Ant-lion
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The antlions are a group of about 2,000
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
in the
neuroptera The insect order (biology), order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera is grouped together with the Megaloptera (alderflies, f ...
n
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Myrmeleontidae. They are known for the
predatory Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
habits of their
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e, which mostly dig pits to trap passing
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
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
dragonflies A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of dragonflies are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threate ...
or
damselflies Damselflies are flying insects of the suborder Zygoptera in the order Odonata. They are similar to dragonflies (which constitute the other odonatan suborder, Epiprocta) but are usually smaller and have slimmer bodies. Most species fold the win ...
. Antlions have a worldwide distribution. The greatest diversity occurs in the tropics, but a few species are found in cold-
temperate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
locations, one such being the European ''
Euroleon nostras ''Euroleon nostras'' is a species of antlion found over most of Europe. The scientific name can be translated as "our European ntlion". Adults resemble dragonflies or damselflies and may reach up to long, with a wingspan of . The larvae prey ...
''. 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 Plant litter (also leaf litter, tree litter, soil litter, litterfall, or duff) is dead plant material (such as leaves, bark, needles, twigs, and cladodes) that has fallen to the ground. This detritus or dead organic material and its constituen ...
. Antlions are poorly represented in the
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
.
Myrmeleontiformia Myrmeleontiformia is an insect clade in the order Neuroptera, and which was historically treated as a suborder. The phylogeny of the Neuroptera has been explored using mitochondrial DNA sequences, and while issues remain for the order as a whole, ...
is generally accepted to be a
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
group, and within the
Myrmeleontoidea Myrmeleontoidea is a neuropteran superfamily in the clade Myrmeleontiformia. The following families are included: Superfamily Myrmeleontoidea (syn Nemopteroidea) * Family Ascalaphidae: owlflies (included in Myrmeleontidae in some classifications ...
, the antlions' closest living relatives are thought to be the
owlflies Ascalaphidae is a family of insects in the order Neuroptera, commonly called owlflies; there are some 450 extant species. They are fast-flying crepuscular or Diurnality, diurnal predators of other flying insects, and have large bulging eyes and ...
(Ascalaphidae). A 2019 study finds Myrmeleontidae to be monophyletic, aside from Stilbopteryginae and
Palparinae Palparinae is an antlion subfamily in the family Myrmeleontidae The antlions are a group of about 2,000 species of insect in the neuropteran family (biology), family Myrmeleontidae. They are known for the predation, predatory habits of their ...
, which form separate clades closer to Ascalaphidae. The predatory actions of the larvae have attracted attention throughout history and antlions have been mentioned in literature since classical times.


Etymology

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". In any case, the term seems to go back to
classical antiquity Classical antiquity, also known as the classical era, classical period, classical age, or simply antiquity, is the period of cultural History of Europe, European history between the 8th century BC and the 5th century AD comprising the inter ...
. The antlion larva is often called a "doodlebug" in
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
because of the odd winding, spiralling trails it leaves in the sand while relocating, which look as if someone has been
doodling A doodle is a drawing made while a person's attention is otherwise occupied. Doodles are simple drawings that can have concrete representational meaning or may just be composed of random and abstract lines or shapes, generally without ever l ...
. The
scientific name In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
of the
type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...
''Myrmeleo'' – and thus, the family as a whole – is derived from
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
''mýrmex'' (μύρμηξ) "ant" + ''léon'' (λέων) "lion", in a
loan translation In linguistics, a calque () or loan translation is a word or phrase borrowed from another language by literal word-for-word or root-for-root translation. When used as a verb, "to calque" means to borrow a word or phrase from another language whil ...
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".


Description

Antlions can be fairly small to very large
neuroptera The insect order (biology), order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera is grouped together with the Megaloptera (alderflies, f ...
ns, with wingspans ranging from . The African genus '' Palpares'' contains some of the largest examples. '' Acanthaclisis occitanica'' is the largest European species, with an wingspan, and most North American species approach this size. The adult has two pairs of long, narrow, multiveined, translucent
wings A wing is a type of fin that produces both lift and drag while moving through air. Wings are defined by two shape characteristics, an airfoil section and a planform. Wing efficiency is expressed as lift-to-drag ratio, which compares the bene ...
and a long, slender
abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
. Although they somewhat resemble
dragonflies A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of dragonflies are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threate ...
or
damselflies Damselflies are flying insects of the suborder Zygoptera in the order Odonata. They are similar to dragonflies (which constitute the other odonatan suborder, Epiprocta) but are usually smaller and have slimmer bodies. Most species fold the win ...
, they belong to a different
infraclass In biology, taxonomic rank (which some authors prefer to call nomenclatural rank because ranking is part of nomenclature rather than taxonomy proper, according to some definitions of these terms) is the relative or absolute level of a group of or ...
of
winged insects Pterygota ( ) is a subclass of insects that includes all winged insects and groups who lost them secondarily. Pterygota group comprises 99.9% of all insects. The orders not included are the Archaeognatha (jumping bristletails) and the Zygent ...
. Antlion adults are easily distinguished from damselflies by their prominent, apically clubbed antennae which are about as long as the head and
thorax The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main di ...
combined. 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
nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...
, and rarely seen by day. 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
oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typica ...
sites, and usually bears tufts of bristles for digging and a finger-like extension. The antlion
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
has a robust
fusiform Fusiform (from Latin ''fusus'' ‘spindle’) means having a spindle (textiles), spindle-like shape that is wide in the middle and tapers at both ends. It is similar to the lemon (geometry), lemon-shape, but often implies a focal broadening of a ...
body, a very plump abdomen, and a thorax bearing three pairs of walking legs. The
prothorax The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum (dorsal), the prosternum ( ventral), and the propleuron (lateral) on ...
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 maxillae and mandibles; the mandibles each contain a deep groove over which the maxilla fits neatly, forming an enclosed canal for injecting
venom Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a sti ...
to immobilise the victim, and
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
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. Antlion larvae are unusual among insects in lacking an
anus In mammals, invertebrates and most fish, the anus (: anuses or ani; from Latin, 'ring' or 'circle') is the external body orifice at the ''exit'' end of the digestive tract (bowel), i.e. the opposite end from the mouth. Its function is to facil ...
. All the
metabolic Metabolism (, from ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cellular processes; the ...
waste generated during the larval stage is stored; some is used to spin the silk for the cocoon and the rest is eventually voided as
meconium Meconium is the earliest stool of a mammalian infant resulting from defecation. Unlike later feces, meconium is composed of materials ingested during the time the infant spends in the uterus: intestinal epithelial cells, lanugo, mucus, am ...
at the end of its
pupa A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...
l stage.


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, scrub-clad
dune A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, flat ...
s, hedge bases, river banks, road verges, under raised buildings and in vacant lots.


Life-cycle

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 An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the ...
-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 Substrate may refer to: Physical layers *Substrate (biology), the natural environment in which an organism lives, or the surface or medium on which an organism grows or is attached ** Substrate (aquatic environment), the earthy material that exi ...
and lays an egg. 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. As
ambush predator Ambush predators or sit-and-wait predators are carnivorous animals that capture their prey via stealth, luring or by (typically instinctive) strategies utilizing an element of surprise. Unlike pursuit predators, who chase to capture prey u ...
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. They can take several years to complete their life-cycle; they mature faster with plentiful food, but can survive for many months without feeding. In cooler climates they dig their way deeper and remain inactive during the winter. When the larva attains its maximum size, it
pupa A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...
tes and undergoes
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth transformation or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and different ...
. It makes a globular cocoon of sand or other local substrate stuck together with fine
silk Silk is a natural fiber, natural protein fiber, some forms of which can be weaving, woven into textiles. The protein fiber of silk is composed mainly of fibroin and is most commonly produced by certain insect larvae to form cocoon (silk), c ...
spun from a slender
spinneret A spinneret is a silk-spinning organ of a spider or the larva of an insect. Some adult insects also have spinnerets, such as those borne on the forelegs of Embioptera. Spinnerets are usually on the underside of a spider's opisthosoma, and ar ...
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
holometabolous Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the clade Holometabola. Immatur ...
insect. This is by virtue of the fact that the
exoskeleton An exoskeleton () . is a skeleton that is on the exterior of an animal in the form of hardened integument, which both supports the body's shape and protects the internal organs, in contrast to an internal endoskeleton (e.g. human skeleton, that ...
of the adult is extremely thin and flimsy, with an exceptionally low
density Density (volumetric mass density or specific mass) is the ratio of a substance's mass to its volume. The symbol most often used for density is ''ρ'' (the lower case Greek letter rho), although the Latin letter ''D'' (or ''d'') can also be u ...
. The adult typically lives for about 25 days, but some insects survive for as long as 45 days.


Ecology

Antlion
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e eat small arthropods – mainly
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
s – while the adults of some species eat
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
and
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
, and others are predators of small arthropods. In certain species of Myrmeleontidae, such as ''
Dendroleon pantherinus ''Dendroleon pantherinus'' is a species of neuropteran insects of the antlion (Myrmeleontidae) family. Larvae are unique among antlions, as they do not create sandy pits, but rather develop in hidden shelters such as hollow trees. Taxonomy T ...
'', the larva, although resembling that of '' Myrmeleon'' structurally, makes no pitfall trap, but hides in
detritus In biology, detritus ( or ) is organic matter made up of the decomposition, decomposing remains of organisms and plants, and also of feces. Detritus usually hosts communities of microorganisms that colonize and decomposition, decompose (Reminera ...
in a hole in a tree and seizes passing prey. 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 A lichen ( , ) is a hybrid colony (biology), colony of algae or cyanobacteria living symbiotically among hypha, filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualism (biology), m ...
, and have been recorded at densities of up to 344 per square metre. The larva is a voracious predator. Within a few minutes of seizing its prey with its jaws and injecting it with
venom Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a sti ...
and
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
s, it begins to suck out the digestion products. 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 In physics, mathematics, engineering, and related fields, a wave is a propagating dynamic disturbance (change from List of types of equilibrium, equilibrium) of one or more quantities. ''Periodic waves'' oscillate repeatedly about an equilibrium ...
s detected by
receptor Receptor may refer to: * Sensory receptor, in physiology, any neurite structure that, on receiving environmental stimuli, produces an informative nerve impulse *Receptor (biochemistry), in biochemistry, a protein molecule that receives and respond ...
s, tufts of hairs on the sides of the two hindmost thoracic segments.


Pit-building species

Funnel-shaped pits are built by members of just 3 antlion tribes: Myrmeleontini, Myrmecaelurini, and Nesoleontini. In these trap-building species, an average-sized larva digs a pit about 2 in (5 cm) deep and 3 in (7.5 cm) wide at the edge. This behavior has also been observed in the
Vermileonidae The Brachyceran family Vermileonidae (the sole family in the infraorder Vermileonomorpha) is a small family of uncertain affinities and unusual biology. It includes fewer than 80 described species, most of them rare and with restricted distributi ...
(
Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advance ...
), whose larvae dig the same sort of pit to feed on ants. Having marked out the chosen site with a circular groove, 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. 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. By digging in a spiral when constructing its pit, the antlion minimises the time needed to complete the pit. 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. 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. The fine grain lining ensures that the avalanches which carry prey are as large as possible. Since the sides of the pit consist of loose sand at its angle of repose, 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. 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 Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s and other
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s, and can even subdue small
spider Spiders (order (biology), order Araneae) are air-breathing arthropods that have eight limbs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude spider silk, silk. They are the largest order of arachnids and ran ...
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. Antlion larvae require loose soil, not necessarily, but often, sand. Antlions can also handle larger granular material which is filtered out of the soil during pit construction. 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. Other arthropods may make use of the antlion larva's ability to trap prey. The larva of the Australian horsefly (''
Scaptia muscula ''Scaptia'' is a genus of horse-fly in the tribe Scionini. Species *'' Scaptia abdominalis'' (Ricardo, 1917) *'' Scaptia alpina'' Mackerras, 1960 *'' Scaptia arnhemensis'' Lessard, 2012 *'' Scaptia auranticula'' Mackerras, 1960 *'' Scaptia aura ...
'') lives in antlion (for example ''
Myrmeleon pictifrons ''Myrmeleon pictifrons'' is a species of antlion. It is native to the Australasian region and is one of over 2000 species of antlion that have been recorded globally. References

Myrmeleontinae Insects of Australia Taxa named by Carl Edua ...
'') pit traps and feeds on the prey caught, and the female
chalcid wasp Chalcid wasps (, , for their metallic colour) are insects within the superfamily Chalcidoidea, part of the order Hymenoptera. The superfamily contains some 22,500 known species, and an estimated total diversity of more than 500,000 species, m ...
('' Lasiochalcidia igiliensis'') purposefully allows itself to be trapped so that it can parasitise the antlion larva by ovipositing between its head and thorax. 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%. Furthermore, they appear to have maximized its usefulness—further increasing the duration is not likely to convey substantial survival benefits to the larvae.


Evolution

The closest living relatives of antlions within the
Myrmeleontoidea Myrmeleontoidea is a neuropteran superfamily in the clade Myrmeleontiformia. The following families are included: Superfamily Myrmeleontoidea (syn Nemopteroidea) * Family Ascalaphidae: owlflies (included in Myrmeleontidae in some classifications ...
are the
owlflies Ascalaphidae is a family of insects in the order Neuroptera, commonly called owlflies; there are some 450 extant species. They are fast-flying crepuscular or Diurnality, diurnal predators of other flying insects, and have large bulging eyes and ...
(Ascalaphidae); the
Nymphidae Nymphidae, sometimes called split-footed lacewings, are a family (biology), family of Pterygota, winged insects of the order (biology), order Neuroptera. There are 35 extant species native to Australia and New Guinea. Nymphidae stand somewhat ap ...
are more distantly related. The extinct Araripeneuridae and
Babinskaiidae Babinskaiidae is an extinct family of neuropterans known from the Cretaceous period of South America and Asia. They are part of the superfamily Myrmeleontoidea. Their distinguishing characters include: "long Insect morphology#Antennae, filiform a ...
are considered likely to be stem groups in the Myrmeleontiformia
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
. The
phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...
of the Neuroptera has been explored using
mitochondrial DNA Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondrion, mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the D ...
sequences, and while issues remain for the group as a whole (the "
Hemerobiiformia The Hemerobiiformia are a suborder of insects in the order Neuroptera that include most of the lacewings, Antlion, antlions and their allies. The phylogeny of the Neuroptera was explored in 2014 using mitochondrial DNA sequences. The results ind ...
" being
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
), the Myrmeleontiformia is generally agreed to be
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
, giving the following
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
: The
subfamilies In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zool ...
are shown below; a few genera, mostly
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
, are of uncertain or basal position. The fossil record of antlions is very small by neuropteran standards. However, some
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
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.


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 Palparinae is an antlion subfamily in the family Myrmeleontidae The antlions are a group of about 2,000 species of insect in the neuropteran family (biology), family Myrmeleontidae. They are known for the predation, predatory habits of their ...
, and
Myrmeleontinae Myrmeleontinae is a subfamily of Myrmeleontidae, the antlions. Tribes and Genera ''BioLib'' includes 3 tribes: ; Maulini Markl, 1954 # '' Isonemurus'' Esben-Petersen, 1928 # ''Maula'' - monotypic '' Maula stigmatus'' Navás, 1912 ; Myrmeleontini ...
. Brachynemurinae, Dendroleontinae, and other subfamilies recognized by prior authors were placed in the
Myrmeleontinae Myrmeleontinae is a subfamily of Myrmeleontidae, the antlions. Tribes and Genera ''BioLib'' includes 3 tribes: ; Maulini Markl, 1954 # '' Isonemurus'' Esben-Petersen, 1928 # ''Maula'' - monotypic '' Maula stigmatus'' Navás, 1912 ; Myrmeleontini ...
. * Stilbopteryginae Newman, 1853 # Stilbopterygini
monotypic: '' Stilbopteryx'' *
Palparinae Palparinae is an antlion subfamily in the family Myrmeleontidae The antlions are a group of about 2,000 species of insect in the neuropteran family (biology), family Myrmeleontidae. They are known for the predation, predatory habits of their ...
Banks, 1911 # Dimarini Navas, 1914 # Palparidiini Markl, 1954 # Palparini Banks, 1911 # Pseudimarini Markl 1954 *
Myrmeleontinae Myrmeleontinae is a subfamily of Myrmeleontidae, the antlions. Tribes and Genera ''BioLib'' includes 3 tribes: ; Maulini Markl, 1954 # '' Isonemurus'' Esben-Petersen, 1928 # ''Maula'' - monotypic '' Maula stigmatus'' Navás, 1912 ; Myrmeleontini ...
Latreille, 1802 # Acanthaclisini Navas, 1912 # Brachynemurini Banks, 1927 # Dendroleontini Banks, 1899 # Gnopholeontini Stange, 1994 # Lemolemini Stange, 1994 # Maulini Markl 1954 # Myrmecaelurini Esben-Petersen, 1918 # Myrmeleontini Latreille, 1802 # Nesoleontini Markl, 1954 # Nemoleontini Banks, 1911


Michel ''et al.'' (2017) classification

Michel ''et.al'' recognized just four subfamilies, consisting of
Acanthaclisinae Acanthaclisini is a tribe in the antlion subfamily Myrmeleontinae. The larvae of most species live in open tracts of fairly deep sand, and do not construct pitfall traps. They burrow beneath the surface of the sand which needs to be deep enou ...
,
Myrmeleontinae Myrmeleontinae is a subfamily of Myrmeleontidae, the antlions. Tribes and Genera ''BioLib'' includes 3 tribes: ; Maulini Markl, 1954 # '' Isonemurus'' Esben-Petersen, 1928 # ''Maula'' - monotypic '' Maula stigmatus'' Navás, 1912 ; Myrmeleontini ...
,
Palparinae Palparinae is an antlion subfamily in the family Myrmeleontidae The antlions are a group of about 2,000 species of insect in the neuropteran family (biology), family Myrmeleontidae. They are known for the predation, predatory habits of their ...
, and Stilbopteryginae. * Stilbopteryginae Newman, 1853 # Stilbopterygini Newman, 1853 *
Acanthaclisinae Acanthaclisini is a tribe in the antlion subfamily Myrmeleontinae. The larvae of most species live in open tracts of fairly deep sand, and do not construct pitfall traps. They burrow beneath the surface of the sand which needs to be deep enou ...
Navas, 1912 # Acanthaclisini Navas, 1912 *
Palparinae Palparinae is an antlion subfamily in the family Myrmeleontidae The antlions are a group of about 2,000 species of insect in the neuropteran family (biology), family Myrmeleontidae. They are known for the predation, predatory habits of their ...
Banks, 1911 # Dimarini Navas, 1914 # Palparidiini Markl, 1954 # Palparini Banks, 1911 # Pseudimarini Markl 1954 *
Myrmeleontinae Myrmeleontinae is a subfamily of Myrmeleontidae, the antlions. Tribes and Genera ''BioLib'' includes 3 tribes: ; Maulini Markl, 1954 # '' Isonemurus'' Esben-Petersen, 1928 # ''Maula'' - monotypic '' Maula stigmatus'' Navás, 1912 ; Myrmeleontini ...
Latreille, 1802 # Brachynemurini Banks, 1927 # Dendroleontini Banks, 1899 # Gepini Markl, 1954 # Gnopholeontini Stange, 1994 # Lemolemini Stange, 1994 # Maulini Markl 1954 # Myrmecaelurini Esben-Petersen, 1918 # Myrmeleontini Latreille, 1802 # Nesoleontini Markl, 1954


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:
Ascalaphinae Ascalaphinae is the type subfamily of the neuropteran owlfly family. Most species are found in the tropics. Their characteristic apomorphy , shared with the Ululodinae, is the ridge which divides each of their large compound eyes; both groups are ...
,
Myrmeleontinae Myrmeleontinae is a subfamily of Myrmeleontidae, the antlions. Tribes and Genera ''BioLib'' includes 3 tribes: ; Maulini Markl, 1954 # '' Isonemurus'' Esben-Petersen, 1928 # ''Maula'' - monotypic '' Maula stigmatus'' Navás, 1912 ; Myrmeleontini ...
, Dendroleontinae, and Nemoleontinae. *
Ascalaphinae Ascalaphinae is the type subfamily of the neuropteran owlfly family. Most species are found in the tropics. Their characteristic apomorphy , shared with the Ululodinae, is the ridge which divides each of their large compound eyes; both groups are ...
Rambur, 1842 # Ascalaphini Lefèbvre, 1842 # Dimarini Navas, 1914 # Haplogleniini Newman, 1853 # Palparini Banks, 1911 # Stilbopterygini Newman, 1853 #
Ululodini Ululodinae is a subfamily or tribe (as Ululodini) of owlflies. Both they and the Ascalaphinae are sometimes known as split-eyed owlflies due to the characteristic ridge that bisects their compound eyes A compound eye is a visual organ found i ...
Van der Weele 1909 *
Myrmeleontinae Myrmeleontinae is a subfamily of Myrmeleontidae, the antlions. Tribes and Genera ''BioLib'' includes 3 tribes: ; Maulini Markl, 1954 # '' Isonemurus'' Esben-Petersen, 1928 # ''Maula'' - monotypic '' Maula stigmatus'' Navás, 1912 ; Myrmeleontini ...
Latreille, 1802 # Acanthaclisini Navas, 1912 # Brachynemurini Banks, 1927 # Myrmecaelurini Esben-Petersen, 1918 # Myrmeleontini Latreille, 1802 # Nesoleontini Markl, 1954 * Dendroleontinae Banks, 1899 # Acanthoplectrini Markl, 1954 # Dendroleontini Banks, 1899 #
Glenurini The Nemoleontinae are a subfamily of ant-lions, erected by Nathan Banks in 1911. Tribes and genera Some authorities place genera elsewhere (such as the Myrmeleontinae) but ''BioLib''
Banks, 1927 # Megistopini Navas, 1912 * Nemoleontinae Banks, 1911 # Nemoleontini Banks, 1911 # Protoplectrini Tillyard, 1916


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''. 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." *
Myrmeleontinae Myrmeleontinae is a subfamily of Myrmeleontidae, the antlions. Tribes and Genera ''BioLib'' includes 3 tribes: ; Maulini Markl, 1954 # '' Isonemurus'' Esben-Petersen, 1928 # ''Maula'' - monotypic '' Maula stigmatus'' Navás, 1912 ; Myrmeleontini ...
Latreille, 1802 # Acanthaclisini Navas, 1912 # Brachynemurini Banks, 1927 # Dendroleontini Banks, 1899 # Myrmecaelurini Esben-Petersen, 1918 # Myrmeleontini Latreille, 1802 # Nemoleontini Banks, 1911 # Nesoleontini Markl, 1954 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 Spiders (order (biology), order Araneae) are air-breathing arthropods that have eight limbs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude spider silk, silk. They are the largest order of arachnids and ran ...
File:Antlion video.ogv, Video of a larva trapping an ant by throwing sand at it


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. Similar practices have been recorded from Africa, the Caribbean, China and Australia. The
Myrmecoleon The Myrmecoleon or Ant-lion is a fantastical animal from classical times, possibly derived from an error in the Septuagint version of the book of Job, reappearing in the Greek Christian ''Physiologus'' of the 3rd or 4th century A.D. It is found ...
was a mythical ant–lion hybrid written about in the 2nd century AD ''
Physiologus The ''Physiologus'' () is a didactic Christian text written or compiled in Greek by an unknown author in Alexandria. Its composition has been traditionally dated to the 2nd century AD by readers who saw parallels with writings of Clement of Alexa ...
'', 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 only eat meat, but the ant half from the mother could only eat grain chaff, thus the offspring could not eat either and would starve. It was paired with the Biblical verse Matthew 5:37. The fictional ant-lion of ''Physiologus'' is probably derived from a misreading of Job 4:11. The French naturalist
Jean-Henri Fabre Jean-Henri Casimir Fabre (; 21 December 1823 – 11 October 1915) was a French naturalist, entomologist, and author known for the lively style of his popular books on the lives of insects. Biography Fabre was born on 21 December 1823 in Saint- ...
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." In
Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...
, both the insect and its pit-traps are popularly known as . This term has since become a stock colloquialism for any "inescapable" trap, whether literal or metaphorical (''e.g.'' an unpleasant social obligation). Antlions appear as antagonists in the 1991
life simulation Life simulation games form a subgenre of simulation video games in which the player lives or controls one or more virtual characters (human or otherwise). Such a game can revolve around "individuals and relationships, or it could be a simulation ...
video game A video game or computer game is an electronic game that involves interaction with a user interface or input device (such as a joystick, game controller, controller, computer keyboard, keyboard, or motion sensing device) to generate visual fe ...
, ''
SimAnt ''SimAnt: The Electronic Ant Colony'' is a 1991 life simulation video game by Maxis and the company's third product, focusing on the lifecycle of ants. It was designed by Will Wright. In 1992, it was named "Best Simulation Game" at the Softwar ...
'', and (in a giant form) in the ''
Final Fantasy is a Japanese fantasy Anthology series, anthology media franchise created by Hironobu Sakaguchi which is owned, developed, and published by Square Enix (formerly Square (video game company), Square). The franchise centers on a series of fanta ...
'' series, '' Grounded'', ''
Terraria ''Terraria'' ( ) is a 2011 action-adventure sandbox game developed by Re-Logic and published by 505 Games. The game features exploration, combat, crafting, building, and mining inside a procedurally generated 2D computer graphics, 2D world. ...
'', '' Don't Starve Together'', ''
Monster Rancher 2 ''Monster Rancher 2'', known in Japan as , is a life simulation role-playing video game developed and published by Tecmo for the PlayStation. It is the second installment in the ''Monster Rancher'' series. It is the first game in the franchise t ...
'', ''
Mother 3 is a 2006 role-playing video game developed by Brownie Brown and HAL Laboratory and published by Nintendo for the Game Boy Advance. It is the third and final entry in the Mother (video game series), ''Mother'' series. The game follows Lucas, a ...
'' and in the ''
Half-Life 2 ''Half-Life 2'' is a 2004 first-person shooter game developed and published by Valve Corporation. It was published for Windows on Valve's digital distribution service, Steam. Like the original ''Half-Life'' (1998), ''Half-Life 2'' is played ent ...
'' video game series as an unrelated alien insect species sharing sand burowing traits with the real antlion larvae. The Trapinch, Vibrava, and Flygon
Pokémon is a Japanese media franchise consisting of List of Pokémon video games, video games, Pokémon (TV series), animated series and List of Pokémon films, films, Pokémon Trading Card Game, a trading card game, and other related media. The fran ...
evolution line is based on an antlion. The fictional
sarlacc The Sarlacc (plural Sarlacci) is a fictional creature in George Lucas's science fiction action film, sci-fi action saga ''Star Wars''. It first appeared in the film ''Return of the Jedi'' (1983) as a multi-tentacled fictional extraterrestrials ...
from the ''
Star Wars ''Star Wars'' is an American epic film, epic space opera media franchise created by George Lucas, which began with the Star Wars (film), eponymous 1977 film and Cultural impact of Star Wars, quickly became a worldwide popular culture, pop cu ...
'' franchise is often compared to the real-life antlion.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, . It also appears as a predator in the film '' Enemy Mine''. In the third book of
Tove Jansson Tove Marika Jansson (; 9 August 1914 – 27 June 2001) was a Swedish-speaking Finnish author, novelist, painter, illustrator and comic strip author. Brought up by artistic parents, Jansson studied art from 1930 to 1938 in Helsinki, Stockholm, ...
's
Moomins The Moomins (, ) are the central characters in a series of novels, short stories, picture books, and a comic strip by the Finnish writer and illustrator Tove Jansson, originally published in Swedish by Finnish publisher Schildts. They are a ...
series, ''
Finn Family Moomintroll ''Finn Family Moomintroll'' (original Swedish title ''Trollkarlens hatt'', ‘The Magician's Hat’; US edition ''The Happy Moomins'') is the third in the series of Tove Jansson's Moomins books, published in Swedish in 1948 and translated to Eng ...
'', a rather large and fanciful antlion appears in the second chapter, depicted as a sand-dwelling predator with the literal head of a
lion The lion (''Panthera leo'') is a large Felidae, cat of the genus ''Panthera'', native to Sub-Saharan Africa and India. It has a muscular, broad-chested body (biology), body; a short, rounded head; round ears; and a dark, hairy tuft at the ...
.


See also

*
List of Myrmeleontidae genera This is a list of genera in the family Myrmeleontidae, antlions. Extant Myrmeleontidae genera * ''Abatoleon'' Banks, 1924 * ''Acanthaclisis'' Rambur, 1842 * ''Acanthoplectron'' Peter Esben-Petersen, Esben-Petersen, 1918 * ''Acratoleon'' Banks, ...
*
Vermileonidae The Brachyceran family Vermileonidae (the sole family in the infraorder Vermileonomorpha) is a small family of uncertain affinities and unusual biology. It includes fewer than 80 described species, most of them rare and with restricted distributi ...
- wormlions


Notes


References


External links

{{Taxonbar, from=Q231439 Neuroptera Articles containing video clips