name '''praying mantis''' is derived from the distinctive posture of the first pair of legs that can be observed in animals in repose. It resembles a [[praying]] attitude.<ref name="Roeder1935">{{cite journal |last1=Roeder |first1=Kenneth D. |date=October 1935 |title=An experimental analysis of the sexual behavior of the praying mantis (Mantis religiosa L.) |journal=The Biological Bulletin  |volume=69 |issue=2 |pages=203–220 |doi=10.2307/1537420|jstor=1537420 }}</ref> Both males and females have elongated bodies with two pairs of wings. The most striking features that all Mantodea share are a very mobile, triangular head with large [[compound eyes]] and their first pair of legs (the '[[raptorial]] legs'), which is highly modified for the efficient capture and restraint of fast-moving or flying prey.<ref name="Roeder1935" />

In [[Germany]], ''M. religiosa'' is listed as ''Gefährdet'' [endangered] on the German [[Red List]] on the basis of an assessment from 1998.<ref>{{cite web |url=https://www.nationalredlist.org/species-information/?speciesID=248765 |title = Species Information {{!}} Regional Red List}}</ref> It is not supposed to be caught or held as a pet.<ref name="ReferenceA">{{cite journal|last1=Linn|first1=Catherine Anne|last2=Griebeler|first2=Eva Maria|title=Habitat Preference of German Mantis religiosa Populations (Mantodea: Mantidae) and Implications for Conservation|journal=Environmental Entomology|date=August 2016|volume=45|issue=4|pages=829–840|doi=10.1093/ee/nvw056|pmid=27271947}}</ref> At a global level, it is assessed by the IUCN as [[least concern]].<ref name="iucn"/>

[[File:Linnaeus 1758 Systema Naturae - Descriptions of mantises.jpg|thumb|right|Copy of the original description of several species of mantis including ''Mantis religiosa'', described by [[Carl Linnaeus]] as ''Gryllus (Mantis) religiosus'' in 1758.]]

==Anatomy==
Though females of  ''M. religiosa'' are usually larger and heavier than males (7–9&nbsp;cm versus 6–7&nbsp;cm), the [[antenna (biology)|antenna]]e and the eyes of male animals outsize those of the females. Along with the forward-directed, compound eyes, simple eyes also are found on the head. These three dorsal [[ocelli]] are also more pronounced in males than in females.<ref name="KralOrientationBehavior" />

Males are often found to be more active and agile, whereas females are physically more powerful.<ref name="Roeder1935" />  Adult females are generally too large and heavy for their wings to enable a take-off.<ref name="KralOrientationBehavior" /><ref name="Lawrence">{{cite journal |last1=Lawrence |first1=S. E. |date=April 1992 |title=Sexual cannibalism in the praying mantid, Mantis religiosa: a field study |journal=Animal Behaviour |volume=43 |issue=4 |pages=569–583 |doi=10.1016/S0003-3472(05)81017-6|s2cid=53199427 }}</ref>

Mantids show strong [[deimatic display]] from very early life stages on.<ref name="Przibram1907">{{cite journal |last1=Przibram |first1=Hans |date=June 1907 |title=Aufzucht, Farbwechsel und Regeneration unsrer europäischen Gottesanbeterin (Mantis religiosa L.) |journal=Archiv für Entwicklungsmechanik der Organismen |volume=23 |issue=4 |pages=600–614 |language=de |doi=10.1007/BF02288315|s2cid=7354286 }}</ref> This behavior can be observed throughout different groups of animals and is used to scare or startle potential predators to give the attacked animals a chance to escape.<ref>{{cite journal|last1=Umbers|first1=Kate D.L.|last2=Lehtonen|first2=Jussi|last3=Mappes|first3=Johanna|title=Deimatic displays|journal=Current Biology|date=January 2015|volume=25|issue=2|pages=R58–R59|doi=10.1016/j.cub.2014.11.011|pmid=25602301|doi-access=free|bibcode=2015CBio...25..R58U }}</ref> The deimatic display in ''M. religiosa'' involves wing spreading and bending of the raptorial legs to reveal two matched black eyespots with a yellow or white center at the base of the [[Arthropod coxa|coxa]]e (legs). It makes the animal appear much larger and more of a threat to the attacker. The black eyespots are also a distinct feature for species discrimination of the European mantis.<ref>{{cite journal|last1=Sureshan|first1=P.M.|last2=Samanta|first2=T|last3=Radhakrishnan|first3=C|title=Mantid (Insecta: Mantodea) fauna of Orissa with some new records for the state.|journal=Zoos' Print Journal|date=2007|volume=22|issue=1|pages=2539–2543|doi=10.11609/jott.zpj.1524.2539-43|doi-access=free}}</ref> 
Another unique feature of ''M. religiosa'' is its midline metathoracic ear (see [[Ultrasound avoidance]]). This “tympanal [[auditory system|auditory]] organ” is an unpaired structure found on the [[ventral]] side of the animal on the metathorax between the third pair of legs. Unlike other sound-processing organs found among different groups of insects, the metathoracic ear has a high sensitivity across high and low frequencies and even [[ultrasound]]. Therefore, the authors call it a ‘true ear’.<ref>{{cite journal|last1=Yager|first1=David|last2=Hoy|first2=Ronald|title=The midline metathoracic ear of the praying mantis, Mantis religiosa|journal=Cell and Tissue Research|date=December 1987|volume=250|issue=3|pages=531–541|doi=10.1007/BF00218944|pmid=3690633|s2cid=37360392}}</ref>

==Coloration==
The great variation in the coloration of ''M. religiosa'' from different shades of yellow, brown, green, and sometimes black has been the cause of numerous hypotheses and studies for over 100 years.<ref>{{cite journal |last1=Ergene |first1=Saadet |title=Homochrome Farbanpassungen bei ''Mantis religiosa'' |trans-title=Homochromic color adjustments in Mantis religiosa |language=de |journal=Zeitschrift für Vergleichende Physiologie |date=1953 |volume=35 |issue=1–2 |pages=36–41 |doi=10.1007/BF00340699 |s2cid=2923351 }}</ref> However, no generally accepted answer about reason, benefit, or mechanism of the coloration or the change of coloration has been found.

[[Alessandro Palma di Cesnola|Di Cesnola]] observed in 1904 that green mantids were found on fresh green grass, whereas brown individuals seemed to prefer brown grass burned by the sun. When forced to change the location as to no longer match their coloration, almost all the ‘not matching’ animals were killed by predators such as birds. This indicates a [[camouflaging]] purpose of the coloration.<ref name="ReferenceB">{{cite journal |last1=Di Cesnola |first1=A. P. |title=Preliminary note on the protective value of colour in Mantis religiosa |journal=Biometrika |date=1904 |volume=3 |issue=1 |pages=58–9 |doi=10.1093/biomet/3.1.58 |url=https://zenodo.org/record/1431603 }}</ref>

Przibram also observed in 1907 that a change in temperature can trigger a change in coloration:<ref name="Przibram1907" /> animals that hatched in a cold environment turned green after moulting when heat and sunlight were provided. Without the change in temperature and only a change in the color of the background, no change in coloration occurred.<ref name="Przibram1907" /> This finding contradicts with Di Cesnola, who claims to have observed the animals within the same time and location (and therefore the same temperature).<ref name="ReferenceB"/>

[[File:Brown Mantis.jpg|thumb|right|alt=Brown European mantis|Brown European mantis]]

==Distribution==
''M. religiosa'' can be found throughout Europe, Asia, and Africa, as well as [[North America]],<ref name=NatureServe>{{cite web |title=''Mantis religiosa'' |url=https://explorer.natureserve.org/Taxon/ELEMENT_GLOBAL.2.808478/Mantis_religiosa |website=NatureServe Explorer An online encyclopedia of life|version=7.1 |publisher=NatureServe |access-date=16 November 2023}}</ref> where it is considered introduced.<ref name="ReferenceC">{{cite journal|last1=Reitze|first1=Margit|last2=Nentwig|first2=Wolfgang|title=Comparative investigations into the feeding ecology of six Mantodea species|journal=Oecologia|date=1991|volume=86|issue=4|pages=568–574|doi=10.1007/BF00318324|pmid=28313339|bibcode=1991Oecol..86..568R|s2cid=22231027}}</ref> Two confirmed stable populations are in [[Germany]]: one in [[Rhineland-Palatinate]] and one in [[Baden-Württemberg]]. In the last years, more reports have been made of the distribution of the animals in [[Northern Europe]] (Latvia, Estonia), as well.<ref>{{cite journal|last1=Liana|first1=Anna|title=Distribution of ''Mantis religiosa'' (L.) and its changes in Poland|journal=Fragmenta Faunistica|date=2007|volume=50|issue=2|pages=91–125|doi=10.3161/00159301ff2007.50.2.091|url=http://rcin.org.pl/Content/45408|doi-access=free}}</ref><ref>{{cite journal|last1=Pupiņš|first1=Mihails|last2=Kalniņš|first2=Mārtiņš|last3=Pupiņa|first3=Aija|last4=Jaundaldere|first4=Ieva|title=FIRST RECORDS OF EUROPEAN MANTID MANTIS RELIGIOSA (LINNAEUS, 1758)(INSECTA: DICTYOPTERA, MANTIDAE) IN LATVIA and also LITHUANIA|journal=Acta Biologica Universitatis Daugavpiliensis|date=2012|volume=12|issue=2|pages=175–184}}</ref><ref>{{Cite web|url=https://novaator.err.ee/980971/eestisse-joudis-uus-lounamaine-loom-palvetajaritsikas|title=Eestisse jõudis uus lõunamaine loom – palvetajaritsikas|last=ERR|first=Katre Tatrik, Tartu Ülikool {{!}}|date=2019-09-16|website=ERR|language=et|access-date=2019-09-17}}</ref>

While it is absent from the [[British Isles]], ''M. religiosa'' occurs - and has been recorded breeding - on the [[Channel Islands]], with a particularly abundant population on the island of [[Jersey]], suggesting that its range may be capable of expanding north to [[Great Britain]].<ref>{{Cite web |date=2022-09-29 |title=Praying mantises set to swarm Britain |url=https://www.independent.co.uk/climate-change/news/praying-mantis-britain-uk-b2178186.html |access-date=2024-03-06 |website=The Independent |language=en}}</ref>

Despite being an introduced species, it is the official state insect of [[Connecticut]].<ref>[http://portal.ct.gov/About/State-Symbols/The-State-Insect CT.gov: The State Insect]; retrieved on March 20, 2018</ref>

==Reproduction==
The sexual behavior of praying mantis in general is curious, so has received interest from scientists over the last century. The differences between the various species are well known.

In ''M. religiosa'', [[courtship display|courtship]] and [[mating]] are separated into two steps: Preliminary courtship begins with the first visual contact between the animals and ends with the first physical contact. [[Copulation (zoology)|Copulation]] begins with physical contact and ends when the [[spermatophore]] is deposited:<ref name="Roeder1935" />

Only a few days after the final moult into adults, the animals begin to show interest in the opposite sex; this point is marked as the achievement of [[sexual maturity]]. Males have been observed to be most attracted to females during the middle of the day when the temperature is hottest. Theories are that the female [[pheromones]] are most [[volatility (chemistry)|volatile]] in the heat and that also the male, as a [[thermophilic]] insect, is most active.<ref name="ReferenceA" /><ref name="Lawrence" />

Instead of just observing them, sexually mature males approach sexually mature females when they see them, but due to the physical superiority of the females, males of ''M. religiosa'' face certain challenges in doing so. When a female spots a male, she is very likely to attack and kill him (see also: [[Sexual cannibalism]]). Therefore, males can be observed to be very slow and cautious in their approach; after spotting a female, the male usually freezes and turns his head to look directly at her. Since the [[fovea (spider)|fovea]]e in his eyes face directly forwards, he has the most accurate and detailed view of her and can watch every one of her moves.<ref name="jeb.biologists.org">{{cite journal|last1=Rossel|first1=Samuel|title=Binocular Spatial Localization in the Praying Mantis|journal=Journal of Experimental Biology|date=1 January 1986|volume=120|issue=1|pages=265–281|doi=10.1242/jeb.120.1.265|url=http://jeb.biologists.org/content/120/1/265.abstract|language=en|issn=0022-0949|doi-access=free}}</ref> He then proceeds to approach her from behind. Males can be observed to stop as soon as the female turns her head or even moves. Mantids are very good at detecting moving structures, but are almost unable to see immobile objects. Using this ‘stop-and-go’ tactic, the male stalks closer to the female. This can often take several hours. Depending on the environment, males sometimes show a light ‘rocking’-behavior which is believed to imitate the leaves of surrounding plants to blend in with the background. Males doing this had no higher probability of being detected and attacked, which supports this hypothesis of concealment.<ref name="Lawrence" />

No mention of male or female courtship has been made in either the field or captivity. The male, in fact, does not win by display; he wins by the absence of display.<ref name="Roeder1935" />
Once the male is close enough to the female, he opens his wings a little to facilitate his jump on the female's back. As soon as he lands, he proceeds to hold on to her with his raptorial legs. His tibiae fit into a pair of grooves on either side of the female's mesothorax in front of her wing bases.<ref name="Roeder1935" />
When the male is in a secure position, copulation is initiated. The [[genitals]], which are found on the end of the [[abdomen]] in both sexes, look quite different; the males possess a pair of asymmetrical claspers, while the females have an ovipositor. The [[ovipositor]] is rather short in comparison to other insects. It is covered in part by the edges of the last [[sternum]]. 
The abdomen of the male curves and twists in a 90° angle around the female's to insert the claspers between the ovipositor and the sternum. The abdomen of the male then contracts in a [[peristaltic]] manner. The animals can stay in this position for four to five hours before a spermatophore is deposited inside the female and the claspers are withdrawn.<ref name="Roeder1935" />
The male then lets go of the female to drop to the ground to get out of her reach for his own safety. After getting away about 50&nbsp;cm, he stops and freezes for about four minutes before he leaves eventually.<ref name="Lawrence" /> This behavior could be interpreted as necessary rest after the efforts of copulating in a safe distance from the female.

==Larval development==
The intervals of oviposition after copulating depend on the food intake and the overall physical fitness of the female. On average, 11 days are needed for the female to form and deposit an [[ootheca]], which contains around 100 to 200 eggs.<ref name="Lawrence" /> Copulation usually takes place in September or October, but the eggs overwinter and the larva do not hatch until the following spring. Hatching is strongly influenced by environmental conditions such as temperature (at least 17&nbsp;°C<ref name="ReferenceD">{{cite journal|last1=Przibram|first1=Hans|title=Aufzucht, Farbwechsel und Regeneration der Gottesanbeterinnen (Mantidae)|journal=Archiv für Entwicklungsmechanik der Organismen|date=November 1909|volume=28|issue=4|pages=561–628|doi=10.1007/BF02162074|s2cid=30725895}}</ref>) and humidity. Females prefer to deposit their eggs on solid substrates at warm and sunny sites. Most eggs from one ootheca hatch at the same time along the entire convex site, as worm-like prelarvae (L1). The hatchings always occur in the morning.<ref name="ReferenceA"/><ref name="KralOrientationBehavior" /><ref name="Przibram1907" />

The L1 just exists for a very short time; the first moult happens on or very close to the ootheca. The now-emerging nymphs already look a lot like the adults, but are maybe a tenth of their size. Very few animals (about 10%) survive this first instar stage due to lack of appropriately sized food, low temperatures, or insufficient humidity.<ref name="KralOrientationBehavior" />

The L2–L6 stages generally last about 14 days each. During this time, a growth around 6&nbsp;mm per stadium can be observed. Each stadium is completed with a [[moult]]. The L7 develops a more compact shape. Wing pads become visible. L7 and L8 show the same growth rate and duration as the previous stadia. After the next moult, the adult animal emerges. It now has wings and is fully developed. While more than eight moults have never been observed in ''M. religiosa'', females usually need one more moult than males under similar circumstances.

Closely related mantids have been reported to be larger than ''M. religiosa'' and require more moults (9–11). The fact that females need a longer time to develop and more moults might be due to the difference in size.<ref name="Przibram1907" />

==Sexual cannibalism==
[[File:Praying Mantis Sexual Cannibalism European-37.jpg|thumb|Praying mantis sexual cannibalism]]
The phenomenon that conspecifics are attacked and eaten after, during, and even before copulation is called [[sexual cannibalism]]. Many examples are known in various groups of [[invertebrates]], including the mantids. Some speculations exist about the benefits of this behaviour, but its causes are not fully understood yet.<ref>{{cite journal|last1=Prokop|first1=Pavol|last2=Maxwell|first2=Michael R.|title=Female predatory response to conspecific males and heterospecific prey in the praying mantis Mantis religiosa: evidence for discrimination of conspecific males|journal=Journal of Ethology|date=10 February 2016|volume=34|issue=2|pages=139–146|doi=10.1007/s10164-016-0458-8|s2cid=15224163}}</ref>
For quite some time, the belief that sexual cannibalism in ''M. religiosa'' only occurred in captivity was widespread. It was believed to be an artefact of improper animal keeping (too little food or space). In 1992, however, the cannibalising behaviour of female ''M. religiosa'' was observed in a field study. Lawrence compared the percentage of cannibalism in natural and manipulated (males and females were put close together in the field) pairings and found rates of 31% and 24%. 
This proves that sexual cannibalism in this praying mantid is not caused by laboratory conditions. They can, however, intensify the phenomenon; females that were starved for more than three days were more likely to attack males, even before copulation.<ref name="Lawrence" /> While the cannibalisation before the copulation is still puzzling to scientists,  some theories about the benefits of sexual cannibalism in general include:
*Eating her mate provides the female with nutrients so she does not have to hunt. She has a prey item available that is bigger than the prey she would be able to catch in the manner she usually hunts. 
*The meal also usually takes place during or shortly after she was fertilised, giving her more resources for the faster production of a large ootheca with large eggs, thus increasing the chance of her offspring to survive. Males have also been known to be more attracted to heavier, well-nourished females for this reason.<ref name="Lawrence" />

At first glance, however, this behaviour does not seem to be very beneficial for the male, since he dies and cannot create more offspring with his genetic material, yet males usually do not mate more than once, anyway. They have a shorter life expectancy than females (7–8 months versus 11–12 months) and since food can begin to become scarce in September and October, the male might starve before he has the chance to mate again. Copulation is also known to take longer when cannibalisation occurs. It seems advantageous to transfer more sperm in this elongated time and additionally provide the female with nutrients. Doing this, the male might just pass on his genes once, but he can ‘make sure’ that his offspring are successful, since they come from a well-nourished female.<ref name="Lawrence" />

Some controversy exists, though, concerning this theory. The absence of elaborate display behaviour does not support this ‘selfless’ behaviour of the males. It could have also been observed that males are more likely to approach a female that is distracted by feeding or cleaning herself. The approach is also happening faster in these cases. The male seems to try to avoid his cannibalisation very actively.<ref>{{cite journal|last1=Gemeno|first1=César|last2=Claramunt|first2=Jordi|title=Sexual Approach in the Praying Mantid Mantis Religiosa (L.)|journal=Journal of Insect Behavior|date=4 December 2006|volume=19|issue=6|pages=731–740|doi=10.1007/s10905-006-9058-8|bibcode=2006JIBeh..19..731G |s2cid=10647665}}</ref>

The fact that sexual cannibalisation happens so often is surely also supported because a male without a head can continue and even initiate copulation. The [[Cerebrum|cerebral]] [[ganglion]] might have an [[inhibitory]] effect on copulatory reflexes. Once it is removed by removing the head, copulation might even be longer, since the male can never ‘decide’ to let go.<ref name="Roeder1935" /><ref>{{cite journal|last1=Roeder|first1=Kenneth D|title=The control of tonus and locomotor activity in the praying mantis (Mantis religiosa L.)|journal=Journal of Experimental Zoology|date=August 1937|volume=76|issue=3|pages=353–374|doi=10.1002/jez.1400760302|bibcode=1937JEZ....76..353R }}</ref> A few mantids can also start copulating when the heads of both animals get removed artificially beforehand. The beheaded animals can live up to five hours and usually just die when they bleed out or starve to death.<ref name="Roeder1935" />

==Hunting==
[[File:European Mantis (Mantis religiosa) - Guelph, Ontario 02.jpg|thumb|right|''M. religiosa'' hunting for prey, using a plant as a vantage point]]
''M. religiosa'' is a [[carnivorous]] [[ambush predator]] that actively scans its environment and feeds on most insects that are not too large to be captured by rapid extension of its raptorial legs. Only living and moving prey is captured and consumed immediately using their powerful mandibles.<ref name="Roeder1935" /> Grasshoppers seem to be rather popular, probably because of their type of movement (flying or leaping),<ref>{{cite journal|last1=Mook|first1=L. J.|last2=Davies|first2=D. M.|title=The European Praying Mantis (Mantis religiosa L.) as a Predator of the Red-legged Grasshopper (Melanoplus femurrubrum (De Geer))|journal=The Canadian Entomologist|date=31 May 2012|volume=98|issue=9|pages=913–918|doi=10.4039/Ent98913-9|s2cid=86161013 }}</ref> but [[crickets]] and [[cockroaches]] are also frequently preyed upon. The attack or consumption of [[Mollusca]], [[Orb-weaver spider|Aranea]], [[Myriapoda]], or [[Oligochaeta]] is also not unknown. Caught prey gets trapped between the spikes on the [[femur|femora]] and [[tibia]]e and is now immobilized. The mantis then starts eating the still-living and preferably moving prey. Some years ago, the general opinion was that they killed their prey with a bite in the neck before consuming it, but current observations do not support this; the animals just start eating the body parts closest to their mouths.<ref name="ReferenceC"/>

''M. religiosa'' is also known to eat [[conspecifics]] outside a sexual context. Because of this, they usually need to be kept isolated from each other in captivity.<ref name="Przibram1907" /><ref name="ReferenceD"/>

==Visual system==
The large [[compound eyes]] that account for a great portion of the head make clear that [[visual perception|vision]] seems to be important for ''M. religiosa''. Their hunting behaviour and their sexual interactions rely almost exclusively on sight and the detection of movement. The capture of fast-flying prey out of the air would not be possible without a high level of temporal resolution. The localisation of the direction and distance of the prey are crucial here.

The eyes of ''M. religiosa'' are [[apposition eye]]s with eight types of [[photoreceptor cell]]s, so are best adapted for daylight vision.<ref name="eje.cz">{{cite journal|last1=Kral|first1=Karl|title=The functional significance of mantis peering behaviour|journal=European Journal of Entomology|date=2012|volume=109|issue=3|page=295|url=http://www.eje.cz/scripts/viewabstract.php?abstract=1711|doi=10.14411/eje.2012.039|doi-access=free}}</ref> One compound eye of an adult mantid consists of 8,000 to 10,000 optically isolated [[ommatidia]] with an interommatidial angle of 2° in the [[peripheral vision|periphery]] and 0.7° in the fovea.<ref name="KralOrientationBehavior" /><ref name="jeb.biologists.org" /> Interommatidial angles in insects vary from tens of degrees to 0.24° in [[dragonflies]], which puts the mantids on the upper end of [[spatial resolution]].<ref name="eje.cz" /><ref>{{cite journal|last1=Land|first1=Michael F.|s2cid=10924209|title=Visual Acuity in Insects|journal=Annual Review of Entomology|date=January 1997|volume=42|issue=1|pages=147–177|doi=10.1146/annurev.ento.42.1.147|pmid=15012311}}</ref> The overlap of the visual field of the two eyes is 40° in nymphs and up to 70° in adults.<ref name="KralOrientationBehavior" /><ref name="jeb.biologists.org" />

The [[pseudopupil]], a small, dark region in the eye, is rather conspicuous. It moves when the animal's head is moving and represents the area of parallel ommatidia, which absorb the incident light, so appear darker than the rest of the eye.<ref>{{cite journal|last1=Zeil|first1=J|last2=Al-Mutairi|first2=M|title=The variation of resolution and of ommatidial dimensions in the compound eyes of the fiddler crab Uca lactea annulipes (Ocypodidae, Brachyura, Decapoda)|journal=The Journal of Experimental Biology|date=1 January 1996|volume=199|issue=Pt 7|pages=1569–1577|doi=10.1242/jeb.199.7.1569|issn=1477-9145|pmid=9319471}}</ref>

The peering behaviour observed in ''M. religiosa'' is believed to be essential for the measurement of distances and depth perception; a side-to-side [[pendulum]]-like movement of the head or the whole body in a [[horizontal plane]] is used to scan the environment. [[Motion parallax]] is a depth cue that describes the fact that the closer the object is, the faster it seems to move when turning one's head while looking at it compared to slower-moving objects that are more distant.<ref>{{cite journal|last1=Kral|first1=Karl|last2=Devetak|first2=Dusan|title=The visual orientation strategies of Mantis religiosa and Empusa fasciata reflect differences in the structure of their visual surroundings|journal=Journal of Insect Behavior|date=1999|volume=12|issue=6|pages=737–752|doi=10.1023/a:1020948809024|s2cid=19745022}}</ref> Animals that were blinded on one eye did not strike for prey, showing that [[binocular vision]] is essential, since the disparity between the information of each eye is used to estimate distances, as well.<ref name="jeb.biologists.org" />

The development of the visual system was reviewed by Karl Kral in 2014:<ref name="KralOrientationBehavior" /> while high-contrast cues could be perceived by adults and nymphs (two hours to three days) alike, the differentiation between lower-contrast cues was much less successful in the two-hour-old animals, but a great improvement occurred after only three days of life. The eyes of newly hatched animals are less than half the size of the eyes of adult animals and have fewer ommatidia. Also, the [[Compound eye|facets]] of the frontal ommatidia, which are usually used for the detection of partners and prey, are not enlarged yet. While they already have all the necessary structures, very young animals have a restricted visual field and lower resolution and sensitivity to light. The great improvement of vision after only three days happens due to the [[sclerotisation]] of the [[cuticle]] that includes the corneal [[lens (anatomy)|lens]]es of the ommatidia. The improved lenses are not able to focus the light on the [[retina]].<ref name="KralOrientationBehavior" />

Like other mantid species, ''M. religiosa has''  [[stereopsis]] vision.<ref name="l409">{{cite journal |last=Rossel |first=Samuel |date=1983 |title=Binocular stereopsis in an insect |journal=Nature |volume=302 |issue=5911 |pages=821–822 |doi=10.1038/302821a0 |issn=0028-0836}}</ref> 

==Subspecies==
* ''M. r. beybienkoi'' is found in [[Kazakhstan]], [[Tajikistan]], [[Mongolia]], [[West Siberian Plain|West Siberia]], and [[Primorsky Krai]] ([[Russian Far East]])<ref>{{cite news | title=К ФАУНЕ БОГОМОЛОВ (MANTOPTERA) ДАЛЬНЕГО ВОСТОКА | issn= 1561-7858 | publisher= биолого-почвенный институт ДВНЦ АН СССР Владивосток | volume= С. 3-5 | year=1981 | first=С. Ю. | last=Стороженко | url=http://molbiol.ru/forums/index.php?act=Attach&type=post&id=155511  }}</ref>
* ''M. r. caucasica'' is found in [[Stavropol]].
* ''M. r. eichleri'' is found in [[Ethiopia]], [[Mauritania]], [[Niokolo-Koba]], [[Ghana]], [[Cameroon]], [[Kenya]], the [[Congo (area)|Congo]], [[Niger]], [[Senegal]], [[Somalia]], [[Sudan]], [[Tanzania]], [[Togo]], [[Zimbabwe]], [[Burkina Faso]], [[Zambia]], and [[Anatolia]].
* ''M. r. inornata'' is found in [[India]], [[Iran]], [[Nepal]], and [[Pakistan]],<ref name=MantodeaOfNepal>{{Cite web |url=http://www.orion-berlin.de/bibl/ehrmann/Ehrmann%2C%20R.%20%26%20Borer%2C%20M.%20%282015%29%20-%20%28E-159%29-Mantodea%20%28Insecta%29%20of%20Nepal%20an%20annotated%20checklist.PDF |title=Mantodea (Insecta) of Nepal: an annotated checklist By Reinhard Ehrmann, Karlsruhe & Matthias Borer, Basel |access-date=2022-01-26 |archive-date=2016-06-25 |archive-url=https://web.archive.org/web/20160625173811/http://www.orion-berlin.de/bibl/ehrmann/Ehrmann%2C%20R.%20%26%20Borer%2C%20M.%20%282015%29%20-%20%28E-159%29-Mantodea%20%28Insecta%29%20of%20Nepal%20an%20annotated%20checklist.PDF |url-status=dead }}</ref>
** syn. <small> ''M. r.  akbari'' (Soomro et al., 2001)</small>
** syn. <small> ''M. inornata'' (Werner, 1930)</small><ref name=MantodeaOfNepal/>
* ''M. r. langoalata'' is found in [[Uzbekistan]].
* ''M. r. latinota'' if found in Kazakhstan.
* ''M. r.  macedonica'' is found in [[North Macedonia]].
* ''M. r. major'' is found in [[Mombasa]] ([[Kenya]]).<ref name=MantodeaSpeciesFile>[http://mantodea.speciesfile.org/Common/specimen/ShowSpecimen.aspx?Router=NewPage Mantodea.SpeciesFile.org Page Mantis religiosa major Gerstaecker, 1873: Specimen records]</ref>
* ''M. r. polonica'' is found in [[Poland]] and [[Russia]].<ref>http://mantidforum.net/forums/index.php?showtopic=30736&hl=%2Bmantis+%2Breligiosa{{dead link|date=December 2017 |bot=InternetArchiveBot |fix-attempted=yes }}</ref>
* ''M. r.  religiosa'' is found in Africa, North America ([[Canada]], [[United States]]), Europe,<ref>[[:ko:항라사마귀]]</ref> and Asia ([[Nepal]]), but not in South America or Australia.<ref name=MantodeaOfNepal/>
** syn. <small> ''M. r.  maroccana'' (Thunberg, 1815)</small>
** syn. <small> ''M. r.  radiata'' (Fischer-Waldheim, 1846)</small>
** syn. <small> ''M. r.  sancta'' (Fabricius, 1787)</small><ref name=MantodeaSpeciesFile/>
* ''M. r. siedleckii'' is found in [[Southeast Asia]]: [[Taiwan]], [[Thailand]], [[Malaysia]], [[Java]], and [[Sulawesi]].
* ''M. r.  sinica'' is found in [[China]], [[Japan]], [[Korea]], [[Myanmar]], and [[Vietnam]].<ref>{{cite web|url=  http://mantodea.speciesfile.org/Common/basic/Taxa.aspx?TaxonNameID=1184170 |title= Taxa hierarchy}}</ref><ref>[[:es:Mantis religiosa]]</ref>

==Gallery==
<gallery>
File:Mantis religiosa Fore Coxa Marking.JPG|''M. religiosa'', female with indicative marking (black spot) on fore coxa
File:MantisLegGBMNH.jpg|The raptorial foreleg of a praying mantis
File:Mantis religiosa couple.JPG|Mating pair of ''M. religiosa'', Lower Austria
File:Mantis religiosa (egg case).jpg|Ootheca [[Sardinia]], [[Italy]]
File:Mantis religiosa (AF) left 01.jpg|Adult male ''M. religiosa'' in [[Saône-et-Loire]] (71, France) in September, 2008
File:Mantidae - Mantis religiosa-3.JPG|Brown adult male ''M.  religiosa'' 2008 October, Cerreto Ratti, [[Alessandria]]
File:Mantis religiosa2.jpg|''M. religiosa'' nymph, [[France]]
File:Mantis religiosa MHNT entière.jpg|Adult male ''M.  religiosa''
File:Brown praying mantis.jpg|Brown adult female M. religiosa
File:European-mantis-eats-grasshopper.JPG|Brown adult female ''M.  religiosa''
File:Mantis en Escorial, España; 2020.jpg|12&nbsp;cm. female adult (El Escorial, Spain)
File:Praying Mantis (Mantis religiosa religiosa) 07.jpg|Adult ''M. r. religiosa'' in Guelph, Ontario
File:Praying Mantis (Mantis religiosa religiosa) Ootheca 02.jpg|Ootheca in Guelph, Ontario
File:Praying Mantis (Mantis religiosa religiosa) Ootheca 06.jpg|Damaged ootheca in Guelph, Ontario
File:Europäische Gottesanbeterin.jpg|At the [[Aristotle]]park of [[Stageira Chalkidikis|Stagira]]
File:Mantide geotermica.jpg
File:Mantis religiosa - 720x576.ogv|Praying Mantis (Mantis religiosa) preying and eating a grasshopper.
Mantis religiosa female molting subadult.png|Subadult female Mantis religiosa molting to its adult stage.
File:European mantis.jpg|European mantis (Mantis religiosa) near the Black Sea, Odesa oblast, Ukraine
</gallery>

==References==
{{Reflist|25em}}

==External links==
* {{Commons category-inline|Mantis religiosa|''Mantis religiosa''}}
* {{Wikispecies-inline|Mantis religiosa|''Mantis religiosa''}}
* [http://blaisephoto.fr/mantes_religieuses.html Photographs of European mantis (''M.  religiosa'')]
* [https://www.youtube.com/watch?v=gF8nOj5WhcQ A European mantis feeding on a variety of insects]

{{Taxonbar|from=Q268319}}
{{Authority control}}

[[Category:Mantis (genus)]]
[[Category:Mantodea of Africa]]
[[Category:Mantodea of Asia]]
[[Category:Mantodea of Europe]]
[[Category:Mantodea of North America]]
[[Category:Mantodea of Southeast Asia]]
[[Category:Insects used as insect pest control agents]]
[[Category:Insects of the United States]]
[[Category:Symbols of Connecticut]]
[[Category:Insects described in 1758]]
[[Category:Taxa named by Carl Linnaeus]]