Antlion
1108210
223646160
2008-07-05T01:47:05Z
Dysmorodrepanis
303688
/* EtymologySwanson (2007) */
{{otheruses4|the insect|the alien creature in the computer game Half-Life 2|Antlion (Half-Life 2)}}
{{Taxobox
| name = Antlions
|fossil_range=[[Mesozoic]] - Recent
| image = Distoleon tetragrammicus01.jpg
| image_width = 240px
| image_caption = Adult ''[[Distoleon tetragrammicus]]''
| regnum = [[Animal]]ia
| phylum = [[Arthropod]]a
| subphylum = [[Hexapoda]]
| classis = [[Insect]]a
| subclassis = [[Pterygota]]
| infraclassis = [[Neoptera]]
| superordo = [[Endopterygota]] or [[Neuropterida]]
| ordo = [[Neuroptera]]
| subordo = [[Myrmeleontiformia]]
| superfamilia = [[Myrmeleontoidea]]
| familia = '''Myrmeleontidae'''
| familia_authority =
| subdivision_ranks = [[Subfamilies]]
| subdivision =
[[Acanthaclisinae]]<br />
[[Brachynemurinae]]<br />
[[Dendroleontinae]]<br />
[[Dimarinae]]<br />
[[Echthromyrmicinae]]<br />
[[Glenurinae]]<br />
[[Myrmecaelurinae]]<br />
[[Myrmeleontinae]]<br />
[[Nemoleontinae]]<br />
[[Palparinae]]<!-- includes Araripeneurinae --><br />
[[Pseudimarinae]]<br />
[[Stilbopteryginae]]<br />
| synonyms =
Myrmeleonidae <small>(''[[lapsus]]'')</small><br />
Palaeoleontidae<br />
and see [[#Systematics|text]]
}}
'''Antlions''' are a [[family (biology)|family]] of [[insect]]s in the order [[Neuroptera]] with the [[scientific name]] '''Myrmeleontidae''' (sometimes misspelled as '''"Myrmeleonidae"'''); the most well-known genus is ''[[Myrmeleo]]''. There are about 2,000 [[species]]. Strictly speaking, the term "antlion" applies to the larval form of the members of this family, but while several languages have own terms for the adult, there is no widely used word for them in [[English (language)|English]]. Very rarely, the adults are called '''"antlion lacewings"'''.
The antlion larva is often called '''"[[doodlebug]]"''' in [[North America]] because of the odd winding, spiralling trails it leaves in the sand while looking for a good location to build its trap, as these trails look like someone has [[doodle]]d in the sand. It is also sometimes called '''"sand dragon"'''.
==Description and ecology==
[[Image:Euroleon fg06.jpg|thumb|left|''[[Euroleon nostras]]'' imago]]
[[Image:Euroleon fg02.jpg|thumb|left|''Euroleon nostras'' larva]]
[[Image:Euroleon fg02b.jpg|thumb|left|''Euroleon nostras'' cocoon]]
[[Image:Antlion2.jpg|thumb|left|Pupa in cocoon (unknown species)]]
[[Image:Euroleon fg03.jpg|thumb|right|''Euroleon nostras'', cocoon with pupal integument]]
Antlions are worldwide in distribution, most common in arid and sandy habitats. A few species occur in cold-[[temperate]] places; a famous example is the European ''[[Euroleon nostras]]'', whose [[scientific name]] means "our European [ant]lion". They can be fairly small to very large Neuroptera (wingspan range of 2-15 cm). The antlion [[larva]]e eat small [[arthropod]]s - mainly [[ant]]s -, while the adults of some species eat small [[pollen]] and [[nectar]], while others are predators of small arthropods in the adult stage too<ref>Engel & Grimaldi (2007)</ref>. In certain species of Myrmeleontidae, such as ''[[Dendroleon pantheormis]]'', the larva, although resembling that of ''Myrmeleon'' structurally, makes no pitfall, but seizes passing prey from any nook or crevice in which it shelters.
The adult has two pairs of long, narrow, multi-veined [[insect wing|wings]] in which the [[apex|apical]] veins enclose regular oblong spaces, and a long, slender [[abdomen]]. Although they greatly resemble [[dragonfly|dragonflies]] or [[damselfly|damselflies]], they belong to an entirely different [[infraclass]] among the [[pterygota|winged insects]]. Antlions are easily distinguished from damselflies by their prominent, apically clubbed [[Antenna (biology)|antennae]] which are about as long as head and [[thorax]] combined. Also, the pattern of wing venation differs, with the very long hypostigmatic cell (behind the fusion point of Sc and R1) being several times as long as wide. They also are very feeble fliers and are normally found fluttering about in the night, in search of a mate. The adult is thus rarely seen in the wild because it is typically active only in the evening.
The life cycle of the antlion begins with oviposition ([[egg (biology)|egg]]-laying). The female antlion repeatedly taps the sand surface with the tip of her [[abdomen]]. She then inserts her abdomen into the sand and lays an egg. The antlion larva is a ferocious-appearing creature with a robust, fusiform body, a very plump [[abdomen]], the [[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 sicklelike jaws with several sharp, hollow projections. The jaws are formed by the [[insect mouthparts|maxillae and mandibles]], which in each pincer enclose a canal for injecting venom between them. Depending on species and where it lives, the larvae will either hide under leaves or pieces of wood, in cracks of rocks, or dig pits in sandy areas.
The [[pupa]]l stage of the antlion is [[quiescent]]. The larva makes a globular [[Cocoon (silk)|cocoon]] of sand stuck together with fine [[silk]] spun from a slender spinneret at the posterior end of the body. These cocoons may be buried several centimeters deep in the sand. It remains there for one month, until the completion of the transformation into the sexually mature insect, which then emerges from the case, leaving the pupal integument behind, and climbs to the surface. After about 20 minutes the adult's wings are fully opened and it will fly off in search of a mate. The adult is considerably larger than the larva; they exhibit the greatest disparity in size between larva and adult of any type of [[holometabolous]] insects, by virtue of the adults having an extremely thin, flimsy [[exoskeleton]] - in other words, they have extremely low mass per unit of volume.
==Systematics==
The closest living relatives of antlions are the [[owlflies]] (Ascalaphidae). The extinct [[Babinskaiidae]], known only from [[fossil]]s, were also very closely related. These three form the most strongly [[apomorph|derived]] lineages of the [[superfamily]] [[Myrmeleontoidea]].<ref name = haaramo2008>See references in Haaramo (2008)</ref>
The numerous [[genera]] and [[species]] of antlions are for the largest part assigned to a diversity of [[subfamilies]]. A few genera, mostly [[fossil]], are of more uncertain or [[basal (evolution)|basal]] position. The fossil record of antlions is very small by neuropteran standards. However, there are some [[Mesozoic]] fossils which attest to the antlions' origin more than 150 million years ago. These were at one time separated as [[family (biology)|family]] '''Palaeoleontidae''', but are now usually recognized as the most primitive examples of actual antlions.<ref>Engel & Grimaldi (2007)</ref>
Antlions of uncertain systematic position are:<ref name = haaramo2008 />
<!-- * ''[[Dendroleon]]'' (fossil) -> Dendroleontidae? -->
* ''[[Palaeoleon]]'' (fossil)
* ''[[Porrerus]]''<!-- "Porrerus" is a lapsus -->
* ''[[Samsonileon]]'' (fossil)
==Etymology<ref>Swanson (2007)</ref>==
The exact meaning of the name "antlion" is uncertain. It has been thought that it refers to the fact that ants form a large percentage of the prey of the insect, the suffix "lion" merely suggesting destroyer or eater. Perhaps, however, the name as a whole only signifies a large terrestrial biting apterous insect, surpassing ants in size and predatory habits, similar to the concept of the lion being "king of the animals" in European mythology and lore. In any case, the term seems to go back to [[classical antiquity|Antiquity]].
In most European languages, at least the larvae are known under the local term corresponding to "antlion":
* [[Romance languages]]: e.g. ''formicaleo'' ([[Medieval Latin]]), ''formicaleone'' (Italian), ''formiga-leão'' (Portuguese), ''formiga león'' (Galician), ''formigues lleó'' (Catalan), ''fourmilion'' or ''fourmis-lion'' (French), ''hormiga león'' (Spanish), ''leii furnicilor'' (Romanian)
* [[Germanic languages]]: e.g. ''Ameisenlöwe'' (German), ''maurløve'' (Norwegian), ''mierenleeuw'' (Dutch), ''mierleeu'' (Afrikaans), ''myreløve'' (Danish), ''myrlejon'' (Swedish)
* [[Slavic languages]]: e.g. ''mravkolev'' (Czech), ''mravkol'v'' (мравкольв, Bulgarian), ''mravlji lav'' (Serbo-Croatian), ''mrówkolew'' (Polish), ''murav'iniy lev'' (муравьиный лев, Russian)
* [[Baltic languages]]: e.g. ''skudrulauva'' (Latvian), ''skruzdžių liūtas''{{Verify source|date=May 2008}}<!-- not sure about -ių and -tas --> (Lithuanian)
* [[Baltic-Finnic languages]]: e.g. ''muurahaisleijona'' (Finnish), ''sipelgalõvi'' (Estonian)
* [[Albanian (language)|Albanian]]: ''luani thnegslave''
In [[Hebrew]], antlions are called ''arinamal'' (ארינמל), in [[Arabic (language)|Arabic]] ''asad al-naml'' (أسد النمل{{Verify source|date=May 2008}}), and in [[Turkish (language)|Turkish]] their name is ''karınca arslanı''. These are also literal equivalents of the English name, though at least the last would seem to date to more recent times, being adapted from some European or Near Eastern language.
The [[scientific name]] of the [[type genus]] ''Myrmeleo'' - and thus, the family as a whole - is derived from [[Ancient Greek]] ''léon'' (λέων, [[Modern Greek]]: ''lión'', λιόν) "lion" + ''mýrmex'' (μύρμηξ, Modern Greek: ''myrmígia'', μυρμήγκια) "ant", in a literal translation of the names common across Europe.
But in [[Hungarian (language)|Hungarian]] - whose progenitor separated from the ancestral Baltic-Finnic languages in the [[Urals]] region the eastern border of Europe - antlions are called ''hangyaleső'', "ant-ambushers". And terms meaning "antlion" are not found as the animals' original names far outside Europe. So it seems that the concept of and words meaning "antlion" were known across Europe by about [[500 AD]], when the [[Magyars]] began their migration to present-day Hungary - and that they at that time already had embraced the concept of and term for "ant-ambusher" so firmly that modern Hungarian is apparently the only language in the whole of Europe and the Mediterranean region whose name for Myrmeleontidae has an [[etymology]] different from all others'. It stands to note that one of the first attested occurrences of ''μυρμηκολέων'' is a mistranslation of a rarely-used Hebrew term for "lion" in the [[Septuagint]]<ref>{{bibleverse||Job|4:11|9}}, translated more correctly as "old lion" in the [[King James Version]].</ref>, dating back a few centuries BC. Therefore the "antlion" concept would seem to have spread across Europe from the Mediterranean region during the first few centuries AD, about the time of the decline of the [[Roman Empire]], when cultural exchange between Mediterranean and Northern European peoples was vigorous.
[[Image:Antlion1 by Jonathan Numer.jpg|thumb|left|Closeup of an antlion larva]]
Outside Europe, antlions are sometimes given names that compare their jaws to a [[bull]]'s horns. In [[Mandarin (language)|Mandarin]], ''dǎotuìniú'' (倒退牛, "backward-moving bull"), ''dìgǔniú'' (地古牛, "ancient soil-bull") and ''yǐniú'' (蚁牛, "ant-bull") are known. In southwestern North America, the colloquial Spanish name is ''torito'' ("little bull"). Other names refer to the larvae's behavior: ''aMbututu'' ("hider-and-digger", [[Xitsonga]]), ''undur-undur'' ("backwards-walker", [[Malay (language)|Malay]]), ''kumhar'' ("potter", [[Punjabi (language)|Punjabi]]) or ''bingundha'' ("tunnel digger", [[Sinhala (language)|Sinhala]]).
In some regions of [[East Asia]], antlions - or more specifically, their traps - are locally known by terms translating approximately as "ant-hell"<ref>The [[Diyu|East Asian "hell"]] is imagined as a prison for souls, unlike the [[Hell|Christian place of damnation]].</ref>. In Korean, this is ''gaemi jiog'' (개미지옥), and the corresponding Japanese word is ''ari-jigoku'' (アリジゴク)<ref>Spahn & Hadamitzky (2003): p.1416</ref>. The term ''ari-jigoku'' is also used to describe a situation in which one has fallen into inevitable doom, and must languish while awaiting disaster.
[[Image:Adult antlion.JPG|thumb|right|The [[damselfly]]-like adult]]
In some languages, there are different names for the larva and the adult. In German, the adults are called ''Ameisenjungfer'' (approximately "ant-[[damselfly]]"<ref>Literally, ''Jungfer'' means "[[maiden]]", but as part of insect names it means "damselfly".</ref>); the Finnish term ''muurahaiskorento'' and the Estonian ''sipelgakiillaste''{{Verify source|date=May 2008}}<!-- possibly plural --> mean the same. In Dutch ''mierenleeuwjuffer'' ("antlion-damselfly") is sometimes used. In Japanese, the term for the adult animals is ''usuba-kagerou'' (ウスバカゲロウ), which means "thin-winged [[mayfly]]". In Korea, the adult is known as ''Yeongju jamjari'' (명주잠자리), meaning approximately "Spirit Mountain damselfly". All these terms refer to the [[habitus]] and the somewhat clumsy flight of the adult animals, which somewhat resembles the unrelated insects they are compared to.
There are also a number of nicknames for the larvae, most of which have no readily discernible etymology. Apart from the [[North America]]n "doodlebug" mentioned above, these include "shuntie" and ''joerie'' (pronounced "yoory") in [[South Africa]], "devil-devil" in [[Australia]], ''jampeepee'' on [[Antigua and Barbuda]], "tortee" or "torty" (probably from "[[tortoise]]") on [[St Vincent and the Grenadines]], and ''chanchito'' in [[Costa Rica]].
==Sand pit traps==
[[Image:Antlion trap.jpg|thumb|left|Sand pit trap of an antlion]]
An average-sized larva digs a pit about 2 inches deep and 3 inches wide at the edge. This behavior has also [[evolve]]d in a family of [[fly|flies]], the [[Vermileonidae]], whose larvae dig the same sort of pit to feed on ants. Having marked out the chosen site by a circular groove, the antlion larva starts to crawl backwards, using its abdomen as a plough to shovel up the soil. By the aid of one front leg it places consecutive heaps of loosened particles upon its head, then with a smart jerk throws each little pile clear of the scene of operations. Proceeding thus it gradually works its way from the circumference towards the center. As it slowly moves round and round, the pit gradually gets deeper and deeper, until the slope angle reaches the [[angle of repose]] (that is, the steepest angle the sand can maintain, where it is on the verge of collapse from slight disturbance). 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.
[[Image:AmeisenlöwenTrichter.jpg|thumb|right|Sand pit trap with remains of an ant]]
Since the sides of the pit consist of loose sand at its angle of repose<ref>Botz ''et al.'' (2003)</ref>, they afford an insecure foothold to any small insects that inadvertently venture over the edge, such as [[ant]]s. Slipping to the bottom, the prey is immediately seized by the lurking antlion; or if it attempts to scramble again 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. 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.
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 will suck 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.
Antlions are especially abundant in soft sand beneath trees or under overhanging [[rock (geology)|rocks]]. Apparently the larvae prefer dry places that are protected from the rain. Eventually the larva attains its maximum size and undergoes [[Metamorphosis (biology)|metamorphosis]]. The entire length of time from egg-laying to adulthood may take two or three years due to the uncertainty and irregular nature of its food supply. When it first hatches, the tiny larva specializes in very small insects, but as it grows larger, it constructs larger pits and thus catches larger prey.
==See also==
* [[:de:Ameisenlöwe]] Featured article on antlion larvae on German Wikipedia.
==Footnotes==
{{Reflist}}
==References==
{{commons|Myrmeleontidae}}
:{{1911}}
* {{aut|Botz, Jason T.; Loudon, Catherine; Barger, J. Bradley; Olafsen, Jeffrey S. & Steeples, Don W.}} (2003): Effects of slope and particle size on ant locomotion: Implications for choice of substrate by antlions. ''Journal of the Kansas Entomological Society'' '''76'''(3): 426-435 [http://cat.inist.fr/?aModele=afficheN&cpsidt=14973602 abstract]
* {{aut|Engel, Michael S. & Grimaldi, David A.}} (2007): The neuropterid fauna of Dominican and Mexican amber (Neuropterida, Megaloptera, Neuroptera). ''American Museum Novitates'' '''3587''': 1-58. [http://digitallibrary.amnh.org/dspace/bitstream/2246/5880/1/N3587.pdf PDF fulltext]
* {{aut|Haaramo, Mikko}} (2008): [[Mikko's Phylogeny Archive]]: [http://www.helsinki.fi/~mhaaramo/metazoa/protostoma/arthropoda/insecta/neuropterida/neuroptera.html Neuroptera]. Version of 2008-MAR-11. Retrieved 2008-APR-27.
* {{aut|Spahn, Mark & Hadamitzky, Wolfgang}} (2003): ''The Kanji Dictionary''. Tuttle. <small>ISBN 0-8048-2058-9</small>
* {{aut|Swanson, Mark}} (2007): The Antlion Pit - [http://www.antlionpit.com/language.html "Antlion" in the World's Languages]. Retrieved 2008-MAY-04.
[[Category:Neuroptera]]
[[ar:ليث عفرين]]
[[su:Undur-undur]]
[[cs:mravkolev]]
[[es:Hormiga león]]
[[de:Ameisenjungfern]]
[[fa:شیرمور]]
[[fr:Fourmilion]]
[[id:Antlion]]
[[it:Formicaleone]]
[[ja:ウスバカゲロウ]]
[[ko:명주잠자리]]
[[lv:Skudrulauva]]
[[hu:Hangyalesők]]
[[nl:Mierenleeuw]]
[[no:Maurløver]]
[[pl:Mrówkolew]]
[[pt:Formiga-leão]]
[[ru:Муравьиные львы]]
[[fi:Muurahaiskorennot]]
[[sv:Myrlejonsländor]]