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{{Short description|Family of insects}}
{{About|the insect}}
{{Good article}}
{{Automatic taxobox
| name = Antlions
| fossil_range = {{fossil range|122|0}} [[Early Cretaceous]] – Recent
| image = Distoleon tetragrammicus01.jpg
| image_caption = ''[[Distoleon tetragrammicus]]''
| image2 = Myrmeleontidae - Distoleon tetragrammicus.JPG
| image2_caption = ''Distoleon tetragrammicus'' larva
| taxon = Myrmeleontidae
| authority = [[Pierre André Latreille|Latreille]], 1802
| subdivision_ranks = Subfamilies
| subdivision = see [[#Taxonomy|text]]
}}

The '''antlions''' are a group of about 2,000 [[species]] of [[insect]] in the [[neuroptera]]n [[family (biology)|family]] '''Myrmeleontidae'''. They are known for the [[predation|predatory]] habits of their [[larva]]e, which mostly dig pits to trap passing [[ant]]s or other prey. In North America, the larvae are sometimes referred to as '''doodlebugs''' because of the marks they leave in the sand. The adult insects are less well known due to their relatively short lifespans in comparison with the larvae. Adults, sometimes known as '''antlion lacewings''', mostly fly at dusk or just after dark and may be mistakenly identified as [[dragonfly|dragonflies]] or [[damselfly|damselflies]].

Antlions have a worldwide distribution. The greatest diversity occurs in the tropics, but a few species are found in cold-[[temperate]] locations, one such being the European ''[[Euroleon nostras]]''. They most commonly occur in dry and sandy habitats where the larvae can easily excavate their pits, but some larvae hide under debris or ambush their prey among [[leaf litter]].

Antlions are poorly represented in the [[fossil|fossil record]]. [[Myrmeleontiformia]] is generally accepted to be a [[monophyly|monophyletic]] group, and within the [[Myrmeleontoidea]], the antlions' closest living relatives are thought to be the [[owlfly|owlflies]] (Ascalaphidae). A 2019 study finds Myrmeleontidae to be monophyletic, aside from [[Stilbopteryginae]] and [[Palparinae]], which form separate clades closer to Ascalaphidae.<ref>{{cite journal |last=Jones |first=Joshua R. |date=November 2019 |title=Total-evidence phylogeny of the owlflies (Neuroptera, Ascalaphidae) supports a new higher-level classification |journal=Zoologica Scripta |language=en |volume=48 |issue=6 |pages=761–782 |issn=0300-3256 |doi=10.1111/zsc.12382 |doi-access=free |s2cid=208581837}}</ref> The predatory actions of the larvae have attracted attention throughout history and antlions have been mentioned in literature since classical times.

==Etymology==
[[File:Doodlebug doodles.jpg|thumb|left|Doodlebug (antlion) doodles and pit traps in the Grand Canyon]]

The exact meaning of the name "antlion" is uncertain. It has been thought to refer to ants forming a large percentage of the prey of the insect, the suffix "lion" merely suggesting "destroyer" or "hunter".<ref name="EB1911">{{EB1911|inline=y|wstitle=Ant-lion|volume=2|page=147|last=Pocock|first=Reginald Innes|author-link=R. I. Pocock}}</ref> In any case, the term seems to go back to [[classical antiquity]].<ref name=Swanson2007>{{cite web |title="Antlion" in the World's Languages |author=Swanson, Mark |year=2007 |work=The Antlion Pit |url=https://www.antlionpit.com/language.html |access-date=23 February 2016}}</ref> The antlion larva is often called a "doodlebug" in [[North America]] because of the odd winding, spiralling trails it leaves in the sand while relocating, which look as if someone has been [[doodle|doodling]].<ref name=Swanson>{{cite web |title=What are Antlions? |author=Swanson, Mark |year=2007 |website=The Antlion Pit |url=https://www.antlionpit.com/what.html |access-date=23 February 2016}}</ref>

The [[scientific name]] of the [[type genus]] ''Myrmeleo''&nbsp;– and thus, the family as a whole&nbsp;– is derived from [[Ancient Greek]] ''mýrmex'' (μύρμηξ) "ant" + ''léon'' (λέων) "lion", in a [[calque|loan translation]] of the names common across Europe. In most European and Middle Eastern languages, at least the larvae are known under the local term corresponding to "antlion".<ref name=Swanson2007/>

==Description==
[[File:Antlion (Myrmecaelurus trigrammus) Macedonia.jpg|thumb|Adult of ''Myrmecaelurus trigammus''|left]]
[[File:Common Antlion Myrmeleon immaculatus mature nymph.jpg|left|thumb|Larva of ''[[Myrmeleon immaculatus]]'']]
Antlions can be fairly small to very large [[neuroptera]]ns, with wingspans ranging from {{convert|2|to|15|cm|in|1|abbr=on}}. The African genus ''[[Palpares]]'' contains some of the largest examples. ''[[Acanthaclisis occitanica]]'' is the largest European species, with an {{convert|11|cm|in|1|abbr=on}} wingspan, and most North American species approach this size.<ref name=Swanson/>

The adult has two pairs of long, narrow, multiveined, translucent [[insect wing|wings]] and a long, slender [[abdomen]]. Although they somewhat resemble [[dragonfly|dragonflies]] or [[damselfly|damselflies]], they belong to a different [[class (biology)|infraclass]] of [[pterygota|winged insects]]. Antlion adults are easily distinguished from damselflies by their prominent, apically clubbed [[antenna (biology)|antennae]] which are about as long as the head and [[thorax (insect anatomy)|thorax]] combined.<ref name=Mares>{{cite book |author1=Mares, Michael A. |title=Encyclopedia of Deserts |year=1999 |publisher=University of Oklahoma Press |isbn=978-0-8061-3146-7 |page=29 |url=https://books.google.com/books?id=g3CbqZtaF4oC&pg=PA29}}</ref> Also, the pattern of wing venation differs, and compared to damselflies, the adults are very feeble fliers and are normally found fluttering about at night in search of a mate. Adult antlions are typically [[nocturnality|nocturnal]], and rarely seen by day.<ref name="EB1911"/>

Males of most species have a unique structure, a bristle-bearing knob known as a "pilula axillaris", at the base of the rear wing. The abdomen in males is usually longer than in females and often has an extra lobe. The tip of the abdomen of females shows greater variation than that of males, depending perhaps on [[ovipositor|oviposition]] sites, and usually bears tufts of bristles for digging and a finger-like extension.<ref name=Miller>{{cite web |title=''Glenurus gratus'' (Say) (Insecta: Neuroptera: Myrmeleontidae) |author1=Miller, Robert B. |author2=Stange, Lionel A. |date=1 November 2015 |work=Featured Creatures |publisher=University of Florida |url=https://entnemdept.ufl.edu/creatures/misc/neuroptera/Glenurus_gratus.htm |access-date=22 February 2016}}</ref>
[[File:Antlion mandible with a visible maxilla for injecting venom.jpg|left|thumb|225x225px|The bottom side of larval mandible with a visible maxilla for injecting venom]]
The antlion [[larva]] has a robust [[wikt:fusiform|fusiform]] body, a very plump abdomen, and a thorax bearing three pairs of walking legs. The [[prothorax]] forms a slender mobile "neck" for the large, square, flattened head, which bears an enormous pair of sickle-like jaws with several sharp, hollow projections. The jaws are formed by the [[insect mouthparts|maxillae and mandibles]]; the mandibles each contain a deep groove over which the maxilla fits neatly, forming an enclosed canal for injecting [[venom]] to immobilise the victim,<ref name=Hawkeswood>{{cite journal |author=Hawkeswood, Trevor J. |year=2006 |title=Effects of envenomation to a human finger and arm by the larva of an unidentified species of ''Myrmeleon'' (Neuroptera: Myrmeleontidae) |journal=Calodema |volume=7 |pages=32–33 |url=http://www.calodema.com/freefiles/307.pdf |url-status=usurped |archive-url=https://web.archive.org/web/20160305041826/http://www.calodema.com/freefiles/307.pdf |archive-date=2016-03-05}}</ref><ref name=Nardi>{{cite book |author=Nardi, James B. |title=Life in the Soil: A Guide for Naturalists and Gardeners |year=2009 |publisher=University of Chicago Press |isbn=978-0-226-56853-9 |page=170 |url=https://books.google.com/books?id=d0MXBkFY07UC&pg=PA170}}</ref> and [[enzyme]]s to digest its soft parts. The larva is clad in forward-pointing bristles which help it to anchor itself and exert greater traction, enabling it to subdue prey considerably larger than itself.<ref name=Camp>{{cite web |title=Beneficials in the garden: Antlion |author=Camp, Donya |year=2005 |work=Galveston County Master Gardeners |publisher=Extension Horticulture at Texas A&M University |url=http://aggie-horticulture.tamu.edu/galveston/beneficials/beneficial-32_ant_lion.htm |access-date=28 February 2016}}</ref>

Antlion larvae are unusual among insects in lacking an [[anus]]. All the [[metabolism|metabolic]] waste generated during the larval stage is stored; some is used to spin the silk for the [[pupa#Cocoon|cocoon]] and the rest is eventually voided as [[meconium]] at the end of its [[pupa]]l stage.<ref name="SwansonDiscard">{{cite web |title=Antlion Larvae Behavior: Discarding the Body |author=Swanson, Mark |year=2007 |website=The Antlion Pit |url=https://www.antlionpit.com/discard.html |access-date=24 February 2016}}</ref>

==Distribution==
There are about 2,000 species of antlion found in most parts of the world, with the greatest diversity being in warmer areas. The best known species are those in which the larvae dig pits to trap their prey, but not all species do this. Antlions live in a range of usually dry habitats including open woodland floors, [[scrubland|scrub]]-clad [[dune]]s, hedge bases, river banks, road verges, under raised buildings and in vacant lots.<ref name=SwansonHabitat>{{cite web |title=Where do Antlions Live? |author=Swanson, Mark |year=2007 |website=The Antlion Pit |url=https://www.antlionpit.com/where.html |access-date=24 February 2016}}</ref>
[[File:Antlion sp. from Sri Lanka.jpg|right|thumb|from Sri Lanka]]

==Life-cycle==
[[File:Antlion life cycle.svg|thumb|upright=0.75|left|The life-cycle of ''[[Distoleon tetragrammicus]]'']]

Apart from pit-trap-forming taxa, the biology of members of the family Myrmeleontidae, to which the antlions belong, has been little studied. The life-cycle begins with oviposition ([[egg]]-laying) in a suitable location. The female antlion repeatedly taps the prospective laying site with the tip of her abdomen and then inserts her ovipositor into the [[substrate (biology)|substrate]] and lays an egg.<ref name=McEwen>{{cite book |author1=McEwen, P.K. |author2=New, T.R. |author3=Whittington, A.E. |title=Lacewings in the Crop Environment |year=2007 |publisher=Cambridge University Press |isbn=978-0-521-03729-7 |page=4 |url=https://books.google.com/books?id=ZIfF6ho1ac4C&pg=PA4}}</ref>

Depending on the species and where it lives, the larva either conceals itself under leaves, debris or pieces of wood, hides in a crack or digs a funnel-shaped pit in loose material.<ref name=Hawkeswood/> As [[ambush predator]]s, catching prey is risky because food arrives unpredictably and, for those species that make traps, maintaining one is costly. The larvae therefore have low metabolic rates and can survive for long periods without food.<ref name=Jervis>{{cite book |author=Jervis, Mark A. |title=Insects as Natural Enemies: A Practical Perspective |year=2007 |publisher=Springer Science & Business Media |isbn=978-1-4020-6587-3 |page=28 |url=https://books.google.com/books?id=RsVt_inIAJUC&pg=PA28}}</ref> They can take several years to complete their life-cycle; they mature faster with plentiful food, but can survive for many months without feeding.<ref>{{cite book |title=Insects as Predators |author=New, T. |publisher=NSW University Press |date=1991 |page=69}}</ref><ref name=SwansonReproduction>{{cite web |last1=Swanson |first1=Mark |title=Reproductive Behavior |website=The Antlion Pit |date=2012 |url=https://www.antlionpit.com/reproduce.html#span |access-date=23 February 2016 |url-status=live |archive-url=https://web.archive.org/web/20160304141617/http://www.antlionpit.com/reproduce.html#span |archive-date=4 March 2016}}<!--Website is properly cited, and mentioned by Coelho--></ref> In cooler climates they dig their way deeper and remain inactive during the winter.<ref name=SwansonDiscard/>

[[File:Ant Lion Cocoon.jpg|thumb|upright=0.75|An antlion cocoon on the side of a house]]
When the larva attains its maximum size, it [[pupa]]tes and undergoes [[metamorphosis]].<ref name=SwansonReproduction/> It makes a globular cocoon of sand or other local substrate stuck together with fine [[silk]] spun from a slender [[spinneret (spider)|spinneret]] at the rear end of the body. The cocoon may be buried several centimetres deep in sand. After completing its transformation into an adult insect over the course of about one month, it emerges from the case, leaving the pupal integument behind, and works its way to the surface. After about twenty minutes, the adult's wings are fully opened and it flies off in search of a mate. The adult is considerably larger than the larva as antlions exhibit the greatest disparity in size between larva and adult of any type of [[holometabolism|holometabolous]] insect. This is by virtue of the fact that the [[exoskeleton]] of the adult is extremely thin and flimsy, with an exceptionally low [[density]].<ref name="SwansonMetamorphosis">{{cite web |last1=Swanson |first1=Mark |title=Metamorphosis |website=The Antlion Pit |date=2012 |url=https://www.antlionpit.com/metamor.html |access-date=23 February 2016}}</ref> The adult typically lives for about 25 days, but some insects survive for as long as 45 days.<ref name=SwansonReproduction/>

==Ecology==
[[File:Antlion trap.jpg|thumb|Sand pit trap of an antlion]]
Antlion [[larva]]e eat small arthropods – mainly [[ant]]s – while the adults of some species eat [[pollen]] and [[nectar]], and others are predators of small arthropods.<ref name=Engel2007>{{cite journal |last1=Engel |first1=Michael S. |last2=Grimaldi |first2=David A. |year=2007 |title=The neuropterid fauna of Dominican and Mexican amber (Neuropterida, Megaloptera, Neuroptera) |journal=American Museum Novitates |issue=3587 |pages=1–58 |doi=10.1206/0003-0082(2007)3587[1:tnfoda]2.0.co;2 |s2cid=49393365 |url=http://digitallibrary.amnh.org/dspace/bitstream/2246/5880/1/N3587.pdf}}</ref> In certain species of Myrmeleontidae, such as ''[[Dendroleon pantherinus]]'', the larva, although resembling that of ''[[Myrmeleon]]'' structurally, makes no pitfall trap, but hides in [[wikt:detritus|detritus]] in a hole in a tree and seizes passing prey.<ref>{{cite journal |author1=Devetak, Dušan |author2=Podlesnik, Jan |author3=Janževocič, Franc |year=2014 |title=Antlion ''Dendroleon pantherinus'' (Fabricius, 1787) (Neuroptera: Myrmeleontidae) in Slovenia |journal=Acta Entomologica Slovenica |volume=18 |issue=2 |pages=159–162 |url=https://www.researchgate.net/publication/261497922}}</ref> In Japan, ''[[Gatzara jezoensis]]'' larvae lurk on the surface of rocks for several years while awaiting prey; during this time they often become coated with [[lichen]], and have been recorded at densities of up to 344 per square metre.<ref>{{cite web |title=Predatory insects |author=O'Neal, Matt |publisher=MSU |url=https://www.msu.edu/course/ent/815/webpredation.pdf |access-date=23 February 2016}}</ref>

The larva is a voracious predator. Within a few minutes of seizing its prey with its jaws and injecting it with [[venom]] and [[enzyme]]s, it begins to suck out the digestion products.<ref name=Hawkeswood/><ref name=Camp/> The larva is extremely sensitive to ground vibrations, the low-frequency sounds made by an insect crawling across the ground; the larva locates the source of the vibrations by the differences in timing of the arrival of [[wave]]s detected by [[sensory receptor|receptor]]s, tufts of hairs on the sides of the two hindmost thoracic segments.<ref name=Cocroft>{{cite book |author=Cocroft, Reginald Bifield |title=Studying Vibrational Communication |year=2014 |publisher=Springer |isbn=978-3-662-43607-3 |page=319 |url=https://books.google.com/books?id=snMrBAAAQBAJ&pg=PA319}}</ref>

===Pit-building species===
[[File:Antlion larva.jpg|thumb|Antlion larva extracted from sand pit]]
Funnel-shaped pits are built by members of just 3 antlion tribes: [[Myrmeleontini]], [[Myrmecaelurini]], and [[Nesoleontini]].<ref name="Machado2018"/> In these trap-building species, an average-sized larva digs a pit about 2&nbsp;in (5&nbsp;cm) deep and 3&nbsp;in (7.5&nbsp;cm) wide at the edge. This behavior has also been observed in the [[Vermileonidae]] ([[Diptera]]), whose larvae dig the same sort of pit to feed on ants. Having marked out the chosen site with a circular groove,<ref>{{cite journal |last1=Scharf |first1=Inon |last2=Ovadia |first2=Ofer |year=2006 |title=Factors influencing site abandonment and site selection in a sit-and-wait Predator: A review of pit-building antlion Larvae |journal=Journal of Insect Behavior |volume=19 |issue=2 |pages=197–218 |bibcode=2006JIBeh..19..197S |citeseerx=10.1.1.565.1350 |doi=10.1007/s10905-006-9017-4 |s2cid=36532488 |url=http://www.bgu.ac.il/~oferovad/library/Scharf%26Ovadia_2006.pdf}}</ref> the antlion larva begins to crawl backward, using its abdomen as a plough to dig up the soil. Using one of its front legs, it places heaps of loosened particles upon its head which it then flicks clear of the scene of operations. Continuing in this way, it gradually works its way from the circumference toward the center.<ref name="EB1911"/> As it slowly moves round and round, the pit gradually gets deeper and deeper, until the slope angle reaches the [[critical angle of repose]] (that is, the steepest angle the sand can maintain, where it is on the verge of collapse from slight disturbance), and the pit is solely lined by fine grains.<ref name=Franks/> By digging in a spiral when constructing its pit, the antlion minimises the time needed to complete the pit.<ref name=Franks>{{cite journal |title=Digging the optimum pit: antlions, spirals and spontaneous stratification |author1=Franks, Nigel R. |author2=Worley, Alan |author3=Falkenberg, Max |author4=Sendova-Franks, Ana B. |author5=Christensen, K |journal=Proceedings of the Royal Society B: Biological Sciences |date=2019 |volume=286 |issue=1899 |article-number=20190365 |pmid=30900535 |doi=10.1098/rspb.2019.0365 |pmc=6452065}}</ref>

[[File:Antlion pincers in Martin Frobenius Ledermüller’s Microscopic Delights (1759–63).jpg|thumb|Microscopic illustration of an antlion larva's pincer by [[Martin Frobenius Ledermüller]] (1759–1763), showcasing the serrated jaws used to seize prey at the bottom of its sand trap.]]

When the pit is completed, the larva settles down at the bottom, buried in the soil with only the jaws projecting above the surface, often in a wide-opened position on either side of the very tip of the cone.<ref name=Coelho>{{cite web |title=The natural history and ecology of antlions (Neuroptera: Myrmeleontidae) |author=Coelho, Joseph R. |url=https://www.academia.edu/1082341 |access-date=23 February 2016}}</ref> The steep-sloped trap that guides prey into the larva's mouth while avoiding crater avalanches is one of the simplest and most efficient traps in the animal kingdom.<ref>{{cite journal |last1=Fertin |first1=A. |title=Efficiency of antlion trap construction |journal=Journal of Experimental Biology |date=2006 |volume=209 |issue=18 |pages=3510–3515 |pmid=16943491 |doi=10.1242/jeb.02401 |doi-access=free}}</ref> The fine grain lining ensures that the avalanches which carry prey are as large as possible.<ref name=Franks/>
Since the sides of the pit consist of loose sand at its angle of repose,<ref>{{cite journal |last1=Botz |first1=Jason T. |last2=Loudon |first2=Catherine |last3=Barger |first3=J. Bradley |last4=Olafsen |first4=Jeffrey S. |last5=Steeples |first5=Don W. |year=2003 |title=Effects of slope and particle size on ant locomotion: Implications for choice of substrate by antlions |journal=Journal of the Kansas Entomological Society |volume=76 |issue=3 |pages=426–435 |url=http://cat.inist.fr/?aModele=afficheN&cpsidt=14973602}}</ref> they afford an insecure foothold to any small insects that inadvertently venture over the edge, such as ants. Slipping to the bottom, the prey is immediately seized by the lurking antlion; if it attempts to scramble up the treacherous walls of the pit, it is speedily checked in its efforts and brought down by showers of loose sand which are thrown at it from below by the larva.<ref name="EB1911"/> By throwing up loose sand from the bottom of the pit, the larva also undermines the sides of the pit, causing them to collapse and bring the prey with them. Thus, it does not matter whether the larva actually strikes the prey with the sand showers.<ref name=Coelho/>

Antlion larvae are capable of capturing and killing a variety of [[insect]]s and other [[arthropod]]s, and can even subdue small [[spider]]s. The projections in the jaws of the larva are hollow and through this, the larva sucks the fluids out of its victim. After the contents are consumed, the dry carcass is flicked out of the pit. The larva readies the pit once again by throwing out collapsed material from the center, steepening the pit walls to the angle of repose.<ref name=Coelho/>

[[File:AmeisenlöwenTrichter.jpg|thumb|Sand pit trap with remains of an ant]]
Antlion pit construction requires loose soil, sand being a common choice. Antlions can also handle larger granular material which is filtered out of the soil during construction.<ref name=Franks/> The larvae prefer dry places protected from the rain. When it first hatches, the tiny larva specialises in very small insects, but as it grows larger, it constructs larger pits, and thus catches larger prey, sometimes much larger than itself.<ref name=Coelho/>

Other arthropods may make use of the antlion larva's ability to trap prey. The larva of the Australian horsefly (''[[Scaptia muscula]]'') lives in antlion (for example ''[[Myrmeleon pictifrons]]'') pit traps and feeds on the prey caught,<ref name=Swanson/> and the female [[Chalcididae|chalcid wasp]] (''[[Lasiochalcidia igiliensis]]'') purposefully allows itself to be trapped so that it can parasitise the antlion larva by ovipositing between its head and thorax.<ref>{{cite book |author=Piek, Tom |title=Venoms of the Hymenoptera: Biochemical, Pharmacological and Behavioural Aspects |year=2013 |publisher=Elsevier |isbn=978-1-4832-6370-0 |page=74 |url=https://books.google.com/books?id=xBQlBQAAQBAJ&pg=PA74}}</ref>

Recent research has found that antlion larvae often "play dead" for a variable amount of time (from a few minutes up to an hour) when disturbed to hide from predators. The method is effective; it increased survival rates in patches that use it by 20%.<ref>{{cite news |last=Giaimo |first=Cara |date=2021-03-07 |title=The Power of Playing Dead |language=en-US |work=The New York Times |issn=0362-4331 |url=https://www.nytimes.com/2021/03/07/science/playing-dead-antlion-larvae.html |access-date=2021-03-08}}</ref> Furthermore, they appear to have maximized its usefulness—further increasing the duration is not likely to convey substantial survival benefits to the larvae.<ref>{{cite journal |last1=Franks |first1=Nigel R. |last2=Worley |first2=Alan |last3=Sendova-Franks |first3=Ana B. |title=Hide-and-seek strategies and post-contact immobility |journal=Biology Letters |year=2021 |volume=17 |issue=3 |article-number=20200892 |pmid=33653098 |doi=10.1098/rsbl.2020.0892 |pmc=8086978}}</ref>

==Evolution==
The closest living relatives of antlions within the [[Myrmeleontoidea]] are the [[owlflies]] (Ascalaphidae); the [[Nymphidae]] are more distantly related. The extinct [[Araripeneuridae]] and [[Babinskaiidae]] are considered likely to be stem groups in the Myrmeleontiformia [[clade]].<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 |date=2007 |publisher=Cambridge University Press |isbn=978-1-139-46776-6 |page=334 |url=https://books.google.com/books?id=5ZBh_-QvX2MC&pg=PA334}}</ref> The [[phylogeny]] of the Neuroptera has been explored using [[mitochondrial DNA]] sequences, and while issues remain for the group as a whole (the "[[Hemerobiiformia]]" being [[paraphyletic]]), the Myrmeleontiformia is generally agreed to be [[monophyletic]], giving the following [[cladogram]]:<ref name=Yan>{{cite journal |author1=Yan, Y. |author2=Wang Y |author3=Liu, X. |author4=Winterton, S.L. |author5=Yang, D. |title=The First Mitochondrial Genomes of Antlion (Neuroptera: Myrmeleontidae) and Split-footed Lacewing (Neuroptera: Nymphidae), with Phylogenetic Implications of Myrmeleontiformia |journal=Int J Biol Sci |date=2014 |volume=10 |issue=8 |pages=895–908 |pmid=25170303 |doi=10.7150/ijbs.9454 |pmc=4147223}}</ref>

{{clade
|label1=[[Neuroptera]]
|1={{clade
   |1=[[Osmylidae]] (formerly in "Hemerobiiformia") [[File:Oedosmylus sp crop.jpg|100px]]
   |2={{clade
      |label1=Most of former "Hemerobiiformia"
      |1={{clade
         |1=[[Mantispidae]] (mantidflies) [[File:Mantispidae fg1.jpg|75px<!--|''[[Ditaxis biseriata]]'' ([[Mantispidae]])-->]]
         |2={{clade
            |1=[[Ithonidae]] (moth lacewings) [[File:Rapisma sp- India crop.jpg|75px<!--Ithonidae-->]]
            |2=[[Chrysopidae]] (green lacewings) [[File:Chrysoperla carnea Guldoeje.jpg|100px<!--|''[[Chrysoperla carnea]]'' ([[Chrysopidae]])-->]]
            }}
         }}
      |label2=[[Myrmeleontiformia]]
      |2={{clade
         |1=[[Nymphidae]] (split-footed lacewings) [[File:Nymphes myrmeleonoides (3155078680) crop.jpg|75px]]
         |2={{clade
            |1='''Myrmeleontidae (antlions)''' [[File:Distoleon tetragrammicus01.jpg|75px<!--|''[[Distoleon tetragrammicus]]'', ([[Myrmeleontidae]])-->]]
            |2=[[Ascalaphidae]] (owlflies) [[File:Libelloides coccajus 210505.jpg|75px<!--|''[[Libelloides coccajus]]'' ([[Ascalaphidae]])-->]]
            }}
         }}
      }}
   }}
}}

The [[subfamilies]] are shown below; a few genera, mostly [[fossil]], are of uncertain or [[basal (evolution)|basal]] position. The fossil record of antlions is very small by neuropteran standards. However, some [[Mesozoic]] fossils attest to the antlions' origin more than 150 million years ago. These were at one time separated as the Palaeoleontidae, but are now usually recognized as early antlions.<ref name=Engel2007/>

==Taxonomy==
The supra-generic classification within the Myrmeleontidae is disputed by different authors. Several different schematics have been proposed.

===Stange (2004) classification===
Stange recognized three subfamilies, consisting of [[Stilbopteryginae]], [[Palparinae]], and [[Myrmeleontinae]].<ref name="Machado2018"/> [[Brachynemurinae]], [[Dendroleontinae]], and other subfamilies recognized by prior authors{{who|date=July 2024}} were placed in the [[Myrmeleontinae]].

* '''[[Stilbopteryginae]]''' <small>Newman, 1853</small>
# [[Stilbopterygini]] {{Au|Newman, 1853}} [then monotypic, later two genera]<br/>
* '''[[Palparinae]]''' <small>Banks, 1911</small>
# [[Dimarini]] <small>Navas, 1914</small>
# [[Palparidiini]] <small>Markl, 1954</small>
# [[Palparini]] <small>Banks, 1911</small>
# [[Pseudimarini]] <small>Markl 1954</small>
* '''[[Myrmeleontinae]]''' <small>Latreille, 1802</small>
# [[Acanthaclisini]] <small>Navas, 1912</small>
# [[Brachynemurini]] <small>Banks, 1927</small>
# [[Dendroleontini]] <small>Banks, 1899</small>
# [[Gnopholeontini]] <small>Stange, 1994</small>
# [[Lemolemini]] <small>Stange, 1994</small>
# [[Maulini]] <small>Markl 1954</small>
# [[Myrmecaelurini]] <small>Esben-Petersen, 1918</small>
# [[Myrmeleontini]] <small>Latreille, 1802</small>
# [[Nesoleontini]] <small>Markl, 1954</small>
# [[Nemoleontini]] <small>Banks, 1911</small>

===Michel ''et al.'' (2017) classification===
Michel ''et.al'' recognized just four subfamilies, consisting of [[Acanthaclisinae]], [[Myrmeleontinae]], [[Palparinae]], and [[Stilbopterygini|Stilbopteryginae]].<ref name="Michel2017">{{cite journal |last1=Michel |first1=Bruno |last2=Clamens |first2=Anne-Laure |last3=Béthoux |first3=Olivier |last4=Kergoat |first4=Gael J. |last5=Condamine |first5=Fabien L. |title=A first higher-level time-calibrated phylogeny of antlions (Neuroptera: Myrmeleontidae) |journal=Molecular Phylogenetics and Evolution |date=February 2017 |volume=107 |pages=103–116 |bibcode=2017MolPE.107..103M |pmid=27780793 |doi=10.1016/j.ympev.2016.10.014}}</ref>

* '''[[Stilbopterygini|Stilbopteryginae]]''' <small>Newman, 1853</small>
# [[Stilbopterygini]] <small>Newman, 1853</small>
* '''[[Acanthaclisinae]]''' <small>Navas, 1912</small>
# [[Acanthaclisini]] <small>Navas, 1912</small>
* '''[[Palparinae]]''' <small>Banks, 1911</small>
# [[Dimarini]] <small>Navas, 1914</small>
# [[Palparidiini]] <small>Markl, 1954</small>
# [[Palparini]] <small>Banks, 1911</small>
# [[Pseudimarini]] <small>Markl 1954</small>
* '''[[Myrmeleontinae]]''' <small>Latreille, 1802</small>
# [[Brachynemurini]] <small>Banks, 1927</small>
# [[Dendroleontini]] <small>Banks, 1899</small>
# [[Gepini]] <small>Markl, 1954</small>
# [[Gnopholeontini]] <small>Stange, 1994</small>
# [[Lemolemini]] <small>Stange, 1994</small>
# [[Maulini]] <small>Markl 1954</small>
# [[Myrmecaelurini]] <small>Esben-Petersen, 1918</small>
# [[Myrmeleontini]] <small>Latreille, 1802</small>
# [[Nesoleontini]] <small>Markl, 1954</small>

===Machado ''et al.'' (2018) classification===
A subsequent revision by Machado ''et.al'' recognized a different four subfamilies, notable in the inclusion of the family Ascalaphidae as a subfamily instead of as a sister taxon to Myrmeleontidae: [[owlfly|Ascalaphinae]], [[Myrmeleontinae]], [[Dendroleontinae]], and [[Nemoleontinae]].<ref name="Machado2018">{{cite journal |last1=Machado |first1=R. J. P. |last2=Gillung |first2=J. P. |last3=Winterton |first3=S. L. |last4=Garzon-Orduña |first4=I. J. |last5=Lemmon |first5=A. R. |last6=Lemmon |first6=E. M. |last7=Oswald |first7=J. D. |title=Owlflies are derived antlions: Anchored phylogenomics supports a new phylogeny and classification of Myrmeleontidae (Neuroptera) |journal=Systematic Entomology |volume=44 |year=2018 |issue=2 |pages=418–450 |doi=10.1111/syen.12334 |doi-access=free}}</ref>

* '''[[owlfly|Ascalaphinae]]''' Rambur, 1842
# [[Ascalaphini]] <small>Lefèbvre, 1842</small>
# [[Dimarini]] <small>Navas, 1914</small>
# [[Haplogleniini]] <small>Newman, 1853</small>
# [[Palparini]] <small>Banks, 1911</small>
# [[Stilbopterygini]] <small>Newman, 1853</small>
# [[Ululodini]] <small>Van der Weele 1909</small>
* '''[[Myrmeleontinae]]''' <small>Latreille, 1802</small>
# [[Acanthaclisini]] <small>Navas, 1912</small>
# [[Brachynemurini]] <small>Banks, 1927</small>
# [[Myrmecaelurini]] <small>Esben-Petersen, 1918</small>
# [[Myrmeleontini]] <small>Latreille, 1802</small>
# [[Nesoleontini]] <small>Markl, 1954</small>
* '''[[Dendroleontinae]]''' <small>Banks, 1899</small>
# [[Acanthoplectrini]] <small>Markl, 1954</small>
# [[Dendroleontini]] <small>Banks, 1899</small>
# [[Glenurini]] <small>Banks, 1927</small>
# [[Megistopini]] <small>Navas, 1912</small>
* '''[[Nemoleontinae]]''' <small>Banks, 1911</small>
# [[Nemoleontini]] <small>Banks, 1911</small>
# [[Protoplectrini]] <small>Tillyard, 1916</small>

===Jones (2019) classification===
Jones dissented with Machado ''et al.'' and presented an alternative classification, restoring the Ascalaphidae to its traditional family-rank placement, and elevating Palparinae and Stilbopteryginae to family level, leaving only a single subfamily in Myrmeleontidae ''sensu stricto''.<ref name="Jones 2019">{{cite journal |last=Jones |first=Joshua R. |title=Total-evidence phylogeny of the owlflies (Neuroptera, Ascalaphidae) supports a new higher-level classification |journal=Zoologica Scripta |volume=48 |issue=6 |date=6 October 2019 |pages=761–782 |doi=10.1111/zsc.12382}}</ref> However, this classification has been rejected by Hévin ''et al.'' (2023), finding the taxonomic decisions to be "not substantiated, with only the family name Palparidae being mentioned in isolation."<ref name=v2023>{{cite journal |last1=Hévin |first1=Noémie M.-C |last2=Kergoat |first2=Gael. J. |last3=Clamens |first3=Anne-Laure |last4=Bruno |first4=Le Ru |last5=Mervyn W. |first5=Mansell |last6=Michel |first6=Bruno |date=2023 |title=Evolution, systematics and historical biogeography of Palparini and Palparidiini antlions (Neuroptera: Myrmeleontidae): Old origin and in situ diversification in Southern Africa |journal=Systematic Entomology |volume=48 |issue=4 |pages=600–617 |bibcode=2023SysEn..48..600H |doi=10.1111/syen.12593 |doi-access=free |hdl=2263/93981 |hdl-access=free}}</ref>

* '''[[Myrmeleontinae]]''' <small>Latreille, 1802</small>
# [[Acanthaclisini]] <small>Navas, 1912</small>
# [[Brachynemurini]] <small>Banks, 1927</small>
# [[Dendroleontini]] <small>Banks, 1899</small>
# [[Myrmecaelurini]] <small>Esben-Petersen, 1918</small>
# [[Myrmeleontini]] <small>Latreille, 1802</small>
# [[Nemoleontini]] <small>Banks, 1911</small>
# [[Nesoleontini]] <small>Markl, 1954</small>

<gallery mode="nolines">
File:Antlion doodles.jpg|Antlion larva trails (doodles) in sand
File:2015.10-434-146bp antlion(unidentified sp.,adult,L(wing tip-antnena tip)4.8cm) DD11.176086,-4.335053@Bobo-Dioulasso,BF thu29oct2015-1054h.jpg|Thorax and head (with club-shaped antenna) of antlion adult
File:Antlion1 by Jonathan Numer.jpg|Larva
File:Formicaleone.jpg|20x closeup of larva
File:Antlion.ogv|Video of antlion larva trying to catch prey with sand traps and eating a small [[spider]]
File:Antlion video.ogv|Video of a larva trapping an ant by throwing sand at it
</gallery>

[[File:Myrmecoleon 1491.jpg|thumb|upright|[[Myrmecoleon]], from the ''[[Hortus Sanitatis]]'' of Jacob Meydenbach, 1491]]

==In culture and folklore==
In popular folklore in the southern United States, people recite a poem or chant to make the antlion come out of its hole.<ref>{{cite book |last=Howell |first=Jim |title=Hey, Bug Doctor!: The Scoop on Insects in Georgia's Homes and Gardens |publisher=University of Georgia Press |year=2006 |pages=15–16 |isbn=0-8203-2804-9 |url=https://books.google.com/books?id=9h1JAVPOmM8C&q=antlion+folklore&pg=PA15}}</ref> Similar practices have been recorded from Africa, the Caribbean, China and Australia.<ref>{{cite book |last=Ouchley |first=Kelby |title=Bayou-Diversity: Nature and People in the Louisiana Bayou Country |publisher=LSU Press |year=2011 |page=85 |isbn=978-0-8071-3861-8 |url=https://books.google.com/books?id=J1N12aH0810C&q=antlion+folklore&pg=PA85}}</ref>

The [[Myrmecoleon]] was a mythical ant–lion hybrid written about in the 2nd century AD ''[[Physiologus]]'', where animal descriptions were paired with Christian morals. The ant-lion as described was said to starve to death because of its dual nature – the lion nature of the father could [[obligate carnivore|only eat meat]], but the ant half from the mother could only [[herbivore|eat grain chaff]], thus the offspring could not eat either and would starve.<ref name="Dekkers2000">{{cite book |last=Dekkers |first=Midas |title=Dearest Pet: On Bestiality |date=2000 |publisher=Verso |isbn=978-1-85984-310-9 |page=78 |url=https://books.google.com/books?id=wF772tvr6YEC&pg=PA78}}</ref> It was paired with the Biblical verse [[Matthew 5:37]].<ref name="Grant1999">{{cite book |last=Grant |first=Robert M. |title=Early Christians and Animals |date=1999 |publisher=Psychology Press |isbn=978-0-415-20204-6 |pages=61– |url=https://books.google.com/books?id=FKr4N2nECZEC&pg=PA61}}</ref> The fictional ant-lion of ''Physiologus'' is probably derived from a misreading of Job 4:11.{{efn|The verse runs "The old lion perisheth for lack of prey, and the stout lion's whelps are scattered abroad." (Authorised Version)}}<ref name="Dekkers2000"/>

The French naturalist [[Jean-Henri Fabre]] wrote that "The Ant-lion makes a slanting funnel in the sand. Its victim, the Ant, slides down the slant and is then stoned, from the bottom of the funnel, by the hunter, who turns his neck into a catapult."<ref name="Fabre2013">{{cite book |last=Fabre |first=Jean-Henri |title=Fabre's Book of Insects |date=2013 |publisher=Courier Corporation |isbn=978-0-486-32011-3 |page=108 |url=https://books.google.com/books?id=g3LCAgAAQBAJ&pg=PA108}}</ref><ref name="Fabre2002">{{cite book |last=Fabre |first=Jean-Henri |title=The Wonder Book of Science |date=2002 |publisher=The Minerva Group |isbn=978-0-89875-711-8 |pages=45– |url=https://books.google.com/books?id=oB7LJRlnpbMC&pg=PA45}}</ref>

In [[Japan]], both the insect and its pit-traps are popularly known as {{nihongo|''Arijigoku''|蟻地獄||lit. "Ant Hell"}}. This term has since become a stock colloquialism for any "inescapable" trap, whether literal or metaphorical (''e.g.'' an unpleasant social obligation).<ref>{{Cite web |url=https://reichanjapan.wordpress.com/2016/02/29/arijigoku-nintendos-puzzling-preoccupation-with-ant-hell/ |title=Arijigoku: Nintendo's Puzzling Preoccupation with Ant Hell |author=<!--Not stated--> |date=February 29, 2016 |website=Reichan Japan |publisher=Wordpress |quote=}}</ref>

Antlions appear as antagonists in the 1991 [[life simulation game|life simulation]] [[video game]], ''[[SimAnt]]'', and (in a giant form) in the ''[[Final Fantasy]]'' series, ''[[Grounded (video game)|Grounded]]'', ''[[Terraria]]'', ''[[Don't Starve Together]]'', ''[[Monster Rancher 2]]'', ''[[Mother 3]]'' and in the ''[[Half-Life (series)|Half-Life 2]]'' video game series as an unrelated alien insect species sharing sand burrowing traits with the real antlion larvae. The [[list of generation III Pokémon#Trapinch|Trapinch, Vibrava, and Flygon]] [[Pokémon]] evolution line is based on an antlion. The fictional [[sarlacc]] from the ''[[Star Wars]]'' franchise is often compared to the real-life antlion.<ref>Jeanne Cavelos, ''The Science of Star Wars: An Astrophysicist's Independent Examination of Space Travel, Aliens, Planets, and Robots as Portrayed in the ''Star Wars'' Films and Books''(New York: St. Martin's Press, 1999), p. 71, {{ISBN|0-312-20958-4}}.</ref><ref name="starwars">{{cite web |last1=Moore |first1=Chris |title=You Don't Have To Go To A Galaxy Far, Far Away To See These 5 Real Life Star Wars Aliens! |website=Moviepilot.com |url=https://moviepilot.com/posts/3680732 |access-date=4 February 2017 |archive-url=https://web.archive.org/web/20170204171316/https://moviepilot.com/posts/3680732 |archive-date=4 February 2017}}</ref> It also appears as a predator in the film ''[[Enemy Mine (film)|Enemy Mine]]''.

In the third book of [[Tove Jansson]]'s [[The Moomins|Moomins]] series, ''[[Finn Family Moomintroll]]'', a rather large and fanciful antlion appears in the second chapter, depicted as a sand-dwelling predator with the literal head of a [[lion]].<ref name="Jansson2010">{{cite book |last=Jansson |first=Tove |title=Finn Family Moomintroll |date=1950 |publisher=Penguin Books |isbn=978-0-14-030150-2 |page=33 |url=https://books.google.com/books?id=BFNABAAAQBAJ}}</ref>

==See also==
* [[List of Myrmeleontidae genera]]
* [[Vermileonidae]] - wormlions

==Notes==
{{Notelist}}

==References==
{{Reflist}}

==External links==
{{Wiktionary|antlion}}
{{Commons-inline}}

{{Neuroptera|3}}
{{Taxonbar|from=Q231439}}

[[Category:Myrmeleontidae| ]]
[[Category:Neuroptera]]
[[Category:Articles containing video clips]]