View source for Biology:Mantis

{{Short description|Order of insects}}
{{Automatic taxobox
| taxon = Mantodea
| fossil_range = {{fossilrange|Early Cretaceous | Recent}}
| image = European praying mantis (Mantis religiosa) green female Dobruja.jpg
| image_caption = ''Mantis religiosa'', Romania
| authority = Burmeister, 1838
| subdivision_ranks = Families
| subdivision = See text
| synonyms = * Manteodea <small>Burmeister, 1829</small>
* Mantearia
* Mantoptera
}}

'''Mantises''' are an [[Biology:Order|order]] ('''Mantodea''') of [[Biology:Insect|insect]]s that contains [[Biology:List of mantis genera and species|over 2,400 species in about 460 genera]] in 33 families. The largest family is the [[Biology:Mantidae|Mantidae]] ("mantids"<!-- {{efn|"Mantid" was once almost a synonym for Mantodea, given that few species outside the Mantidae were known; that has changed with recent discoveries.}} -->). Mantises are distributed worldwide in [[Earth:Temperate climate|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 [[Biology:Termite|termite]]s and [[Biology:Cockroach|cockroach]]es ([[Biology:Blattodea|Blattodea]]), which are all within the superorder [[Biology:Dictyoptera|Dictyoptera]]. Mantises are sometimes confused with stick insects ([[Biology:Phasmatodea|Phasmatodea]]), other elongated insects such as [[Biology:Grasshopper|grasshopper]]s ([[Biology:Orthoptera|Orthoptera]]), or other more distantly related insects with [[Biology:Raptorial|raptorial]] forelegs such as mantisflies ([[Biology:Mantispidae|Mantispidae]]). Mantises are mostly [[Biology:Ambush predator|ambush predator]]s, 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 [[Biology:Sexual cannibalism|sexual cannibalism]], eating their mates after copulation.

Mantises were considered to have [[Philosophy:Supernatural|supernatural powers]] by early civilizations, including {{wipe|Ancient Greece}}, ancient Egypt, and [[Place:Assyria|Assyria]]. A cultural trope popular in [[Engineering:Cartoon|cartoon]]s imagines the female mantis as a ''[[Social:Femme fatale|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.<ref>{{cite web |url=http://mantodea.speciesfile.org/HomePage.aspx |title=Mantodea Species File Online |first1=Daniel |last1=Otte |first2=Lauren |last2=Spearman |access-date=17 July 2012}}</ref> They are predominantly found in tropical regions, but some live in temperate areas.<ref name="Prete1999" /><ref>{{cite book |editor1=Prete, Fredrick R. |editor2=Wells, Harrington |editor3=Wells, Patrick H. |editor4=Hurd, Lawrence E. |title=The Praying Mantids |contribution=Mantid in Ecological Research |author=Hurd, I. E. |year=1999 |publisher=Johns Hopkins University Press |isbn=978-0-8018-6174-1 |page=[https://archive.org/details/prayingmantids0000unse/page/231 231] |url=https://archive.org/details/prayingmantids0000unse/page/231}}</ref> The [[Biology:Systematics|systematics]] of mantises have long been disputed. Mantises, along with stick insects ([[Biology:Phasmatodea|Phasmatodea]]), were once placed in the order [[Biology:Orthoptera|Orthoptera]] with the cockroaches (now [[Biology:Blattodea|Blattodea]]) and ice crawlers (now [[Biology:Grylloblattodea|Grylloblattodea]]). Kristensen (1991) combined the Mantodea with the cockroaches and [[Biology:Termite|termite]]s into the order [[Biology:Dictyoptera|Dictyoptera]], suborder Mantodea.<ref name="Costa">{{cite book |last=Costa |first=James |title=The Other Insect Societies |publisher=Harvard University Press |pages=135–136 |isbn=978-0-674-02163-1 |url=https://books.google.com/books?id=PYRFDrZs9QAC&pg=PA135 |year=2006}}</ref><ref name=Capinera2008>{{cite book |author=Capinera, John L. |title=Encyclopedia of Entomology |url=https://books.google.com/books?id=i9ITMiiohVQC&pg=PA3033 |year=2008 |volume=4 |publisher=Springer |isbn=978-1-4020-6242-1 |pages=3033–3037}}</ref> The name ''mantodea'' is formed from the Ancient Greek words {{lang|el|μάντις}} (''mantis'') meaning "prophet", and {{lang|el|εἶδος}} (''eidos'') meaning "form" or "type". It was coined in 1838 by the German entomologist Hermann Burmeister.<ref>{{cite book |title=College entomology |last=Essig |first=Edward Oliver |year=1947 |publisher=Macmillan Company |oclc=809878 |pages=[https://archive.org/details/in.ernet.dli.2015.1341/page/n136 124], 900 |url=https://archive.org/details/in.ernet.dli.2015.1341}}</ref><ref name="OED">{{OEtymD|mantis}}</ref> 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 [[Biology:Mantidae|Mantidae]], which was, historically, the only family in the order. The other common name, praying mantis, applied to any species in the order<ref>{{cite book |author=Bullock, William |year=1812 |url=https://archive.org/details/companiontomrbul00bull |title=A companion to the London Museum and Pantherion |edition=12th}}</ref> (though in Europe mainly to ''Mantis religiosa''), comes from the typical "[[Unsolved:Prayer|prayer]]-like" posture with folded forelimbs.<ref>{{cite book |author=Partington, Charles Frederick |year=1837 |url=https://archive.org/details/britishcyclopae00partgoog |title=The British Cyclopædia of Natural History |publisher=W. S. Orr |volume=1}}</ref><ref name=NatGeo>{{cite web |title=Praying Mantis |date=10 September 2010 |publisher=National Geographic Society |url=http://animals.nationalgeographic.com/animals/bugs/praying-mantis/ |archive-url=https://web.archive.org/web/20100111130055/http://animals.nationalgeographic.com/animals/bugs/praying-mantis/ |url-status=dead |archive-date=11 January 2010 |access-date=28 August 2015}}</ref> 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.<ref>Bragg, P. E. (1996). "Mantis, Mantid, Mantids, Mantises". ''Mantis Study Group Newsletter'', 1:4.</ref><ref name="Ehrmann">Ehrmann, R. 2002. Mantodea: Gottesanbeterinnen der Welt. Natur und Tier, Münster.</ref>

=== Phylogeny ===

==== External ====

Evolutionary relationships based on Evangelista et al. 2019 are shown in the [[Biology:Cladogram|cladogram]]:<ref name="Evangelista 2019">{{cite journal |last1=Evangelista |first1=D.A.|last2=Wipfler |first2=B. |last3=O. |first3=Bethoux |last4=Donath |first4=A. |last5=Fujita |first5=M. |last6=Kohli |first6=M.K. |last7=Legendre |first7=F. |last8=Liu |last9=Machida |last10=Misof |last11=Peters |last12=Podsiadlowski |last13=Rust |last14=Schuette |last15=Tollenaar |last16=Ware |last17=Wappler |last18=Zhou |last19=Meusemann |last20=Simon |title=An integrative phylogenomic approach illuminates the evolutionary history of cockroaches and termites (Blattodea) |journal=Proc. R. Soc. B |date=23 January 2019 |volume=286 |issue=1895 |doi=10.1098/rspb.2018.2076 |pmid=30963947 |pmc=6364590 |doi-access=free }}</ref> 

{{clade
|label1=[[Biology:Dictyoptera|Dictyoptera]]
|1={{clade
   |label1='''Mantodea''' 
   |1= (Mantises)
   |label2=[[Biology:Blattodea|Blattodea]]
   |2= (Cockroaches and termites)
   }}
}}

==== Internal ====

One of the earliest classifications splitting an all-inclusive Mantidae into multiple families was that proposed by Beier in 1968, recognizing eight families,<ref name=Beier1968>{{cite journal |author=Beier, M. |year=1968 |title=Ordnung Mantodea (Fangheuschrecken) |journal=Handbuch der Zoologie |volume=4 |issue=2 |pages=3–12}}</ref> though it was not until Ehrmann's reclassification into 15 families in 2002<ref name="Ehrmann"/> that a multiple-family classification became universally adopted. Klass, in 1997, studied the external male genitalia and postulated that the families [[Biology:Chaeteessidae|Chaeteessidae]] and [[Biology:Metallyticidae|Metallyticidae]] diverged from the other families at an early date.<ref name="Klass1997">{{cite book |last=Klass |first=Klaus-Dieter |title=The external male genitalia and phylogeny of Blattaria and Mantodea |url=https://books.google.com/books?id=LfJMAAAAYAAJ |year=1997 |publisher=Zoologisches Forschungsinstitut |isbn=978-3-925382-45-1}}</ref> However, as previously configured, the [[Biology:Mantidae|Mantidae]] and [[Biology:Thespidae|Thespidae]] especially were considered polyphyletic,<ref name="MartillBechly2007">{{cite book |last1=Martill |first1=David M.|last2=Bechly |first2=Günter |last3=Loveridge |first3=Robert F. |title=The Crato Fossil Beds of Brazil: Window into an Ancient World |url=https://books.google.com/books?id=5ZBh_-QvX2MC&pg=PA236 |year=2007 |publisher=Cambridge University Press |isbn=978-1-139-46776-6 |pages=236–238}}</ref> so the Mantodea have been revised substantially as of 2019 and now includes 29 families.<ref name="Schwarz">[https://www.tandfonline.com/doi/full/10.1080/00379271.2018.1556567 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 &#91;2&#93;''': 101–196.]</ref>

<div style="overflow:auto;">{{cladogram
|align=left
|title=Cladogram of extant Mantodea families<ref>{{Cite journal |last1=Grimaldi |first1=David |date= 28 July 2003 |title=A Revision of Cretaceous Mantises and Their Relationships, Including New Taxa (Insecta: Dictyoptera: Mantodea) |journal=American Museum Novitates |volume= |issue=3412 |hdl=2246/2838 |pages=1–47|doi=10.1206/0003-0082(2003)412<0001:AROCMA>2.0.CO;2 |s2cid=56007623 |url=https://www.biodiversitylibrary.org/bibliography/178207 }}</ref><ref name="Schwarz" />
|{{clade|style=font-size:80%;
|label1=Mantodea
|1={{clade 
    |1=† Extinct Genera
    |label2=Eumantodea
    |2={{clade
         |label1=Chaeteessoidea
         |1=[[Biology:Chaeteessidae|Chaeteessidae]]
         |label2=Spinomantodea
         |2={{clade
                |label1=Mantoidoidea
                |1=[[Biology:Mantoididae|Mantoididae]]
                |label2=Schizomantodea
                |2={{clade
                  |label1=Metallyticoidea
                  |1=[[Biology:Metallyticidae|Metallyticidae]]
                  |label2=Artimantodea
                  |2={{clade
                    |label1=Amerimantodea
                    |1={{clade
                      |label1=Thespoidea
                      |1=[[Biology:Thespidae|Thespidae]]
                        |label2=Acanthopoidea
                        |2={{clade
                          |1=[[Biology:Angelidae|Angelidae]]
                          |2={{clade
                            |1=[[Biology:Coptopterygidae|Coptopterygidae]]
                            |2={{clade
                              |1=[[Biology:Liturgusidae|Liturgusidae]]
                              |2={{clade
                                |1=[[Biology:Photinaidae|Photinaidae]]
                                |2=[[Biology:Acanthopidae|Acanthopidae]]
                                 }}
                               }}
                             }}
                        }}  }}
              |label2=Cernomantodea
              |2={{clade
                      |label1=Nanomantodea
                      |1={{clade
                        |label1=Chroicopteroidea
                        |1=[[Biology:Chroicopteridae|Chroicopteridae]]
                        |label2=Nanomantoidea
                        |2={{clade
                          |1=[[Biology:Leptomantellidae|Leptomantellidae]]
                          |2={{clade
                            |1=[[Biology:Amorphoscelidae|Amorphoscelidae]]
                            |2=[[Biology:Nanomantidae|Nanomantidae]]
                             }}
                           }}  }}
                      |label2=Metamantodea
                      |2={SUBCLADE_Metamantodea}
                       }}
                     }}
                   }}
                 }}
               }} }}
  |targetA={SUBCLADE_Metamantodea}
  |subcladeA={{Clade
  |1={{clade
     |label1=Gonypetoidea
     |1=[[Biology:Gonypetidae|Gonypetidae]]
     |label2=Lobipedia
     |2={{clade
       |label1=Epaphroditoidea
       |1={{clade
         |1=[[Biology:Epaphroditidae|Epaphroditidae]]
         |2=[[Biology:Majangidae|Majangidae]]
          }}
       |label2=Mantimorpha
       |2={{clade
         |label1=Haanioidea
         |1=[[Biology:Haaniidae|Haaniidae]]
         |label2=Heteromantodea
         |2={{clade
           |label1=Eremiaphiloidea
           |1={{clade
             |1=[[Biology:Rivetinidae|Rivetinidae]]
             |2={{clade
               |1=[[Biology:Amelidae|Amelidae]]
               |2={{clade
                 |1=[[Biology:Eremiaphilidae|Eremiaphilidae]]
                 |2=[[Biology:Toxoderidae|Toxoderidae]]
                }}
              }}  }}
           |label2=Pareumantodea
           |2={{clade
             |label1=Hoplocoryphoidea
             |1=[[Biology:Hoplocoryphidae|Hoplocoryphidae]]
             |label2=Calomantodea
             |2={{clade
               |label1=Miomantoidea
               |1=[[Biology:Miomantidae|Miomantidae]]
               |label2=Promantidea
               |2={{clade
                 |label1=Galinthiadoidea
                 |1=[[Biology:Galinthiadidae|Galinthiadidae]]
                 |label2=Mantidea
                 |2={{clade
                   |label1=Hymenopodoidea
                   |1={{clade
                     |1=[[Biology:Empusidae|Empusidae]]
                     |2=[[Biology:Hymenopodidae|Hymenopodidae]]
                    }}
                   |label2=Mantoidea
                   |2={{clade
                     |1=[[Biology:Dactylopterygidae|Dactylopterygidae]]
                     |2={{clade
                       |1=[[Biology:Deroplatyidae|Deroplatyidae]]
                       |2=[[Biology:Mantidae|Mantidae]]
                        }}
                      }}
                    }}
                  }}
                }}
              }}
            }}
          }}
        }}
      }}
  }}  }}
|}}</div>


===Fossil mantises===
[[File:Santanmantis reconstruction.png|thumb|Life restoration of ''[[Biology:Santanmantis|Santanmantis]]'', a primitive fossil mantis known from the Early Cretaceous of Brazil, and one of the oldest members of the group]]
Mantises are thought to have evolved from cockroach-like ancestors.<ref>{{Cite journal |last=Li |first=Xinran |date=2019-08-30 |title=Disambiguating the scientific names of cockroaches |url=https://www.mapress.com/j/pe/article/view/palaeoentomology.2.4.13 |journal=Palaeoentomology |volume=2 |issue=4 |pages=390–402 |doi=10.11646/palaeoentomology.2.4.13 |s2cid=202789239 |issn=2624-2834}}</ref> The earliest confidently identified mantis fossils date to the [[Earth:Early Cretaceous|Early Cretaceous]].<ref name="MartillBechly2007" /> Fossils of the group are rare: by 2022, 37 fossil species are known.<ref name="MartillBechly2007" /><ref>{{Cite journal |last1=Terríquez-Beltrán |first1=J. |last2=Riquelme |first2=F. |last3=Varela-Hernández |first3=F. |date=2022-10-31 |title=A new species of mantis (Insecta: Mantodea: Amelidae) from the Miocene Amber-Lagerstätte in Mexico |url=https://www.tandfonline.com/doi/full/10.1080/08912963.2022.2134782 |journal=Historical Biology |language=en |pages=1–8 |doi=10.1080/08912963.2022.2134782 |s2cid=253280592 |issn=0891-2963}}</ref> Fossil mantises, including one from Japan with spines on the front legs as in modern mantises, have been found in [[Earth:Cretaceous|Cretaceous]] amber.<ref>{{cite web |last1=Ryall |first1=Julian |title=Ancient Praying Mantis Found in Amber |url=http://news.nationalgeographic.com/news/2008/04/080425-amber-mantis.html |archive-url=https://web.archive.org/web/20080429193405/http://news.nationalgeographic.com/news/2008/04/080425-amber-mantis.html |url-status=dead |archive-date=29 April 2008 |publisher=National Geographic Society |access-date=30 July 2015 |date=25 April 2008}}</ref> Most fossils in amber are nymphs; compression fossils (in rock) include adults. Fossil mantises from the [[Earth:Crato Formation|Crato Formation]] in Brazil include the {{convert|10|mm|in|adj=on|abbr=on}} long ''[[Biology:Santanmantis|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 [[Astronomy:X-ray computed tomography|X-ray computed tomography]].<ref name="MartillBechly2007" />  Extinct families and genera include:

{{div col|colwidth=22em}}
* †[[Biology:Baissomantidae|Baissomantidae]]
* †Gryllomantidae
* †[[Biology:Cretomantidae|Cretomantidae]]
* †[[Biology:Santanmantidae|Santanmantidae]]
* †[[Biology:Amelidae|Amelidae]]
* ''[[Biology:Incertae sedis|Incertae sedis]]'': 
** †''Jersimantis''
** †''Chaeteessites''
** †''Cretophotina''
** †''Ambermantis''
{{div col end}}

===Similar insects in the [[Biology:Neuroptera|Neuroptera]]===
Because of the superficially similar [[Biology:Raptorial|raptorial]] forelegs, [[Biology:Mantispidae|mantidflies]] may be confused with mantises, though they are unrelated. Their similarity is an example of [[Biology:Convergent evolution|convergent evolution]]; mantidflies do not have [[Biology:Tegmen|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.<ref>{{cite journal |last1=Boyden |first1=Thomas C. |title=Mimicry, predation and potential pollination by the mantispid, ''Climaciella brunnea'' var. ''instabilis'' (Say) (Mantispidae: Neuroptera) |journal=Journal of the New York Entomological Society |year=1983 |volume=91 |issue=4 |pages=508–511 |jstor=25009393}}</ref> Another example of confusion caused by convergent evolution is [[Biology:Titanoptera|Titanoptera]], an order of insect that lived in the Triassic period and also shared the [[Biology:Raptorial|Raptorial forelegs]] of a mantis.

== 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 spines
|caption2=The raptorial foreleg, showing the unusually long coxa, which, together with the trochanter, gives the impression of a femur. The femur itself is the [[Biology:Anatomical 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 [[Biology:Mandible (insect mouthpart)|mandibles]]. They have two bulbous compound eyes, three small simple eyes, and a pair of [[Biology:Antenna|antennae]]. The articulation of the neck is also remarkably flexible; some species of mantis can rotate their heads nearly 180°.<ref name=NatGeo /> 
The mantis thorax consists of a [[Biology:Prothorax|prothorax]], a [[Biology:Mesothorax|mesothorax]], and a [[Biology:Metathorax|metathorax]]. In all species apart from the genus ''[[Biology:Mantoida|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.<ref name="prete" /><ref name="close up">{{cite book |last=Siwanowicz |first=Igor |year=2009 |title=Animals Up Close |url=https://archive.org/details/Animals_Up_Close_by_Igor_Siwanowicz_DK |publisher=Dorling Kindersley |pages=[https://archive.org/details/Animals_Up_Close_by_Igor_Siwanowicz_DK/page/n38 38]–39 |isbn=-978-1-405-33731-1}}</ref> Mantises also are unique to the [[Biology:Dictyoptera|Dictyoptera]] in that they have tympanate hearing, with two [[Biology:Tympanum (anatomy)|tympana]] in an auditory chamber in their metathorax. Most mantises can only hear [[Physics:Ultrasound|ultrasound]].<ref>{{cite journal |last1=Yager |first1=David |last2=Svenson |first2=Gavin |title=Patterns of praying mantis auditory system evolution based on morphological, molecular, neurophysiological, and behavioural data. |journal=Biological Journal of the Linnean Society |date=1 July 2008 |volume=94 |issue=3 |pages=541–568 |doi=10.1111/j.1095-8312.2008.00996.x |publisher=Oxford University Press |language=en|doi-access=free}}</ref>

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.<ref name="prete">{{Cite book |title=The Praying Mantids |editor1=Prete, Fredrick R. |editor2=Wells, Harrington |editor3=Wells, Patrick H. |editor4=Hurd, Lawrence E. |contribution=Morphology and Taxonomy |author=Roy, Roger |year=1999 |pages=[https://archive.org/details/prayingmantids0000unse/page/21 21–33] |publisher=Johns Hopkins University Press |isbn=978-0-8018-6174-1 |url=https://archive.org/details/prayingmantids0000unse/page/21}}</ref><ref name="Corrette" />

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 [[Biology:Tegmen|tegmina]], are usually narrow and leathery. They function as camouflage and as a shield for the hindwings, which are clearer and more delicate.<ref name="prete" /><ref>{{cite web |last=Kemper |first=William T. |title=Insect Order ID: Mantodea (Praying Mantises, Mantids) |url=http://www.missouribotanicalgarden.org/Portals/0/Gardening/Gardening%20Help/Insect%20Orders/Mantodea.pdf |publisher=Missouri Botanical Garden |access-date=21 September 2015}}</ref>
The abdomen of all mantises consists of 10 [[Biology:Tergum|tergites]], with a corresponding set of nine [[Biology:Sternum (arthropod anatomy)|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 [[Biology:Cercus|cerci]] in both sexes.<ref name="prete" />

=== Vision ===
[[File:Large brown mantid close up nohair.jpg|thumb|left|Head of ''[[Biology:Archimantis latistyla|Archimantis latistyla]]'', showing the compound eyes and labrum|alt=Head of a mantis with large compound eyes and labrum]]

Mantises have [[Stereopsis|stereo vision]].<ref>{{cite journal |doi=10.1038/srep18718|pmid=26740144|pmc=4703989|title=Insect stereopsis demonstrated using a 3D insect cinema|journal=Scientific Reports|volume=6|pages=18718|year=2016|last1=Nityananda|first1=Vivek|last2=Tarawneh|first2=Ghaith|last3=Rosner|first3=Ronny|last4=Nicolas|first4=Judith|last5=Crichton|first5=Stuart|last6=Read|first6=Jenny|bibcode=2016NatSR...618718N}}</ref><ref>{{cite journal | last1 = Rossel | first1 = S | year = 1983 | title = Binocular stereopsis in an insect | journal = Nature | volume = 302 | issue = 5911| pages = 821–822 | doi=10.1038/302821a0| bibcode = 1983Natur.302..821R | s2cid = 4264254}}</ref><ref>{{cite journal |pages=13229–13232|pmc=24075|year=1996|last1=Rossel|first1=S|title=Binocular vision in insects: How mantids solve the correspondence problem|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=93|issue=23|pmid=11038523|doi=10.1073/pnas.93.23.13229|bibcode=1996PNAS...9313229R|doi-access=free}}</ref> They locate their prey by sight; their compound eyes contain up to 10,000 [[Biology:Ommatidium|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.<ref name="Corrette">{{cite journal |last1=Corrette |first1=Brian J. |title=Prey capture in the praying mantis ''Tenodera aridifolia sinensis'': coordination of the capture sequence and strike movements |journal=Journal of Experimental Biology |date=1990 |volume=148 |pages=147–180 |doi=10.1242/jeb.148.1.147 |url=http://jeb.biologists.org/content/148/1/147.full.pdf |pmid=2407798}}</ref><ref name=Simmons>{{cite book |author1=Simmons, Peter J. |author2=Young, David |title=Nerve Cells and Animal Behaviour |url=https://books.google.com/books?id=WRBHlAYfADUC&pg=PA89 |year=1999 |publisher=Cambridge University Press |isbn=978-0-521-62726-9 |page=8990}}</ref> The eyes are widely spaced and [[Biology:Anatomical terms of location|laterally]] situated, affording a wide [[Binocular vision|binocular field of vision]] and precise stereoscopic vision at close range.<ref name=Howard>{{cite book |author1=Howard, Ian P. |author2=Rogers, Brian J. |title=Binocular Vision and Stereopsis |url=https://books.google.com/books?id=I8vqITdETe0C&pg=PA646 |year=1995 |publisher=Oxford University Press |isbn=978-0-19-508476-4 |page=646}}</ref> The dark spot on each eye that moves as it rotates its head is a [[Biology:Pseudopupil|pseudopupil]]. This occurs because the ommatidia that are viewed "head-on" absorb the incident light, while those to the side reflect it.<ref name="Zeil">{{cite journal |author1=Zeil, Jochen |author2=Al-Mutairi, Maha M. |year=1996 |title=Variations in the optical properties of the compound eyes of ''Uca lactea annulipes'' |journal=The Journal of Experimental Biology |volume=199 |issue=7 |pages=1569–1577 |doi=10.1242/jeb.199.7.1569 |url=http://jeb.biologists.org/cgi/reprint/199/7/1569.pdf |pmid=9319471}}</ref>

As their hunting relies heavily on vision, mantises are primarily [[Biology:Diurnality|diurnal]]. Many species, however, fly at night, and then may be [[Biology:Phototaxis#Insects|attracted to artificial lights]]. Mantises in the family [[Biology:Liturgusidae|Liturgusidae]] collected at night have been shown to be predominately males;<ref>{{cite journal | last1 = Bragg | first1 = P.E. | year = 2010 | title = A review of the Liturgusidae of Borneo (Insecta: Mantodea) | journal = Sepilok Bulletin | volume = 12 | pages = 21–36}}</ref> this is probably true for most mantises. Nocturnal flight is especially important to males in locating less-mobile females by detecting their [[Biology:Pheromone|pheromone]]s. 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 [[Philosophy:Animal echolocation|echolocation]] calls and responding evasively.<ref name="Yager">{{Cite book |title=The Praying Mantids |editor1=Prete, Fredrick R. |editor2=Wells, Harrington |editor3=Wells, Patrick H. |editor4=Hurd, Lawrence E. |year=1999 |contribution=Hearing |author=Yager, David |pages=[https://archive.org/details/prayingmantids0000unse/page/101 101–103] |publisher=Johns Hopkins University Press |isbn=978-0-8018-6174-1 |url=https://archive.org/details/prayingmantids0000unse/page/101}}</ref><ref>{{cite book |last1=Grimaldi |first1=David |last2=Engel |first2=Michael, S. |year=2005 |title=Evolution of the Insects |url=https://archive.org/details/evolutioninsects00grim_110 |url-access=limited |publisher=Cambridge University Press |page=[https://archive.org/details/evolutioninsects00grim_110/page/n271 257] |isbn=978-0-521-82149-0}}</ref>

=== Diet and hunting ===
[[File:Tenodera sinensis 2 Luc Viatour.jpg|thumb|''Tenodera sinensis'' feeding on a cricket|alt=Mantis eating a cricket]]
Mantises are generalist predators of [[Biology:Arthropod|arthropod]]s.<ref name="Prete1999" /> The majority of mantises are [[Biology:Ambush predator|ambush predator]]s 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.<ref>{{cite journal |title=Mantids – The Praying Predators |journal=National Geographic |first=Edward S. |last=Ross |pages=268–280 |volume=165 |issue=2 |date=February 1984 |issn=0027-9358 |oclc=643483454}}</ref> Larger mantises sometimes eat smaller individuals of their own species,<ref>{{cite book |author=Capinera, John L. |title=Encyclopedia of Entomology |url=https://books.google.com/books?id=i9ITMiiohVQC&pg=PA1509 |year=2008 |publisher=Springer |isbn=978-1-4020-6242-1 |page=1509}}</ref> as well as small vertebrates such as lizards, frogs, fish, and particularly small birds.<ref>{{cite book |last1=Grimaldi |first1=David |last2=Engel |first2=Michael S. |title=Evolution of the Insects |url=https://books.google.com/books?id=Ql6Jl6wKb88C&pg=PA254 |year=2005 |publisher=Cambridge University Press |isbn=978-0-521-82149-0 |page=254}}</ref><ref>{{cite journal |author1=Nyffeler, Martin|author2=Maxwell, Michael R.|author3=Remsen, J. V., Jr. |title=Bird predation by praying mantises: a global perspective |journal=The Wilson Journal of Ornithology |year=2017 |volume=129 |issue=2 |pages=331–344 |doi=10.1676/16-100.1|s2cid=90832425}}</ref><ref>{{cite journal |last1=Battison |first1=Roberto |display-authors=0|title=The fishing mantid: predation on fish as a new adaptive strategy for praying mantids (Insecta: Mantodea) |journal=Journal of Orthoptera Research |volume=27 |issue=2 |pages=155–158 |doi=10.3897/jor.27.28067 |url=https://jor.pensoft.net/article/28067/element/8/49047/|year=2018 |doi-access=free}}</ref>

Most mantises stalk tempting prey if it strays close enough, and will go further when they are especially hungry.<ref name=Gelperin>{{cite journal |last1=Gelperin |first1=Alan |title=Feeding behaviour of the praying mantis: a learned modification |journal=Nature |date=July 1968 |volume=219 |issue=5152 |pages=399–400 |doi=10.1038/219399a0 |pmid=5667084 |bibcode=1968Natur.219..399G |s2cid=10951932}}</ref> Once within reach, mantises strike rapidly to grasp the prey with their spiked raptorial forelegs.<ref>{{cite journal |last1=Prete |first1=F. R. |last2=Cleal |first2=K. S. |title=The predatory strike of free ranging praying mantises, ''Sphodromantis lineola'' (Burmeister). I: Strikes in the mid-sagittal plane |journal=Brain, Behavior and Evolution |date=1996 |volume=48 |issue=4 |pages=173–190 |pmid=8886389 |doi=10.1159/000113196}}</ref> 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 [[Biology:Tiger beetle|tiger beetle]]s do.<ref name="prete" />

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.<ref>{{cite book |author=Capinera, John L. |title=Encyclopedia of Entomology |url=https://books.google.com/books?id=i9ITMiiohVQC&pg=PA116 |year=2008 |publisher=Springer |isbn=978-1-4020-6242-1 |page=116}}</ref> [[Biology:Chinese mantis|Chinese mantis]]es live longer, grow faster, and produce more young when they are able to eat [[Biology:Pollen|pollen]].<ref>{{cite journal |doi=10.1603/0046-225X-32.4.881 |title=Pollen feeding and fitness in praying mantids: the vegetarian side of a tritrophic predator |journal=Environmental Entomology |volume=32 |issue=4 |pages=881–885 |year=2003 |last1=Beckman |first1=Noelle |last2=Hurd |first2=Lawrence E.|doi-access=free}}</ref>

=== Antipredator adaptations ===
{{further|Biology:Flower mantis}}

Mantises are preyed on by vertebrates such as frogs, lizards, and birds, and by invertebrates such as spiders, [[Biology:Asian giant hornet#Predation|large species of hornets]], and ants.<ref>{{cite journal |last1=Iyer |first1=Geetha |title=Down to earth |url=http://www.frontline.in/static/html/fl2817/stories/20110826281706800.htm |journal=Frontline Magazine |volume=28 |issue=17 |date=13 August 2011}}</ref> Some hunting wasps, such as some species of ''[[Biology:Tachytes|Tachytes]]'' also paralyze some species of mantis to feed their young.<ref name="Fabre1921">{{cite book|author=Jean-Henri Fabre|title=More Hunting Wasps|url=https://archive.org/details/morehuntingwasps00fabrrich|year=1921|publisher=Dodd, Mead}}</ref> Generally, mantises protect themselves by camouflage, most species being cryptically colored to resemble [[Biology:Leaf|foliage]] or other backgrounds, both to avoid predators and to better snare their prey.<ref>{{cite book |author1=Gullan, P. J. |author2=Cranston, P. S. |year=2010 |title=The Insects: An Outline of Entomology |url=https://archive.org/details/insectsoutlineen00pjgu |url-access=limited |publisher=Wiley |edition=4th |isbn=978-1-118-84616-2 |page=[https://archive.org/details/insectsoutlineen00pjgu/page/n394 370]}}</ref> 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.<ref name=Prete282>{{Cite book |title=The Praying Mantids |editor1=Prete, Fredrick R. |editor2=Wells, Harrington |editor3=Wells, Patrick H. |editor4=Hurd, Lawrence E. |year=1999 |pages=[https://archive.org/details/prayingmantids0000unse/page/282 282–293] |author1=Edmunds, Malcolm |author2=Brunner, Dani |contribution=Ethology of Defenses against Predators |publisher=Johns Hopkins University Press |isbn=978-0-8018-6174-1 |url=https://archive.org/details/prayingmantids0000unse/page/282}}</ref> The species from different families called [[Biology:Flower mantis|flower mantis]]es are [[Biology:Aggressive mimicry|aggressive mimics]]: they resemble flowers convincingly enough to attract prey that come to collect pollen and nectar.<ref name=Cott>{{cite book| last=Cott | first=Hugh | title=Adaptive Coloration in Animals | publisher=Methuen | year=1940 | pages=[https://archive.org/details/adaptivecolorati00cott/page/392 392–393] |  title-link=Adaptive Coloration in Animals}}</ref><ref>{{cite journal | last=Annandale | first=Nelson | title=Observations on the habits and natural surroundings of insects made during the 'Skeat Expedition' to the Malay Peninsula, 1899–1900| journal=Proceedings of the Zoological Society of London |volume=69 | year=1900 | pages=862–865}}</ref><ref>{{cite journal|last1=O'Hanlon |first1=James C. |last2=Holwell |first2=Gregory I. |last3=Herberstein |first3=Marie E. |title=Pollinator deception in the orchid mantis |journal=The American Naturalist|year=2014 |volume=183 |issue=1 |pages=126–132 |doi=10.1086/673858 |pmid=24334741|s2cid=2228423 |doi-access=free }}</ref> 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).<ref name=Prete282 />

[[File:Mantis Hymenopus coronatus 2 Luc Viatour.jpg|thumb|upright|[[Biology:Aggressive mimicry|Aggressive mimicry]]: Malaysian orchid mantises are camouflaged pink or yellow, matching the coloration of local orchids.]]

When directly threatened, many mantis species stand tall and spread their forelegs, with their wings fanning out wide. The fanning of the wings makes the mantis seem larger and more threatening, with some species 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 [[Biology:Spiracle (arthropods)|spiracle]]s. 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.<ref name=Prete282 /><ref>{{cite journal |last1=Yager |first1=D. |last2=May |first2=M. |year=1993 |title=Coming in on a wing and an ear |journal=Natural History |volume=102 |issue=1 |pages=28–33}}</ref><ref>{{cite web |url=http://news.nationalgeographic.com/news/2002/11/1119_021119_TVbats_2.html |archive-url=https://web.archive.org/web/20060521084215/http://news.nationalgeographic.com/news/2002/11/1119_021119_TVbats_2.html |url-status=dead |archive-date=21 May 2006 |title=Praying Mantis Uses Ultrasonic Hearing to Dodge Bats |date=19 November 2002 |publisher=National Geographic Society |access-date=17 August 2012}}</ref>

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 [[Biology:Crypsis|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.<ref>{{cite journal |author=O'Dea, J. D. |title=Eine zusatzliche oder alternative Funktion der 'kryptischen' Schaukelbewegung bei Gottesanbeterinnen und Stabschrecken (Mantodea, Phasmatodea) |journal=Entomologische Zeitschrift |volume=101 |issue=1–2 |year=1991 |pages=25–27}}</ref> As ants may be predators of mantises, genera such as ''Loxomantis'', ''[[Biology:Orthodera|Orthodera]]'', and ''[[Biology:Statilia|Statilia]]'', like many other arthropods, avoid attacking them. A variety of arthropods, including some early-instar mantises, exploit this behavior and mimic [[Biology:Ant mimicry|ants]] to evade their predators.<ref>{{Cite journal | last1=Nelson | first1=Ximena J. | last2=Jackson | first2=Robert R. | year=2006 | title=Innate aversion to ants (Hymenoptera: Formicidae) and ant mimics: experimental findings from mantises (Mantodea) | journal=Biological Journal of the Linnean Society | volume=88 | issue=1 | pages=23–32 | doi=10.1111/j.1095-8312.2006.00598.x| doi-access=free}}</ref>

<gallery mode="nolines" widths="180px">
Choeradodis stalii MHNT dos.jpg|Leaf [[Biology:Mimicry|mimicry]]: ''[[Biology:Choeradodis|Choeradodis]]'' has leaf-like forewings and a widened green [[Biology:Thorax (insect anatomy)|thorax]].
Gottesanbeterin Abwehr.JPG|Adult female ''[[Biology:Iris oratoria|Iris oratoria]]'' performs a bluffing threat display, rearing back with the forelegs and wings spread and mouth opened.
Gemmatus.jpg|The jeweled flower mantis, ''[[Biology:Creobroter gemmatus|Creobroter gemmatus]]'': the brightly colored wings are opened suddenly in a deimatic display to startle predators.
Ant-Mantis.jpg|Some mantis [[Biology:Nymph|nymphs]] [[Biology:Ant mimicry|mimic ants]] to [[Biology:Anti-predator adaptation|avoid predators]].
</gallery>

=== Reproduction and life history ===

The mating season in temperate climates typically takes place in autumn,<ref name="Ene 1964" /><ref name="Maxwell" /> while in tropical areas, mating can occur at any time of the year.<ref name="Maxwell">{{cite book |editor1=Prete, Fredrick R. |editor2=Wells, Harrington |editor3=Wells, Patrick H. |editor4=Hurd, Lawrence E. |title=The Praying Mantids |contribution=Mating Behavior |author=Michael, Maxwell. R. |year=1999 |publisher=Johns Hopkins University Press |isbn=978-0-8018-6174-1 |page=[https://archive.org/details/prayingmantids0000unse/page/70 70] |url=https://archive.org/details/prayingmantids0000unse/page/70}}</ref> 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 [[Biology:Ootheca|ootheca]]. Depending on the species, the ootheca can be attached to a flat surface, wrapped around a plant, or even deposited in the ground.<ref name="Ene 1964" /> Despite the versatility and durability of the eggs, they are often preyed on, especially by several species of [[Biology:Parasitoid wasp|parasitoid wasp]]s. In a few species, mostly ground and bark mantises in the family [[Biology:Tarachodidae|Tarachodidae]], the mother guards the eggs.<ref name="Ene 1964">{{cite journal |doi=10.1080/00222936408651488 |author=Ene, J. C. |year=1964 |title=The distribution and post-embryonic development of ''Tarachodes afzelli'' (Stal) (Mantodea : Eremiaphilidae) |journal=Journal of Natural History |volume=7 |issue=80 |pages=493–511}}</ref> The cryptic ''[[Biology:Tarachodes maurus|Tarachodes maurus]]'' positions herself on bark with her abdomen covering her egg capsule, ambushing passing prey and moving very little until the eggs hatch.<ref name="Costa" /> An unusual reproductive strategy is adopted by [[Biology:Brunneria borealis|Brunner's stick mantis]] from the southern United States: no males have ever been found in this species, and the females breed [[Biology:Parthenogenesis|parthenogenetically]].<ref name="Prete1999" /> The ability to reproduce by parthenogenesis has been recorded in at least two other species, ''[[Biology:Sphodromantis viridis|Sphodromantis viridis]]'' and ''Miomantis'' sp., although these species usually reproduce sexually.<ref>Bragg, P.E. (1987) A case of parthenogenesis in a mantis. ''Bulletin of the Amateur Entomologists' Society'', 46 (356): 160.</ref><ref>{{cite journal|last1=Bragg|first1=P. E.|title=Parthenogenetic mantid named|journal=Bulletin of the Amateur Entomologists' Society|url=http://mantodea.myspecies.info/parthenogenetic-mantid-named|publisher=Mantis Study Group|access-date=8 January 2017|date=December 1989|volume=48|issue=367|page=242}}</ref><ref>Dickie, S. (1996) Parthenogenesis in mantids. ''Mantis Study Group Newsletter'', 1: 5.</ref> In temperate climates, adults do not survive the winter and the eggs undergo a [[Biology:Diapause|diapause]], hatching in the spring.<ref name=Capinera2008 />

As in closely related insect groups in the superorder Dictyoptera, mantises go through three life stages: egg, nymph, and adult (mantises are among the [[Biology:Hemimetabolism|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 [[Biology:Ecdysis|molts]] its [[Biology:Exoskeleton|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.<ref name="Prete1999">{{cite book |editor1=Prete, Fredrick R. |editor2=Wells, Harrington |editor3=Wells, Patrick H. |editor4=Hurd, Lawrence E. |title=The Praying Mantids |contribution=Ecology of Praying Mantids |author=Hurd, I. E. |url=https://books.google.com/books?id=DpQtuB-EVSUC&pg=PA43 |year=1999 |publisher=Johns Hopkins University Press |isbn=978-0-8018-6174-1 |pages=43–49}}</ref><ref name="close up" />

<gallery mode="nolines" widths="180px" heights="180px">
File:Mantis religiosa couple.JPG|''Mantis religiosa'' mating (brown male, green female)
File:Mantis laying ootheca.jpg|''Stagmomantis carolina'' laying [[Biology:Ootheca|ootheca]]
File:Mantis religiosa (egg case).jpg|Recently laid ''M. religiosa'' ootheca
File:PrayingMantisesJustBorn (cropped).jpg|Hatching from the ootheca
File:Sphodromantis Lineola molting.jpg|''[[Biology:Sphodromantis lineola|Sphodromantis lineola]]'' molting
</gallery>

=== Sexual cannibalism ===
{{further|Biology: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.<ref>{{cite journal |last1=Lawrence |first1=S. E. |title=Sexual cannibalism in the praying mantid, ''Mantis religiosa'': a field study |journal=Animal Behaviour |year=1992 | volume=43 | issue=4 | pages=569–583 | doi=10.1016/S0003-3472(05)81017-6|s2cid=53199427}}</ref><ref name=Hurd>{{cite journal |last1=Hurd |first1=L. E. |author2=Eisenberg, R. M.|author3=Fagan, W. F.|author4=Tilmon, K. J.|author5=Snyder, W. E.|author6=Vandersall, K. S.|author7=Datz, S. G.|author8=Welch, J. D.|title=Cannibalism reverses male-biased sex ratio in adult mantids: female strategy against food limitation?|journal=Oikos |date=1994 | volume=69 | issue=2 | pages=193–198 |doi=10.2307/3546137 |jstor=3546137}}</ref><ref>{{cite journal |last1=Maxwell |first1=Michael R. | title=Lifetime mating opportunities and male mating behaviour in sexually cannibalistic praying mantids |journal=Animal Behaviour | volume=55 | issue=4 |pages=1011–1028 |year=1998 |doi=10.1006/anbe.1997.0671 |pmid=9632486 |s2cid=26887566}}</ref> Around 90% of the predatory species of mantises exhibit sexual cannibalism.<ref>{{cite journal |doi=10.1146/annurev.ecolsys.110308.120238 |title=The importance of ecological and phylogenetic conditions for the occurrence and frequency of sexual cannibalism |journal=Annual Review of Ecology, Evolution, and Systematics |volume=40 |pages=21–39 |year=2009 |last1=Wilder |first1=Shawn M. |last2=Rypstra |first2=Ann L. |last3=Elgar |first3=Mark A.}}</ref> Adult males typically outnumber females at first, but their numbers may be fairly equivalent later in the adult stage,<ref name=Capinera2008 /> possibly because females selectively eat the smaller males.<ref>{{cite web |title=Do Female Praying Mantises Always Eat the Males? |url=http://entomologytoday.org/2013/12/22/do-female-praying-mantises-always-eat-the-males/ |publisher=Entomology Today |access-date=31 July 2015 |date=22 December 2013}}</ref> 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.<ref name=Hurd />

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 [[Biology:Ganglion|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 ''[[Biology:Ad libitum|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.<ref name=Liske>{{cite journal |doi=10.1016/S0003-3472(84)80170-0 |title=Sexual behaviour of the Chinese praying mantis |journal=Animal Behaviour |volume=32 |issue=3 |pages=916–918 |year=1984 |last1=Liske |first1=E. |last2=Davis |first2=W. J.|s2cid=53144893}}</ref> 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.<ref name=Lelito>{{cite journal |author1=Lelito, Jonathan P. |author2=Brown, William D. |year=2006 |title=Complicity or conflict over sexual cannibalism? Male risk taking in the praying mantis ''Tenodera aridifolia sinensis'' |journal=The American Naturalist |volume=168 |issue=2 |pages=263–269 |doi=10.1086/505757 |pmid=16874635|s2cid=32185318 }}</ref>

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.<ref name=Liske2>{{cite journal |doi= 10.1016/s0003-3472(87)80024-6|title=Courtship and mating behaviour of the Chinese praying mantis, Tenodera aridifolia sinenesis |journal=Animal Behaviour |volume=35 |issue= 5|pages=1524–1537 |year=1987 |last1=Liske |first1=E. |last2=Davis |first2=W. J.|s2cid=53191218}}</ref> 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.<ref>{{cite journal |doi=10.1111/j.1365-2311.2010.01239.x |title=Effects of female feeding regime in a sexually cannibalistic mantid: fecundity, cannibalism, and male response in ''Stagmomantis limbata'' (Mantodea) |journal=Ecological Entomology |volume=35 |issue=6 |pages=775–787 |year=2010 |last1=Maxwell |first1=Michael R. |last2=Gallego |first2=Kevin M. |last3=Barry |first3=Katherine L.|s2cid=85923330 }}</ref> 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.<ref name=Lelito /> 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.<ref>{{cite journal |last1=Prokop |first1=P |last2=Václav |first2=R |title=Males Respond to the Risk of Sperm Competition in the Sexually Cannibalistic Praying Mantis, ''Mantis religiosa'' |journal=Ethology |date=2005 |volume=111 |issue=9 |pages=836–848|doi=10.1111/j.1439-0310.2005.01113.x }}</ref>

== 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 period, Japan, c. 1800]]

One of the earliest mantis references is in the ancient Chinese dictionary ''Erya'', which gives its attributes in poetry, where it represents courage and fearlessness, and a brief description. A later text, the ''Jingshi Zhenglei Daguan Bencao <!--經史證類大觀本草-->'' ("Great History of Medical Material Annotated and Arranged by Types, Based upon the Classics and Historical Works") from 1108, gives accurate details of the construction of the egg packages, the development cycle, anatomy, and the function of the antennae.
Although mantises are rarely mentioned in Ancient Greek sources, a female mantis in threat posture is accurately illustrated on a series of fifth-century BC silver coins, including didrachms, from Metapontum in [[Earth:Lucania|Lucania]].<ref name="Campbell2014">{{cite book |last=Bodson |first=Liliane |editor=Campbell, Gordon Lindsay |title=The Oxford Handbook of Animals in Classical Thought and Life |url=https://books.google.com/books?id=ofo5BAAAQBAJ&pg=PA557 |year=2014 |publisher=Oxford University Press |isbn=978-0-19-958942-5 |pages=557–558}}</ref>
In the 10th century AD, Byzantine era ''Adages'', Suidas describes an insect resembling a slow-moving green locust with long front legs.<ref>{{cite web |title=Entomomancy |url=http://www.occultopedia.com/e/entomomancy.htm |work=Occultopedia |publisher=California Astrology Association |access-date=30 July 2015}}</ref> He translates [[Biography:Zenobius|Zenobius]] 2.94 with the words ''seriphos'' (maybe a mantis) and ''graus'', an old woman, implying a thin, dried-up stick of a body.<ref name="ErasmusFantazzi1992">{{cite book|last1=Erasmus |first1=Desiderius |last2=Fantazzi |first2=Charles |title=Adages: Iivii1 to Iiiiii100 |url=https://books.google.com/books?id=vwf_62ogNK4C&pg=PA334 |year=1992 |publisher=University of Toronto Press |isbn=978-0-8020-2831-0 |pages=334–335}}</ref>

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".<ref name=":0">{{Cite journal|last1=McCafferty|first1=Geoffrey|last2=McCafferty|first2=Sharisse|date=1 April 2020|title=Praying to the Predator: Symbols of Insect Animism on Luna/El Menco Polychrome from Pre-Columbian Pacific Nicaragua|doi=10.1525/lavc.2020.220004|journal=Latin American and Latinx Visual Culture|volume=2|issue=2|pages=28–46|s2cid=225959388}}</ref>

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 ''{{lang|de|Insekten-Belustigungen}}'' (''Insect Entertainments'').<ref name="PreteWells" />

[[File:Wandering Violin Mantis (Gongylus gongylodes) Photograph By Shantanu Kuveskar.jpg|thumb|upright|In ''Island'', Aldous Huxley reflected on death as a pair of ''Gongylus gongylodes'' mated.|alt=The thin-legged mantis ''Gongylus gongylodes'']]

In the early 1900s, people in the United States Ozarks region referred to them as '''Devil's horses.'''<ref>{{Cite book |last=Randolph |first=Vance |url=https://books.google.com/books?id=txnhppn0SrwC&dq=People+near+Natural+Dam,+Arkansas,+told+me+that+the+Devil%E2%80%99s+horse,+or+praying+mantis,+is+deadly+poison,+and+that+a+boy+near+that+place+died+as+a+result+of+its+bite.+Local+physicians+laughed+at+this+story;+one+doctor+said+that+he+didn%E2%80%99t+know+whether+or+not+the+Devil%E2%80%99s+horse+was+poisonous,+but+he+knew+damned+well+that+it+had+never+killed+anybody+in+his+neighborhood.+Children+in+the+Ozarks+are+often+told,+however,+that+it+is+bad+luck+to+%E2%80%9Cpester%E2%80%9D+a+Devil%E2%80%99s+horse,+as+the+creature+is+likely+to+spit+tobacco+juice+in+one%E2%80%99s+eye+and+perhaps+cause+blindness&pg=PA259 |title=Ozark Magic and Folklore |date=2012-07-31 |publisher=Courier Corporation |isbn=978-0-486-12296-0 |language=en}}</ref>

[[Biography:Aldous Huxley|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 ''[[Biology:Gongylus gongylodes|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 [[Biology:Gecko|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.<ref>{{cite book |last1=Durrell |first1=Gerald |title=My Family and Other Animals |url=https://books.google.com/books?id=Q3dJACk-gsoC |year=2006 |orig-year=1956 |publisher=Penguin Books |isbn=978-0-14-193609-3 |pages=204–208}}</ref><!--end of ref-->}}<!--end quote-->

M. C. Escher's woodcut ''Dream'' depicts a human-sized mantis standing on a sleeping bishop.<ref name="EmmerSchattschneider2007">{{cite book |last1=Emmer |first1=Michele |last2=Schattschneider |first2=Doris |title=M.C. Escher's Legacy: A Centennial Celebration|url=https://books.google.com/books?id=5DDyBwAAQBAJ&pg=PA66 |year=2007 |publisher=Springer |isbn=978-3-540-28849-7 |page=66}}</ref> 

A cultural trope imagines the female mantis as a ''[[Social:Femme fatale|femme fatale]]''. The idea is propagated in cartoons by Cable, Guy and Rodd, LeLievre, T. McCracken, and Mark Parisi, among others.<ref name=Wade>{{cite web |last1=Wade |first1=Lisa |title=Shoddy Research and Cultural Tropes: The Praying Mantis |url=http://thesocietypages.org/socimages/2010/11/29/shoddy-research-and-cultural-tropes-the-case-of-the-praying-mantis/ |publisher=The Society Pages |access-date=9 October 2015 |date=29 November 2010}}</ref><ref>{{cite web |title=Praying Mantises Cartoons and Comics |url=http://www.cartoonstock.com/directory/p/praying_mantises.asp |publisher=CartoonStock |access-date=30 July 2015}}</ref><ref>{{cite web |last1=Parisi |first1=Mark |url=https://www.offthemark.com/index.php?_route_=cartoons/praying+mantis//key/praying+mantis/ |title=Praying Mantis Cartoons |publisher=Off the Mark |access-date=30 July 2015 |url-status=dead |archive-url=https://web.archive.org/web/20150925144024/https://www.offthemark.com/index.php?_route_=cartoons%2Fpraying+mantis%2F%2Fkey%2Fpraying+mantis%2F |archive-date=25 September 2015 }}</ref><ref>{{cite web |last1=McCracken |first1=T. |title=Praying Mantis Should Be Gay |url=https://mchumor.com/cartoon/6814-insect-cartoon/ |publisher=McHumor Cartoons |access-date=30 July 2015 |archive-date=5 March 2016 |archive-url=https://web.archive.org/web/20160305110846/https://mchumor.com/cartoon/6814-insect-cartoon/ |url-status=dead }}</ref> It ends Isabella Rossellini's short film about the life of a praying mantis in her 2008 ''Green Porno'' season for the Sundance Channel.<ref>{{cite web |last1=Rossellini |first1=Isabella |title=Green Porno |url=https://www.youtube.com/watch?v=eePxQM2aWH4  |archive-url=https://ghostarchive.org/varchive/youtube/20211215/eePxQM2aWH4 |archive-date=15 December 2021 |url-status=live|publisher=Sundance Channel |access-date=1 September 2015|date=2008}}{{cbignore}}</ref><ref name="press09">{{cite web | last=Rossellini | first=Isabella | title=Green Porno: a series of short films by Isabella Rossellini | publisher=Sundance Channel | year=2009 | url=http://www.sundance.tv/series/greenporno | format=Press Kit | access-date=1 September 2015}}</ref><!--British goth band [[Engineering:Bauhaus|Bauhaus]]' song "The Passion of Lovers" contains several metaphoric references to mantises (particularly sexual cannibalism), and the artwork associated with the single is a photo of a female mantis devouring her mate.-->

''The Deadly Mantis'' is a 1957 American science fiction monster film, with a giant mantis threatening mankind.<ref>[https://www.imdb.com/title/tt0050294/ Imdb]</ref>
<!--Please do not add any trivia here, nor anything without citing a reliable source that actually discusses the item. This is not a list of "and another one appeared in xxx", that has no value.-->

=== Martial arts ===
[[File:Shi DeRu and Shi DeYang.jpg|thumb|Grandmasters of the Shaolin Temple, Shi DeRu and Shi DeYang, demonstrating the Southern Praying Mantis style of martial art]]

Two martial arts separately developed in China have movements and fighting strategies based on those of the mantis.<ref name=Academy>{{cite web |title=History of Praying Mantis Kung Fu |url=http://www.prayingmantis.com.au/prayingmantis.php |archive-url=https://web.archive.org/web/20151001110604/http://www.prayingmantis.com.au/prayingmantis.php |archive-date=1 October 2015 |publisher=Praying Mantis Kung Fu Academy |access-date=30 September 2015}}</ref><ref name=Hagood /> 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'<ref name=Funk>{{cite web |last1=Funk |first1=Jon |title=Praying Mantis Kung Fu |url=http://www.mantiskungfu.com/praying_mantis_kung_fu_complete_system.php |publisher=Praying Mantis Kung Fu |access-date=31 July 2015}}</ref> and 'Southern Praying Mantis'.<ref name=Hagood>{{cite book |last1=Hagood |first1=Roger D. |title=18 Buddha Hands: Southern Praying Mantis Kung Fu |date=2012 |publisher=Southern Mantis Press |isbn=978-0-9857240-1-6}}</ref> Both are very popular in China, and have also been exported to the West in recent decades.<ref name=Hagood /><ref name=Funk /><ref name=Barnes>{{cite web |last1=Barnes |first1=Bryan |title=Northern Praying Mantis Kung Fu |url=http://www.northernprayingmantiskungfu.com/ |publisher=Shantung Northern Praying Mantis Kung Fu |access-date=30 September 2015}}</ref><ref name=Orum>{{cite web |last1=Orum |first1=Pete |title=Chow Gar |url=http://www.southernmantiskungfu.co.uk/ |publisher=Southern Mantis Kung Fu |access-date=30 September 2015}}</ref>

=== In mythology and religion ===
According to local beliefs in Africa, this insect brings good luck.<ref>[https://www.reference.com/pets-animals/praying-mantis-good-luck-7d4194e515defa22 Is a Praying Mantis Good Luck?], Reference.com, 11 April 2020</ref> 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 ''{{lang|af|Hottentotsgot}}'' ("god of the Hottentots") in the [[Social:Afrikaans|Afrikaans]] language that had developed among the first European settlers.<ref>{{cite web |title=Insek-kaleidoskoop: Die 'skynheilige' hottentotsgot |url=http://www.mieliestronk.com/insek7hotten.html |website=Mieliestronk.com |access-date=9 October 2015 |archive-date=24 May 2019 |archive-url=https://web.archive.org/web/20190524230352/http://www.mieliestronk.com/insek7hotten.html |url-status=dead }}</ref> However, at least for the San, the mantis was only one of the manifestations of a trickster-deity, [[Religion:ǀKaggen|ǀKaggen]], who could assume many other forms, such as a snake, hare or vulture.<ref>{{cite web |title=Siyabona Africa: Kruger National Park: San |url=http://www.krugerpark.co.za/africa_bushmen.html |website=Siyabona Africa |access-date=30 June 2017}}</ref> 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 ''[[Religion:Book of the Dead|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 [[Place:Nineveh|Nineveh]] grasshoppers (''buru''), the mantis is named [[Unsolved:Necromancy|necromancer]] (''buru-enmeli'') and soothsayer (''buru-enmeli-ashaga'').<ref name="PreteWells">{{cite book |editor1=Prete, Fredrick R. |editor2=Wells, Harrington |editor3=Wells, Patrick H. |editor4=Hurd, Lawrence E. |title=The Praying Mantids |contribution=The Predatory Behavior of Mantids: Historical Attitudes and Contemporary Questions|author1=Prete, Frederick R.|author2=Wells, Harrington|author3=Wells, Patrick H. |name-list-style=amp |url=https://books.google.com/books?id=DpQtuB-EVSUC&pg=PA4 |year=1999 |publisher=Johns Hopkins University Press |isbn=978-0-8018-6174-1 |pages=3–8}}</ref><ref>{{cite encyclopedia |title=Mantid |url=https://www.britannica.com/animal/mantid |encyclopedia=Encyclopædia Britannica |access-date=30 July 2015}}</ref> 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.<ref name=":0" />

=== As pets ===
[[File:Gray adult female Carolina Mantis.JPG|thumb|Gray adult female [[Biology:Carolina mantis|Carolina mantis]] in human hand]]

Mantises are among the insects most widely kept as pets.<ref name=Kentucky>{{cite web |title=Pet bugs: Kentucky arthropods |url=http://www.uky.edu/Ag/CritterFiles/casefile/bugconnection/petbugs/Kentucky/Kentucky.htm#mantid |publisher=University of Kentucky |access-date=31 July 2015}}</ref><ref name=VanZomeren>{{cite web |last1=Van Zomeren |first1=Linda |title=European Mantis |date=16 March 2011 |url=http://www.keepinginsects.com/praying-mantis/species/european-mantis/ |publisher=Keeping Insects |access-date=31 July 2015}}</ref> 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.<ref name=UKmantis>{{cite report |title=Breeding Reports: 1st July 2013–1st October 2013 |type=Newsletter |url=http://www.ukmantisforums.co.uk/newsletters/Oct2013.pdf |publisher=UK Mantis Forums |work=UK Mantis Forums Newsletter |issue=12 |access-date=31 July 2015 |date=October 2013 |url-status=dead |archive-url=https://web.archive.org/web/20150923103205/http://www.ukmantisforums.co.uk/newsletters/Oct2013.pdf |archive-date=23 September 2015 }}</ref> In 1996 at least 50 species were known to be kept in captivity by members of the Mantis Study Group.<ref>Bragg, P.E. [editor] (1996) Species in culture. ''Mantis Study Group Newsletter'', '''1''': 2–3.</ref> ''The Independent'' described the "giant Asian praying mantis" as "part stick insect with a touch of Buddhist monk".<ref name="Indie">{{cite news |last1=Buckley |first1=Jamie |date=17 October 2009 |title=Pet of the week: The giant Asian praying mantis |newspaper=The Independent |url=https://www.independent.co.uk/property/house-and-home/pets/features/pet-of-the-week-the-giant-asian-praying-mantis-1803198.html |access-date=8 October 2015}}</ref>

=== For pest control ===

Naturally occurring mantis populations provide plant pest control.<ref name="Zhang-et-al-2007">{{cite journal | last1=Zhang | first1=Wei | last2=Ricketts | first2=Taylor H. | last3=Kremen | first3=Claire | last4=Carney | first4=Karen | last5=Swinton | first5=Scott M. | title=Ecosystem services and dis-services to agriculture | journal=Ecological Economics | publisher=[[Organization:International Society for Ecological Economics|International Society for Ecological Economics]] ([[Company:Elsevier|Elsevier]]) | volume=64 | issue=2 | year=2007 | issn=0921-8009 | doi=10.1016/j.ecolecon.2007.02.024 | pages=253–260 | s2cid=32112906| url=https://repositorio.unal.edu.co/handle/unal/82151 }}</ref> Gardeners who prefer to avoid [[Biology:Pesticide|pesticide]]s may encourage mantises in the hope of controlling insect pests.<ref name=Doutt>{{cite web |last1=Doutt |first1=R. L. |title=The Praying Mantis (Leaflet 21019)|url=http://vric.ucdavis.edu/pdf/PESTS/pests_PrayingMantis.pdf |publisher=University of California Division of Agricultural Sciences |access-date=31 July 2015}}</ref> However, mantises do not have key attributes of [[Biology:Biological pest control|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.<ref name=Doutt />

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.<ref>{{cite web |title=Praying and Chinese Mantises |url=http://www.dec.ny.gov/animals/58926.html |publisher=New York State Department of Environmental Conservation |access-date=26 August 2015}}</ref>

=== Mantis-like robot ===

A prototype [[Engineering:Robot|robot]] inspired by the forelegs of the praying mantis has 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.<ref>{{cite web |last1=Cardona |first1=Ramon |last2=Touretzky |first2=David |title=Leg Design for a Praying Mantis Robot |url=https://www.aaai.org/ocs/index.php/FLAIRS/FLAIRS12/paper/viewFile/4438/4855 |website=Association for the Advancement of Artificial Intelligence |publisher=Florida Artificial Intelligence Research Society |access-date=12 April 2016 |archive-date=24 June 2016 |archive-url=https://web.archive.org/web/20160624070612/https://www.aaai.org/ocs/index.php/FLAIRS/FLAIRS12/paper/viewFile/4438/4855 |url-status=dead }}</ref>

==See also==
* List of Orthopteroid genera containing species recorded in Europe
{{Clear}}

==References==
{{Reflist|28em}}

== External links ==
{{EB1911 poster|Mantis}}
* [http://mantodea.myspecies.info/ Mantis Study Group] – Information on mantises, phylogenetics and evolution.
* [http://mantodea.speciesfile.org/Common/basic/Taxa.aspx?TaxonNameID=1182267 Mantodea Species File]

{{Orders of Insects}}
{{Mantodea}}
{{Insects in culture}}
{{Taxonbar|from=Q131250}}

[[Category:Mantodea| ]]
[[Category:Extant Cretaceous first appearances]]

{{Sourceattribution|Mantis}}