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Ant

May 20, 2023

For other uses, see Ant (disambiguation).

Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22,000 species have been classified. They are easily identified by their geniculate (elbowed) antennae and the distinctive node-like structure that forms their slender waists.

Ants
Temporal range: 100–0 Ma[1] Late Albian – Present
Fire ants
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Infraorder: Aculeata
Superfamily: Formicoidea
Family: Formicidae
Latreille, 1809
Type species
Formica rufa
Subfamilies
Cladogram of
subfamilies

A phylogeny of the extant ant subfamilies.[2][3]
*Cerapachyinae is paraphyletic
‡ The previous dorylomorph subfamilies were synonymized under Dorylinae by Brady et al. in 2014[4]

Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies that may occupy large territories and consist of millions of individuals. Larger colonies consist of various castes of sterile, wingless females, most of which are workers (ergates), as well as soldiers (dinergates) and other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens" (gynes). The colonies are described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.

(video) Ants gathering food

Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and a few remote or inhospitable islands. Ants thrive in moist tropical ecosystems and may exceed the combined biomass of wild birds and mammals. Their success in so many environments has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships.

Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study. Many human cultures make use of ants in cuisine, medication, and rites. Some species are valued in their role as biological pest control agents. Their ability to exploit resources may bring ants into conflict with humans, however, as they can damage crops and invade buildings. Some species, such as the red imported fire ant (Solenopsis invicta) of South America, are regarded as invasive species in other parts of the world, establishing themselves in areas where they have been introduced accidentally.

Etymology

The word ant and the archaic word emmet[5] are derived from ante, emete of Middle English, which come from ǣmette of Old English; these are all related to Low Saxon e(e)mt, empe and varieties (Old Saxon emeta) and to German Ameise (Old High German āmeiza). All of these words come from West Germanic *ǣmaitjōn, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut").[6][7]

The family name Formicidae is derived from the Latin formīca ("ant")[8] from which the words in other Romance languages, such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnică, and French fourmi are derived. It has been hypothesised that a Proto-Indo-European word *morwi- was the root for Sanskrit vamrah, Greek μύρμηξ mýrmēx, Old Church Slavonic mraviji, Old Irish moirb, Old Norse maurr, Dutch mier, Swedish myra, Danish myre, Middle Dutch miere, and Crimean Gothic miera.[9][10]

Taxonomy and evolution

Phylogenetic position of the Formicidae[11]

The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees, and wasps. Ants evolved from a lineage within the stinging wasps, and a 2013 study suggests that they are a sister group of the Apoidea.[11] In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma) that lived in the Cretaceous period. The specimen, trapped in amber dating back to around 92 million years ago, has features found in some wasps, but not found in modern ants.[12] The oldest fossils of ants date to the mid-Cretaceous, around 100 million years ago, which belong to extinct stem-groups such as the Haidomyrmecinae, Sphecomyrminae and Zigrasimeciinae, with modern ant subfamilies appearing towards the end of the Cretaceous around 80–70 million years ago.[13] Ants diversified and assumed ecological dominance around 60 million years ago.[14][1][15][16] Some groups, such as the Leptanillinae and Martialinae, are suggested to have diversified from early primitive ants that were likely to have been predators underneath the surface of the soil.[3][17]

 
Ants fossilised in Baltic amber

During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian supercontinent (the Northern Hemisphere). Their representation in the fossil record is poor, in comparison to the populations of other insects, representing only about 1% of fossil evidence of insects in the era. Ants became dominant after adaptive radiation at the beginning of the Paleogene period. By the Oligocene and Miocene, ants had come to represent 20–40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, around one in 10 genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene).[14][18]

Termites live in colonies and are sometimes called "white ants", but termites are only distantly related to ants. They are the sub-order Isoptera, and together with cockroaches, they form the order Blattodea. Blattodeans are related to mantids, crickets, and other winged insects that do not undergo full metamorphosis. Like ants, termites are eusocial, with sterile workers, but they differ greatly in the genetics of reproduction. The similarity of their social structure to that of ants is attributed to convergent evolution.[19] Velvet ants look like large ants, but are wingless female wasps.[20][21]

Distribution and diversity

Region Number of
species [22]
Neotropics 2,162
Nearctic 580
Europe 180
Africa 2,500
Asia 2,080
Melanesia 275
Australia 985
Polynesia 42

Ants have a cosmopolitan distribution. They are found on all continents except Antarctica, and only a few large islands, such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species.[23][24] Ants occupy a wide range of ecological niches and exploit many different food resources as direct or indirect herbivores, predators and scavengers. Most ant species are omnivorous generalists, but a few are specialist feeders. There is considerable variation in ant abundance across habitats, peaking in the moist tropics to nearly six times that found in less suitable habitats.[25] Their ecological dominance has been examined primarily using estimates of their biomass: myrmecologist E. O. Wilson had estimated in 2009 that at any one time the total number of ants was between one and ten quadrillion (short scale) (i.e., between 1015 and 1016) and using this estimate he had suggested that the total biomass of all the ants in the world was approximately equal to the total biomass of the entire human race.[26] More careful estimates made in 2022 which take into account regional variations puts the global ant contribution at 12 megatons of dry carbon, which is about 20% of the total human contribution, but greater than that of the wild birds and mammals combined. This study also puts a conservative estimate of the ants at about 20 × 1015 (20 quadrillion).[27][28][29]

Ants range in size from 0.75 to 52 millimetres (0.030–2.0 in),[30][31] the largest species being the fossil Titanomyrma giganteum, the queen of which was 6 cm (2+1⁄2 in) long with a wingspan of 15 cm (6 in).[32] Ants vary in colour; most ants are yellow to red or brown to black, but a few species are green and some tropical species have a metallic lustre. More than 13,800 species are currently known[33] (with upper estimates of the potential existence of about 22,000; see the article List of ant genera), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntWeb and the Hymenoptera Name Server, help to keep track of the known and newly described species.[33] The relative ease with which ants may be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.[34][35]

Morphology

 
Diagram of a worker ant (Neoponera verenae)

Ants are distinct in their morphology from other insects in having geniculate (elbowed) antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma, and metasoma are the three distinct body segments (formally tagmata). The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole may be formed by one or two nodes (the second alone, or the second and third abdominal segments).[36] Tergosternal fusion, when the tergite and sternite of a segment fuse together, can occur partly or fully on the second, third and fourth abdominal segment and is used in identification. Fourth abdominal tergosternal fusion was formerly used as character that defined the poneromorph subfamilies, Ponerinae and relatives within their clade, but this is no longer considered a synapomorphic character.[37]

Like other arthropods, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases, such as carbon dioxide, pass through their exoskeleton via tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph toward the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages.[38]

Head

 
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
 
Ant head

An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ant eyes are good for acute movement detection, but do not offer a high resolution image. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization.[39] Compared to vertebrates, ants tend to have blurrier eyesight, particularly in smaller species,[40] and a few subterranean taxa are completely blind.[2] However, some ants, such as Australia's bulldog ant, have excellent vision and are capable of discriminating the distance and size of objects moving nearly a meter away.[41]

Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents, and vibrations; they also are used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence.[38] In some species, a small pocket (infrabuccal chamber) inside the mouth stores food, so it may be passed to other ants or their larvae.[42]

Mesosoma

Both the legs and wings of the ant are attached to the mesosoma ("thorax"). The legs terminate in a hooked claw which allows them to hook on and climb surfaces.[43] Only reproductive ants (queens and males) have wings. Queens shed their wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. In a few species, wingless queens (ergatoids) and males occur.[38]

Metasoma

The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae), and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.[38]

Polymorphism

 
Seven leafcutter ant workers of various castes (left) and two queens (right)

In the colonies of a few ant species, there are physical castes—workers in distinct size-classes, called minor, median, and major ergates. Often, the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. These are known as macrergates while smaller workers are known as micrergates.[44] Although formally known as dinergates, such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they still are workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species, the median workers are absent, creating a sharp divide between the minors and majors.[45] Weaver ants, for example, have a distinct bimodal size distribution.[46][47] Some other species show continuous variation in the size of workers. The smallest and largest workers in Carebara diversa show nearly a 500-fold difference in their dry weights.[48]

Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile, haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes.[49] For instance, these replete workers develop in the North American honeypot ant Myrmecocystus mexicanus. Usually the largest workers in the colony develop into repletes; and, if repletes are removed from the colony, other workers become repletes, demonstrating the flexibility of this particular polymorphism.[50] This polymorphism in morphology and behaviour of workers initially was thought to be determined by environmental factors such as nutrition and hormones that led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp.[51] These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens.[52] The Australian jack jumper ant (Myrmecia pilosula) has only a single pair of chromosomes (with the males having just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.[53][54]

Genome size

Genome size is a fundamental characteristic of an organism. Ants have been found to have tiny genomes, with the evolution of genome size suggested to occur through loss and accumulation of non-coding regions, mainly transposable elements, and occasionally by whole genome duplication.[55] This may be related to colonisation processes, but further studies are needed to verify this.[55]

Life cycle

 
Meat eater ant nest during swarming

The life of an ant starts from an egg; if the egg is fertilised, the progeny will be female diploid, if not, it will be male haploid. Ants develop by complete metamorphosis with the larva stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach". Larvae, especially in the later stages, may also be provided solid food, such as trophic eggs, pieces of prey, and seeds brought by workers.[56]

The larvae grow through a series of four or five moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa.[57] The differentiation into queens and workers (which are both female), and different castes of workers, is influenced in some species by the nutrition the larvae obtain. Genetic influences and the control of gene expression by the developmental environment are complex and the determination of caste continues to be a subject of research.[58] Winged male ants, called drones (termed "aner" in old literature[59]), emerge from pupae along with the usually winged breeding females. Some species, such as army ants, have wingless queens. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so often are moved around among the various brood chambers within the colony.[60]

A new ergate spends the first few days of its adult life caring for the queen and young. She then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants who are older and are likely to die soon of natural causes.[61][62]

Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, being quite short-lived and surviving for only a few weeks.[63] Ant queens are estimated to live 100 times as long as solitary insects of a similar size.[64]

Ants are active all year long in the tropics, but, in cooler regions, they survive the winter in a state of dormancy known as hibernation. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.[65]

 
Alate male ant, Prenolepis imparis

Reproduction

 
Honey ants (Prenolepis imparis) mating

A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis.[66] Secretions from the male accessory glands in some species can plug the female genital opening and prevent females from re-mating.[67] Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens, while others may exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right.[68]

Drones can also mate with existing queens by entering a foreign colony, such as in army ants. When the drone is initially attacked by the workers, it releases a mating pheromone. If recognized as a mate, it will be carried to the queen to mate.[69] Males may also patrol the nest and fight others by grabbing them with their mandibles, piercing their exoskeleton and then marking them with a pheromone. The marked male is interpreted as an invader by worker ants and is killed.[70]

Most ants are univoltine, producing a new generation each year.[71] During the species-specific breeding period, winged females and winged males, known to entomologists as alates, leave the colony in what is called a nuptial flight. The nuptial flight usually takes place in the late spring or early summer when the weather is hot and humid. Heat makes flying easier and freshly fallen rain makes the ground softer for mated queens to dig nests.[72] Males typically take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Males will mount females in the air, but the actual mating process usually takes place on the ground. Females of some species mate with just one male but in others they may mate with as many as ten or more different males, storing the sperm in their spermathecae.[73] In Cardiocondyla elegans, workers may transport newly emerged queens to other conspecific nests where wingless males from unrelated colonies can mate with them, a behavioural adaptation that may reduce the chances of inbreeding.[74]

 
Fertilised meat-eater ant queen beginning to dig a new colony

Mated females then seek a suitable place to begin a colony. There, they break off their wings using their tibial spurs and begin to lay and care for eggs. The females can selectively fertilise future eggs with the sperm stored to produce diploid workers or lay unfertilized haploid eggs to produce drones. The first workers to hatch, known as nanitics,[75] are weaker and smaller than later workers but they begin to serve the colony immediately. They enlarge the nest, forage for food, and care for the other eggs. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site,[73] a process akin to swarming in honeybees.

Behaviour and ecology

Communication

 
Two Camponotus sericeus workers communicating through touch and pheromones

Ants communicate with each other using pheromones, sounds, and touch.[76] Since most ants live on the ground, they use the soil surface to leave pheromone trails that may be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually identifying the best path.[76][77]

Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from farther away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves.[78] Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum, and hind tibia.[64] Pheromones also are exchanged, mixed with food, and passed by trophallaxis, transferring information within the colony.[79] This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to.[80] In ant species with queen castes, when the dominant queen stops producing a specific pheromone, workers begin to raise new queens in the colony.[81]

Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.[82][83]

Defence

See also: Defense in insects
 
A Plectroctena sp. attacks another of its kind to protect its territory.

Ants attack and defend themselves by biting and, in many species, by stinging often injecting or spraying chemicals. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt sting pain index.[84]

The sting of jack jumper ants can be lethal for humans,[85] and an antivenom has been developed for it.[86] Fire ants, Solenopsis spp., are unique in having a venom sac containing piperidine alkaloids.[87] Their stings are painful and can be dangerous to hypersensitive people.[88] Formicine ants secrete a poison from their glands, made mainly of formic acid.[89]

 
A weaver ant in fighting position, mandibles wide open

Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom.[90] One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 and 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backward to escape a threat.[90] Before striking, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory organs resembling hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws also are seen in other ponerines such as Anochetus, as well as some genera in the tribe Attini, such as Daceton, Orectognathus, and Strumigenys,[90][91] which are viewed as examples of convergent evolution.

A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. If combat takes a turn for the worse, a worker may perform a final act of suicidal altruism by rupturing the membrane of its gaster, causing the content of its mandibular glands to burst from the anterior region of its head, spraying a poisonous, corrosive secretion containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies.[92]

Suicidal defences by workers are also noted in a Brazilian ant, Forelius pusillus, where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.[93]

 
Ant mound holes prevent water from entering the nest during rain.

In addition to defence against predators, ants need to protect their colonies from pathogens. Secretions from the metapleural gland, unique to the ants, produce a complex range of chemicals including several with antibiotic properties.[94] Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophoresis, the disposal of dead nest-mates.[95] Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana[96] while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates to trigger similar behaviour.[97]

Nests may be protected from physical threats such as flooding and overheating by elaborate nest architecture.[98][99] Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside.[100] Camponotus anderseni, which nests in the cavities of wood in mangrove habitats, deals with submergence under water by switching to anaerobic respiration.[101]

Learning

 
Two Weaver ants walking in tandem

Many animals can learn behaviours by imitation, but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis can lead a naïve nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. The leader is acutely sensitive to the progress of the follower and slows down when the follower lags and speeds up when the follower gets too close.[102]

Controlled experiments with colonies of Cerapachys biroi suggest that an individual may choose nest roles based on her previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out fewer and fewer times. A month later, the successful foragers continued in their role while the others had moved to specialise in brood care.[103]

Nest construction

Main article: Ant colony

Complex nests are built by many ant species, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests may be found in the ground, under stones or logs, inside logs, hollow stems, or even acorns. The materials used for construction include soil and plant matter,[73] and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.[104]

The army ants of South America, such as the Eciton burchellii species, and the driver ants of Africa do not build permanent nests, but instead, alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together.[105]

Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.[106]

 
Ant bridge

Formica polyctena, among other ant species, constructs nests that maintain a relatively constant interior temperature that aids in the development of larvae. The ants maintain the nest temperature by choosing the location, nest materials, controlling ventilation and maintaining the heat from solar radiation, worker activity and metabolism, and in some moist nests, microbial activity in the nest materials.[107][108]

Some ant species, such as those that use natural cavities, can be opportunistic and make use of the controlled micro-climate provided inside human dwellings and other artificial structures to house their colonies and nest structures.[109][110]

Cultivation of food

Main article: Ant–fungus mutualism
 
Myrmecocystus, honeypot ants, store food to prevent colony famine.

Most ants are generalist predators, scavengers, and indirect herbivores,[15] but a few have evolved specialised ways of obtaining nutrition. It is believed that many ant species that engage in indirect herbivory rely on specialized symbiosis with their gut microbes[111] to upgrade the nutritional value of the food they collect[112] and allow them to survive in nitrogen poor regions, such as rainforest canopies.[113] Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Ergates specialise in related tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is found to be toxic to the fungus, the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria introduced into the nest that may harm the fungi.[114]

Navigation

 
An ant trail

Foraging ants travel distances of up to 200 metres (700 ft) from their nest [115] and scent trails allow them to find their way back even in the dark. In hot and arid regions, day-foraging ants face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants of the genus Cataglyphis such as the Sahara desert ant navigate by keeping track of direction as well as distance travelled. Distances travelled are measured using an internal pedometer that keeps count of the steps taken[116] and also by evaluating the movement of objects in their visual field (optical flow).[117] Directions are measured using the position of the sun.[118] They integrate this information to find the shortest route back to their nest.[119] Like all ants, they can also make use of visual landmarks when available[120] as well as olfactory and tactile cues to navigate.[121][122] Some species of ant are able to use the Earth's magnetic field for navigation.[123] The compound eyes of ants have specialised cells that detect polarised light from the Sun, which is used to determine direction.[124][125] These polarization detectors are sensitive in the ultraviolet region of the light spectrum.[126] In some army ant species, a group of foragers who become separated from the main column may sometimes turn back on themselves and form a circular ant mill. The workers may then run around continuously until they die of exhaustion.[127]

Locomotion

The female worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs.[128] There are several species of gliding ant including Cephalotes atratus; this may be a common trait among arboreal ants with small colonies. Ants with this ability are able to control their horizontal movement so as to catch tree trunks when they fall from atop the forest canopy.[129]

Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods.[130] These rafts may also have a role in allowing ants to colonise islands.[131] Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they go to trapped pockets of air in the submerged nests to breathe.[132]

Cooperation and competition

 
Meat-eater ants feeding on a cicada: social ants cooperate and collectively gather food

Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal of ants. Like virtually all ants, they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using her large eyes instead of chemical senses to find prey.[133]

Some species attack and take over neighbouring ant colonies. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive.[134] Captured workers of enslaved Temnothorax species have evolved a counter-strategy, destroying just the female pupae of the slave-making Temnothorax americanus, but sparing the males (who do not take part in slave-raiding as adults).[135]

 
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen (on top)

Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.[136]

Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species such as Strumigenys xenos are entirely parasitic and do not have workers, but instead, rely on the food gathered by their Strumigenys perplexa hosts.[137][138] This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen may enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest.[139]

A conflict between the sexes of a species is seen in some species of ants with these reproducers apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process whereby a diploid egg loses its maternal contribution to produce haploid males who are clones of the father.[140]

Relationships with other organisms

 
The spider Myrmarachne plataleoides (female shown) mimics weaver ants to avoid predators.

Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They also are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae, and eggs, consuming the food stores of the ants, or avoiding predators. These inquilines may bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.[141][142]

 
An ant collects honeydew from an aphid

Aphids and other hemipteran insects secrete a sweet liquid called honeydew, when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect.[143] In some cases, the aphids secrete the honeydew in response to ants tapping them with their antennae. The ants in turn keep predators away from the aphids and will move them from one feeding location to another. When migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs may become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.[144]

Myrmecophilous (ant-loving) caterpillars of the butterfly family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants.[145] A similar adaptation can be seen in Grizzled skipper butterflies that emit vibrations by expanding their wings in order to communicate with ants, which are natural predators of these butterflies.[146] Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ant nest where it feeds on the ant larvae.[147] A number of specialized bacteria have been found as endosymbionts in ant guts. Some of the dominant bacteria belong to the order Hyphomicrobiales whose members are known for being nitrogen-fixing symbionts in legumes but the species found in ant lack the ability to fix nitrogen.[148][149] Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the genera Leucoagaricus or Leucocoprinus of the family Agaricaceae. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant, Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.[150]

 
Ants may obtain nectar from flowers such as the dandelion, but are only rarely known to pollinate flowers.

Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees, (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees.[151] Although some ants obtain nectar from flowers, pollination by ants is somewhat rare, one example being of the pollination of the orchid Leporella fimbriata which induces male Myrmecia urens to pseudocopulate with the flowers, transferring pollen in the process.[152] One theory that has been proposed for the rarity of pollination is that the secretions of the metapleural gland inactivate and reduce the viability of pollen.[153][154] Some plants have special nectar exuding structures, extrafloral nectaries, that provide food for ants, which in turn protect the plant from more damaging herbivorous insects.[155] Species such as the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) who defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants.[156] In return, the ants obtain food from protein- and lipid-rich Beltian bodies. In Fiji Philidris nagasau (Dolichoderinae) are known to selectively grow species of epiphytic Squamellaria (Rubiaceae) which produce large domatia inside which the ant colonies nest. The ants plant the seeds and the domatia of young seedling are immediately occupied and the ant faeces in them contribute to rapid growth.[157] Similar dispersal associations are found with other dolichoderines in the region as well.[158] Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.[159]

Many plant species have seeds that are adapted for dispersal by ants.[160] Seed dispersal by ants or myrmecochory is widespread, and new estimates suggest that nearly 9% of all plant species may have such ant associations.[161][160] Often, seed-dispersing ants perform directed dispersal, depositing the seeds in locations that increase the likelihood of seed survival to reproduction.[162] Some plants in arid, fire-prone systems are particularly dependent on ants for their survival and dispersal as the seeds are transported to safety below the ground.[163] Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food.[164] Ants can substantially alter rate of decomposition and nutrient cycling in their nest.[165][166] By myrmecochory and modification of soil conditions they substantially alter vegetation and nutrient cycling in surrounding ecosystem.[167]

A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.[168]

 
A meat ant tending a common leafhopper nymph

Most ants are predatory and some prey on and obtain food from other social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants.[115] Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away predatory ant species.[169] The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellent chemical.[170] It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Other wasps, such as A. multipicta, defend against ants by blasting them off the nest with bursts of wing buzzing.[171] Stingless bees (Trigona and Melipona) use chemical defences against ants.[115]

Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants.[172] Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.[172]

 
Oecophylla smaragdina killed by a fungus

Fungi in the genera Cordyceps and Ophiocordyceps infect ants. Ants react to their infection by climbing up plants and sinking their mandibles into plant tissue. The fungus kills the ants, grows on their remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores [173] in a microhabitat that best suits the fungus.[174] Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates.[175]

A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black-coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, causing them to carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits, such as Hyeronima alchorneoides, and eaten. The droppings of the bird are collected by other ants and fed to their young, leading to further spread of the nematode.[176]

 
Spiders (Like this Menemerus jumping spider) sometimes feed on ants

A study of Temnothorax nylanderi colonies in Germany found that workers parasitized by the tapeworm Anomotaenia brevis (ants are intermediate hosts, the definitive hosts are woodpeckers) lived much longer than unparasitized workers and had a reduced mortality rate, comparable to that of the queens of the same species, which live for as long as two decades.[177]

South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants.[178]

Army ants forage in a wide roving column, attacking any animals in that path that are unable to escape. In Central and South America, Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers.[179][180] This behaviour was once considered mutualistic, but later studies found the birds to be parasitic. Direct kleptoparasitism (birds stealing food from the ants' grasp) is rare and has been noted in Inca doves which pick seeds at nest entrances as they are being transported by species of Pogonomyrmex.[181] Birds that follow ants eat many prey insects and thus decrease the foraging success of ants.[182] Birds indulge in a peculiar behaviour called anting that, as yet, is not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may be a means to remove ectoparasites from the birds.

Anteaters, aardvarks, pangolins, echidnas and numbats have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants. About 12%, 16%, and 4% of their faecal volume in spring, summer and autumn, respectively, is composed of ants.[183]

Relationship with humans

 
Weaver ants are used as a biological control for citrus cultivation in southern China.

Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control.[184] On the other hand, ants may become nuisances when they invade buildings or cause economic losses.

In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.[185][186][187] The large heads of the dinergates (soldiers) of the leafcutting ant Atta cephalotes are also used by native surgeons in closing wounds.[188]

Some ants have toxic venom and are of medical importance. The species include Paraponera clavata (tocandira) and Dinoponera spp. (false tocandiras) of South America [189] and the Myrmecia ants of Australia.[190]

In South Africa, ants are used to help harvest the seeds of rooibos (Aspalathus linearis), a plant used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds may be collected from one ant-heap.[191][192]

Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These tend to be island species that have evolved specialized traits and risk being displaced by introduced ant species. Examples include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.[193]

As food

See also: Entomophagy
 
Roasted ants in Colombia
 
Ant larvae for sale in Isaan, Thailand

Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are used in Mexican escamoles. They are considered a form of insect caviar and can sell for as much as US$50 per kg going up to US$200 per kg (as of 2006) because they are seasonal and hard to find.[194] In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.[195] In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry.[196] Weaver ant eggs and larvae, as well as the ants, may be used in a Thai salad, yam (Thai: ยำ), in a dish called yam khai mot daeng (Thai: ยำไข่มดแดง) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".[197]

In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickling, acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the repletes, or living honey-pots, of the honey ant (Myrmecocystus).[197]

As pests

See also: Ants of medical importance
 
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.

Some ant species are considered as pests, primarily those that occur in human habitations, where their presence is often problematic. For example, the presence of ants would be undesirable in sterile places such as hospitals or kitchens. Some species or genera commonly categorized as pests include the Argentine ant, immigrant pavement ant, yellow crazy ant, banded sugar ant, pharaoh ant, red wood ant, black carpenter ant, odorous house ant, red imported fire ant, and European fire ant. Some ants will raid stored food, some will seek water sources, others may damage indoor structures, some may damage agricultural crops directly or by aiding sucking pests. Some will sting or bite.[198] The adaptive nature of ant colonies make it nearly impossible to eliminate entire colonies and most pest management practices aim to control local populations and tend to be temporary solutions. Ant populations are managed by a combination of approaches that make use of chemical, biological, and physical methods. Chemical methods include the use of insecticidal bait which is gathered by ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Management is based on the species and techniques may vary according to the location and circumstance.[198]

In science and technology

See also: Myrmecology, Biomimetics, and Ant colony optimization algorithms
 
Camponotus nearcticus workers travelling between two formicaria through connector tubing

Observed by humans since the dawn of history, the behaviour of ants has been documented and the subject of early writings and fables passed from one century to another. Those using scientific methods, myrmecologists, study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbock in 1881.[199] Studies on ants have tested hypotheses in ecology and sociobiology, and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies.[200] Ant colonies may be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures.[201] Individuals may be tracked for study by marking them with dots of colours.[202]

The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems, for example Ant colony optimization and Ant robotics. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage, and networking algorithms.[203]

As pets

Main article: Ant-keeping

From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Some later commercial versions use transparent gel instead of soil, allowing greater visibility at the cost of stressing the ants with unnatural light.[204]

In culture

 
Aesop's ants: illustration by Milo Winter, 1888–1956

Anthropomorphised ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They also are mentioned in religious texts.[205][206] In the Book of Proverbs in the Bible, ants are held up as a good example of hard work and cooperation.[207] Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. [Quran 27:18],[208][209] In parts of Africa, ants are considered to be the messengers of the deities. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief.[210] Ant bites are used in the initiation ceremonies of some Amazon Indian cultures as a test of endurance.[211][212] In Greek mythology, the goddess Athena turned the maiden Myrmex into an ant when the latter claimed to have invented the plough, when in fact it was Athena's own invention.[213]

 
An ant pictured in the coat of arms of Multia, a town in Finland

Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his 1880 book A Tramp Abroad.[214] Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. H.G. Wells wrote about intelligent ants destroying human settlements in Brazil and threatening human civilization in his 1905 science-fiction short story, The Empire of the Ants. In more recent times, animated cartoons and 3-D animated films featuring ants have been produced including Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Ferdy the Ant and Atom Ant. Renowned myrmecologist E. O. Wilson wrote a short story, "Trailhead" in 2010 for The New Yorker magazine, which describes the life and death of an ant-queen and the rise and fall of her colony, from an ants' point of view.[215] The French neuroanatomist, psychiatrist and eugenicist Auguste Forel believed that ant societies were models for human society. He published a five volume work from 1921 to 1923 that examined ant biology and society.[216]

In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".[217]

Ants also are quite popular inspiration for many science-fiction insectoids, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the films Them! and Empire of the Ants, Marvel Comics' super hero Ant-Man, and ants mutated into super-intelligence in Phase IV. In computer strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.[218]

See also

Main article: Outline of ants

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Cited texts

Further reading

  • Mike Snider (8 Jan 2022). "When the trees where these ants live were damaged, they made some DIY home repairs". USA Today.
  • Bolton, Barry (1995). A New General Catalogue of the Ants of the World. Harvard University Press. ISBN 978-0-674-61514-4.
  • Hölldobler B, Wilson EO (1998). Journey to the Ants: A Story of Scientific Exploration. Belknap Press. ISBN 978-0-674-48526-6.
  • Hölldobler B, Wilson EO (2009). The Superorganism: The Beauty, Elegance and Strangeness of Insect Societies. Norton & Co. ISBN 978-0-393-06704-0.

External links

 
Wikiquote has quotations related to Ant.
 
Wikimedia Commons has media related to Formicidae.
 
Wikispecies has information related to Formicidae.
  • AntWeb from The California Academy of Sciences
  • AntWiki – Bringing Ants to the World
  • Ant Species Fact Sheets from the National Pest Management Association on Argentine, Carpenter, Pharaoh, Odorous, and other ant species
  • Ant Genera of the World – distribution maps
  • The super-nettles. A dermatologist's guide to ants-in-the-plants

May 20, 2023
other, uses, disambiguation, eusocial, insects, family, formicidae, along, with, related, wasps, bees, belong, order, hymenoptera, evolved, from, vespoid, wasp, ancestors, cretaceous, period, more, than, estimated, total, species, have, been, classified, they,. For other uses see Ant disambiguation Ants are eusocial insects of the family Formicidae and along with the related wasps and bees belong to the order Hymenoptera Ants evolved from vespoid wasp ancestors in the Cretaceous period More than 13 800 of an estimated total of 22 000 species have been classified They are easily identified by their geniculate elbowed antennae and the distinctive node like structure that forms their slender waists AntsTemporal range 100 0 Ma 1 PreꞒ Ꞓ O S D C P T J K Pg N Late Albian PresentFire antsScientific classificationKingdom AnimaliaPhylum ArthropodaClass InsectaOrder HymenopteraInfraorder AculeataSuperfamily FormicoideaFamily FormicidaeLatreille 1809Type speciesFormica rufaLinnaeus 1761SubfamiliesAgroecomyrmecinae Amblyoponinae incl Apomyrminae Aneuretinae Brownimeciinae Dolichoderinae Dorylinae Ectatomminae Formiciinae Formicinae Haidomyrmecinae Heteroponerinae Leptanillinae Martialinae Myrmeciinae incl Nothomyrmeciinae Myrmicinae Paraponerinae Ponerinae Proceratiinae Pseudomyrmecinae Sphecomyrminae ZigrasimeciinaeCladogram of subfamiliesMartialinaeLeptanillinaeAmblyoponinaeParaponerinaeAgroecomyrmecinaePonerinaeProceratiinaeEcitoninae Aenictinae Dorylini Aenictogitoninae Cerapachyinae Leptanilloidinae DolichoderinaeAneuretinaePseudomyrmecinaeMyrmeciinaeEctatomminaeHeteroponerinaeMyrmicinaeFormicinaeA phylogeny of the extant ant subfamilies 2 3 Cerapachyinae is paraphyletic The previous dorylomorph subfamilies were synonymized under Dorylinae by Brady et al in 2014 4 Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies that may occupy large territories and consist of millions of individuals Larger colonies consist of various castes of sterile wingless females most of which are workers ergates as well as soldiers dinergates and other specialised groups Nearly all ant colonies also have some fertile males called drones and one or more fertile females called queens gynes The colonies are described as superorganisms because the ants appear to operate as a unified entity collectively working together to support the colony source source source source source source source source source source source source source source video Ants gathering food Ants have colonised almost every landmass on Earth The only places lacking indigenous ants are Antarctica and a few remote or inhospitable islands Ants thrive in moist tropical ecosystems and may exceed the combined biomass of wild birds and mammals Their success in so many environments has been attributed to their social organisation and their ability to modify habitats tap resources and defend themselves Their long co evolution with other species has led to mimetic commensal parasitic and mutualistic relationships Ant societies have division of labour communication between individuals and an ability to solve complex problems These parallels with human societies have long been an inspiration and subject of study Many human cultures make use of ants in cuisine medication and rites Some species are valued in their role as biological pest control agents Their ability to exploit resources may bring ants into conflict with humans however as they can damage crops and invade buildings Some species such as the red imported fire ant Solenopsis invicta of South America are regarded as invasive species in other parts of the world establishing themselves in areas where they have been introduced accidentally Contents 1 Etymology 2 Taxonomy and evolution 3 Distribution and diversity 4 Morphology 4 1 Head 4 2 Mesosoma 4 3 Metasoma 4 4 Polymorphism 4 5 Genome size 5 Life cycle 5 1 Reproduction 6 Behaviour and ecology 6 1 Communication 6 2 Defence 6 3 Learning 6 4 Nest construction 6 5 Cultivation of food 6 6 Navigation 6 7 Locomotion 6 8 Cooperation and competition 6 9 Relationships with other organisms 7 Relationship with humans 7 1 As food 7 2 As pests 7 3 In science and technology 7 4 As pets 7 5 In culture 8 See also 9 References 9 1 Cited texts 10 Further reading 11 External linksEtymologyThe word ant and the archaic word emmet 5 are derived from ante emete of Middle English which come from ǣmette of Old English these are all related to Low Saxon e e mt empe and varieties Old Saxon emeta and to German Ameise Old High German ameiza All of these words come from West Germanic ǣmaitjōn and the original meaning of the word was the biter from Proto Germanic ai off away mait cut 6 7 The family name Formicidae is derived from the Latin formica ant 8 from which the words in other Romance languages such as the Portuguese formiga Italian formica Spanish hormiga Romanian furnică and French fourmi are derived It has been hypothesised that a Proto Indo European word morwi was the root for Sanskrit vamrah Greek myrmh3 myrmex Old Church Slavonic mraviji Old Irish moirb Old Norse maurr Dutch mier Swedish myra Danish myre Middle Dutch miere and Crimean Gothicmiera 9 10 Taxonomy and evolutionAculeata Chrysidoidea VespidaeRhopalosomatidae PompilidaeTiphiidae Scolioidea ApoideaFormicidaePhylogenetic position of the Formicidae 11 The family Formicidae belongs to the order Hymenoptera which also includes sawflies bees and wasps Ants evolved from a lineage within the stinging wasps and a 2013 study suggests that they are a sister group of the Apoidea 11 In 1966 E O Wilson and his colleagues identified the fossil remains of an ant Sphecomyrma that lived in the Cretaceous period The specimen trapped in amber dating back to around 92 million years ago has features found in some wasps but not found in modern ants 12 The oldest fossils of ants date to the mid Cretaceous around 100 million years ago which belong to extinct stem groups such as the Haidomyrmecinae Sphecomyrminae and Zigrasimeciinae with modern ant subfamilies appearing towards the end of the Cretaceous around 80 70 million years ago 13 Ants diversified and assumed ecological dominance around 60 million years ago 14 1 15 16 Some groups such as the Leptanillinae and Martialinae are suggested to have diversified from early primitive ants that were likely to have been predators underneath the surface of the soil 3 17 Ants fossilised in Baltic amber During the Cretaceous period a few species of primitive ants ranged widely on the Laurasian supercontinent the Northern Hemisphere Their representation in the fossil record is poor in comparison to the populations of other insects representing only about 1 of fossil evidence of insects in the era Ants became dominant after adaptive radiation at the beginning of the Paleogene period By the Oligocene and Miocene ants had come to represent 20 40 of all insects found in major fossil deposits Of the species that lived in the Eocene epoch around one in 10 genera survive to the present Genera surviving today comprise 56 of the genera in Baltic amber fossils early Oligocene and 92 of the genera in Dominican amber fossils apparently early Miocene 14 18 Termites live in colonies and are sometimes called white ants but termites are only distantly related to ants They are the sub order Isoptera and together with cockroaches they form the order Blattodea Blattodeans are related to mantids crickets and other winged insects that do not undergo full metamorphosis Like ants termites are eusocial with sterile workers but they differ greatly in the genetics of reproduction The similarity of their social structure to that of ants is attributed to convergent evolution 19 Velvet ants look like large ants but are wingless female wasps 20 21 Distribution and diversityRegion Number ofspecies 22 Neotropics 2 162Nearctic 580Europe 180Africa 2 500Asia 2 080Melanesia 275Australia 985Polynesia 42Ants have a cosmopolitan distribution They are found on all continents except Antarctica and only a few large islands such as Greenland Iceland parts of Polynesia and the Hawaiian Islands lack native ant species 23 24 Ants occupy a wide range of ecological niches and exploit many different food resources as direct or indirect herbivores predators and scavengers Most ant species are omnivorous generalists but a few are specialist feeders There is considerable variation in ant abundance across habitats peaking in the moist tropics to nearly six times that found in less suitable habitats 25 Their ecological dominance has been examined primarily using estimates of their biomass myrmecologist E O Wilson had estimated in 2009 that at any one time the total number of ants was between one and ten quadrillion short scale i e between 1015 and 1016 and using this estimate he had suggested that the total biomass of all the ants in the world was approximately equal to the total biomass of the entire human race 26 More careful estimates made in 2022 which take into account regional variations puts the global ant contribution at 12 megatons of dry carbon which is about 20 of the total human contribution but greater than that of the wild birds and mammals combined This study also puts a conservative estimate of the ants at about 20 1015 20 quadrillion 27 28 29 Ants range in size from 0 75 to 52 millimetres 0 030 2 0 in 30 31 the largest species being the fossil Titanomyrma giganteum the queen of which was 6 cm 2 1 2 in long with a wingspan of 15 cm 6 in 32 Ants vary in colour most ants are yellow to red or brown to black but a few species are green and some tropical species have a metallic lustre More than 13 800 species are currently known 33 with upper estimates of the potential existence of about 22 000 see the article List of ant genera with the greatest diversity in the tropics Taxonomic studies continue to resolve the classification and systematics of ants Online databases of ant species including AntWeb and the Hymenoptera Name Server help to keep track of the known and newly described species 33 The relative ease with which ants may be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies 34 35 Morphology Diagram of a worker ant Neoponera verenae Ants are distinct in their morphology from other insects in having geniculate elbowed antennae metapleural glands and a strong constriction of their second abdominal segment into a node like petiole The head mesosoma and metasoma are the three distinct body segments formally tagmata The petiole forms a narrow waist between their mesosoma thorax plus the first abdominal segment which is fused to it and gaster abdomen less the abdominal segments in the petiole The petiole may be formed by one or two nodes the second alone or the second and third abdominal segments 36 Tergosternal fusion when the tergite and sternite of a segment fuse together can occur partly or fully on the second third and fourth abdominal segment and is used in identification Fourth abdominal tergosternal fusion was formerly used as character that defined the poneromorph subfamilies Ponerinae and relatives within their clade but this is no longer considered a synapomorphic character 37 Like other arthropods ants have an exoskeleton an external covering that provides a protective casing around the body and a point of attachment for muscles in contrast to the internal skeletons of humans and other vertebrates Insects do not have lungs oxygen and other gases such as carbon dioxide pass through their exoskeleton via tiny valves called spiracles Insects also lack closed blood vessels instead they have a long thin perforated tube along the top of the body called the dorsal aorta that functions like a heart and pumps haemolymph toward the head thus driving the circulation of the internal fluids The nervous system consists of a ventral nerve cord that runs the length of the body with several ganglia and branches along the way reaching into the extremities of the appendages 38 Head Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision Ant head An ant s head contains many sensory organs Like most insects ants have compound eyes made from numerous tiny lenses attached together Ant eyes are good for acute movement detection but do not offer a high resolution image They also have three small ocelli simple eyes on the top of the head that detect light levels and polarization 39 Compared to vertebrates ants tend to have blurrier eyesight particularly in smaller species 40 and a few subterranean taxa are completely blind 2 However some ants such as Australia s bulldog ant have excellent vision and are capable of discriminating the distance and size of objects moving nearly a meter away 41 Two antennae feelers are attached to the head these organs detect chemicals air currents and vibrations they also are used to transmit and receive signals through touch The head has two strong jaws the mandibles used to carry food manipulate objects construct nests and for defence 38 In some species a small pocket infrabuccal chamber inside the mouth stores food so it may be passed to other ants or their larvae 42 Mesosoma Both the legs and wings of the ant are attached to the mesosoma thorax The legs terminate in a hooked claw which allows them to hook on and climb surfaces 43 Only reproductive ants queens and males have wings Queens shed their wings after the nuptial flight leaving visible stubs a distinguishing feature of queens In a few species wingless queens ergatoids and males occur 38 Metasoma The metasoma the abdomen of the ant houses important internal organs including those of the reproductive respiratory tracheae and excretory systems Workers of many species have their egg laying structures modified into stings that are used for subduing prey and defending their nests 38 Polymorphism Seven leafcutter ant workers of various castes left and two queens right In the colonies of a few ant species there are physical castes workers in distinct size classes called minor median and major ergates Often the larger ants have disproportionately larger heads and correspondingly stronger mandibles These are known as macrergates while smaller workers are known as micrergates 44 Although formally known as dinergates such individuals are sometimes called soldier ants because their stronger mandibles make them more effective in fighting although they still are workers and their duties typically do not vary greatly from the minor or median workers In a few species the median workers are absent creating a sharp divide between the minors and majors 45 Weaver ants for example have a distinct bimodal size distribution 46 47 Some other species show continuous variation in the size of workers The smallest and largest workers in Carebara diversa show nearly a 500 fold difference in their dry weights 48 Workers cannot mate however because of the haplodiploid sex determination system in ants workers of a number of species can lay unfertilised eggs that become fully fertile haploid males The role of workers may change with their age and in some species such as honeypot ants young workers are fed until their gasters are distended and act as living food storage vessels These food storage workers are called repletes 49 For instance these replete workers develop in the North American honeypot ant Myrmecocystus mexicanus Usually the largest workers in the colony develop into repletes and if repletes are removed from the colony other workers become repletes demonstrating the flexibility of this particular polymorphism 50 This polymorphism in morphology and behaviour of workers initially was thought to be determined by environmental factors such as nutrition and hormones that led to different developmental paths however genetic differences between worker castes have been noted in Acromyrmex sp 51 These polymorphisms are caused by relatively small genetic changes differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens 52 The Australian jack jumper ant Myrmecia pilosula has only a single pair of chromosomes with the males having just one chromosome as they are haploid the lowest number known for any animal making it an interesting subject for studies in the genetics and developmental biology of social insects 53 54 Genome size Genome size is a fundamental characteristic of an organism Ants have been found to have tiny genomes with the evolution of genome size suggested to occur through loss and accumulation of non coding regions mainly transposable elements and occasionally by whole genome duplication 55 This may be related to colonisation processes but further studies are needed to verify this 55 Life cycle Meat eater ant nest during swarming The life of an ant starts from an egg if the egg is fertilised the progeny will be female diploid if not it will be male haploid Ants develop by complete metamorphosis with the larva stages passing through a pupal stage before emerging as an adult The larva is largely immobile and is fed and cared for by workers Food is given to the larvae by trophallaxis a process in which an ant regurgitates liquid food held in its crop This is also how adults share food stored in the social stomach Larvae especially in the later stages may also be provided solid food such as trophic eggs pieces of prey and seeds brought by workers 56 The larvae grow through a series of four or five moults and enter the pupal stage The pupa has the appendages free and not fused to the body as in a butterfly pupa 57 The differentiation into queens and workers which are both female and different castes of workers is influenced in some species by the nutrition the larvae obtain Genetic influences and the control of gene expression by the developmental environment are complex and the determination of caste continues to be a subject of research 58 Winged male ants called drones termed aner in old literature 59 emerge from pupae along with the usually winged breeding females Some species such as army ants have wingless queens Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development and so often are moved around among the various brood chambers within the colony 60 A new ergate spends the first few days of its adult life caring for the queen and young She then graduates to digging and other nest work and later to defending the nest and foraging These changes are sometimes fairly sudden and define what are called temporal castes An explanation for the sequence is suggested by the high casualties involved in foraging making it an acceptable risk only for ants who are older and are likely to die soon of natural causes 61 62 Ant colonies can be long lived The queens can live for up to 30 years and workers live from 1 to 3 years Males however are more transitory being quite short lived and surviving for only a few weeks 63 Ant queens are estimated to live 100 times as long as solitary insects of a similar size 64 Ants are active all year long in the tropics but in cooler regions they survive the winter in a state of dormancy known as hibernation The forms of inactivity are varied and some temperate species have larvae going into the inactive state diapause while in others the adults alone pass the winter in a state of reduced activity 65 Alate male ant Prenolepis imparis Reproduction Honey ants Prenolepis imparis mating A wide range of reproductive strategies have been noted in ant species Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis 66 Secretions from the male accessory glands in some species can plug the female genital opening and prevent females from re mating 67 Most ant species have a system in which only the queen and breeding females have the ability to mate Contrary to popular belief some ant nests have multiple queens while others may exist without queens Workers with the ability to reproduce are called gamergates and colonies that lack queens are then called gamergate colonies colonies with queens are said to be queen right 68 Drones can also mate with existing queens by entering a foreign colony such as in army ants When the drone is initially attacked by the workers it releases a mating pheromone If recognized as a mate it will be carried to the queen to mate 69 Males may also patrol the nest and fight others by grabbing them with their mandibles piercing their exoskeleton and then marking them with a pheromone The marked male is interpreted as an invader by worker ants and is killed 70 Most ants are univoltine producing a new generation each year 71 During the species specific breeding period winged females and winged males known to entomologists as alates leave the colony in what is called a nuptial flight The nuptial flight usually takes place in the late spring or early summer when the weather is hot and humid Heat makes flying easier and freshly fallen rain makes the ground softer for mated queens to dig nests 72 Males typically take flight before the females Males then use visual cues to find a common mating ground for example a landmark such as a pine tree to which other males in the area converge Males secrete a mating pheromone that females follow Males will mount females in the air but the actual mating process usually takes place on the ground Females of some species mate with just one male but in others they may mate with as many as ten or more different males storing the sperm in their spermathecae 73 In Cardiocondyla elegans workers may transport newly emerged queens to other conspecific nests where wingless males from unrelated colonies can mate with them a behavioural adaptation that may reduce the chances of inbreeding 74 Fertilised meat eater ant queen beginning to dig a new colony Mated females then seek a suitable place to begin a colony There they break off their wings using their tibial spurs and begin to lay and care for eggs The females can selectively fertilise future eggs with the sperm stored to produce diploid workers or lay unfertilized haploid eggs to produce drones The first workers to hatch known as nanitics 75 are weaker and smaller than later workers but they begin to serve the colony immediately They enlarge the nest forage for food and care for the other eggs Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site 73 a process akin to swarming in honeybees Behaviour and ecologyCommunication Two Camponotus sericeus workers communicating through touch and pheromones Ants communicate with each other using pheromones sounds and touch 76 Since most ants live on the ground they use the soil surface to leave pheromone trails that may be followed by other ants In species that forage in groups a forager that finds food marks a trail on the way back to the colony this trail is followed by other ants these ants then reinforce the trail when they head back with food to the colony When the food source is exhausted no new trails are marked by returning ants and the scent slowly dissipates This behaviour helps ants deal with changes in their environment For instance when an established path to a food source is blocked by an obstacle the foragers leave the path to explore new routes If an ant is successful it leaves a new trail marking the shortest route on its return Successful trails are followed by more ants reinforcing better routes and gradually identifying the best path 76 77 Ants use pheromones for more than just making trails A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from farther away Several ant species even use propaganda pheromones to confuse enemy ants and make them fight among themselves 78 Pheromones are produced by a wide range of structures including Dufour s glands poison glands and glands on the hindgut pygidium rectum sternum and hind tibia 64 Pheromones also are exchanged mixed with food and passed by trophallaxis transferring information within the colony 79 This allows other ants to detect what task group e g foraging or nest maintenance other colony members belong to 80 In ant species with queen castes when the dominant queen stops producing a specific pheromone workers begin to raise new queens in the colony 81 Some ants produce sounds by stridulation using the gaster segments and their mandibles Sounds may be used to communicate with colony members or with other species 82 83 Defence See also Defense in insects A Plectroctena sp attacks another of its kind to protect its territory Ants attack and defend themselves by biting and in many species by stinging often injecting or spraying chemicals Bullet ants Paraponera located in Central and South America are considered to have the most painful sting of any insect although it is usually not fatal to humans This sting is given the highest rating on the Schmidt sting pain index 84 The sting of jack jumper ants can be lethal for humans 85 and an antivenom has been developed for it 86 Fire ants Solenopsis spp are unique in having a venom sac containing piperidine alkaloids 87 Their stings are painful and can be dangerous to hypersensitive people 88 Formicine ants secrete a poison from their glands made mainly of formic acid 89 A weaver ant in fighting position mandibles wide open Trap jaw ants of the genus Odontomachus are equipped with mandibles called trap jaws which snap shut faster than any other predatory appendages within the animal kingdom 90 One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km h 78 and 143 mph with the jaws closing within 130 microseconds on average The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backward to escape a threat 90 Before striking the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory organs resembling hairs on the inside of the mandibles The mandibles also permit slow and fine movements for other tasks Trap jaws also are seen in other ponerines such as Anochetus as well as some genera in the tribe Attini such as Daceton Orectognathus and Strumigenys 90 91 which are viewed as examples of convergent evolution A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster If combat takes a turn for the worse a worker may perform a final act of suicidal altruism by rupturing the membrane of its gaster causing the content of its mandibular glands to burst from the anterior region of its head spraying a poisonous corrosive secretion containing acetophenones and other chemicals that immobilise small insect attackers The worker subsequently dies 92 Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening 93 Ant mound holes prevent water from entering the nest during rain In addition to defence against predators ants need to protect their colonies from pathogens Secretions from the metapleural gland unique to the ants produce a complex range of chemicals including several with antibiotic properties 94 Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophoresis the disposal of dead nest mates 95 Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana 96 while workers of Linepithema humile react to the absence of characteristic chemicals dolichodial and iridomyrmecin present on the cuticle of their living nestmates to trigger similar behaviour 97 Nests may be protected from physical threats such as flooding and overheating by elaborate nest architecture 98 99 Workers of Cataulacus muticus an arboreal species that lives in plant hollows respond to flooding by drinking water inside the nest and excreting it outside 100 Camponotus anderseni which nests in the cavities of wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration 101 Learning Two Weaver ants walking in tandem Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed A knowledgeable forager of Temnothorax albipennis can lead a naive nest mate to newly discovered food by the process of tandem running The follower obtains knowledge through its leading tutor The leader is acutely sensitive to the progress of the follower and slows down when the follower lags and speeds up when the follower gets too close 102 Controlled experiments with colonies of Cerapachys biroi suggest that an individual may choose nest roles based on her previous experience An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled One group was continually rewarded with prey while it was made certain that the other failed As a result members of the successful group intensified their foraging attempts while the unsuccessful group ventured out fewer and fewer times A month later the successful foragers continued in their role while the others had moved to specialise in brood care 103 Nest construction Main article Ant colony Leaf nest of weaver ants Pamalican Philippines Complex nests are built by many ant species but other species are nomadic and do not build permanent structures Ants may form subterranean nests or build them on trees These nests may be found in the ground under stones or logs inside logs hollow stems or even acorns The materials used for construction include soil and plant matter 73 and ants carefully select their nest sites Temnothorax albipennis will avoid sites with dead ants as these may indicate the presence of pests or disease They are quick to abandon established nests at the first sign of threats 104 The army ants of South America such as the Eciton burchellii species and the driver ants of Africa do not build permanent nests but instead alternate between nomadism and stages where the workers form a temporary nest bivouac from their own bodies by holding each other together 105 Weaver ant Oecophylla spp workers build nests in trees by attaching leaves together first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges Similar forms of nest construction are seen in some species of Polyrhachis 106 Ant bridge Formica polyctena among other ant species constructs nests that maintain a relatively constant interior temperature that aids in the development of larvae The ants maintain the nest temperature by choosing the location nest materials controlling ventilation and maintaining the heat from solar radiation worker activity and metabolism and in some moist nests microbial activity in the nest materials 107 108 Some ant species such as those that use natural cavities can be opportunistic and make use of the controlled micro climate provided inside human dwellings and other artificial structures to house their colonies and nest structures 109 110 Cultivation of food Main article Ant fungus mutualism Myrmecocystus honeypot ants store food to prevent colony famine Most ants are generalist predators scavengers and indirect herbivores 15 but a few have evolved specialised ways of obtaining nutrition It is believed that many ant species that engage in indirect herbivory rely on specialized symbiosis with their gut microbes 111 to upgrade the nutritional value of the food they collect 112 and allow them to survive in nitrogen poor regions such as rainforest canopies 113 Leafcutter ants Atta and Acromyrmex feed exclusively on a fungus that grows only within their colonies They continually collect leaves which are taken to the colony cut into tiny pieces and placed in fungal gardens Ergates specialise in related tasks according to their sizes The largest ants cut stalks smaller workers chew the leaves and the smallest tend the fungus Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material apparently detecting chemical signals from the fungus If a particular type of leaf is found to be toxic to the fungus the colony will no longer collect it The ants feed on structures produced by the fungi called gongylidia Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria introduced into the nest that may harm the fungi 114 Navigation An ant trail Foraging ants travel distances of up to 200 metres 700 ft from their nest 115 and scent trails allow them to find their way back even in the dark In hot and arid regions day foraging ants face death by desiccation so the ability to find the shortest route back to the nest reduces that risk Diurnal desert ants of the genus Cataglyphis such as the Sahara desert ant navigate by keeping track of direction as well as distance travelled Distances travelled are measured using an internal pedometer that keeps count of the steps taken 116 and also by evaluating the movement of objects in their visual field optical flow 117 Directions are measured using the position of the sun 118 They integrate this information to find the shortest route back to their nest 119 Like all ants they can also make use of visual landmarks when available 120 as well as olfactory and tactile cues to navigate 121 122 Some species of ant are able to use the Earth s magnetic field for navigation 123 The compound eyes of ants have specialised cells that detect polarised light from the Sun which is used to determine direction 124 125 These polarization detectors are sensitive in the ultraviolet region of the light spectrum 126 In some army ant species a group of foragers who become separated from the main column may sometimes turn back on themselves and form a circular ant mill The workers may then run around continuously until they die of exhaustion 127 Locomotion The female worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies Therefore unlike their wasp ancestors most ants travel by walking Some species are capable of leaping For example Jerdon s jumping ant Harpegnathos saltator is able to jump by synchronising the action of its mid and hind pairs of legs 128 There are several species of gliding ant including Cephalotes atratus this may be a common trait among arboreal ants with small colonies Ants with this ability are able to control their horizontal movement so as to catch tree trunks when they fall from atop the forest canopy 129 Other species of ants can form chains to bridge gaps over water underground or through spaces in vegetation Some species also form floating rafts that help them survive floods 130 These rafts may also have a role in allowing ants to colonise islands 131 Polyrhachis sokolova a species of ant found in Australian mangrove swamps can swim and live in underwater nests Since they lack gills they go to trapped pockets of air in the submerged nests to breathe 132 Cooperation and competition Meat eater ants feeding on a cicada social ants cooperate and collectively gather food Not all ants have the same kind of societies The Australian bulldog ants are among the biggest and most basal of ants Like virtually all ants they are eusocial but their social behaviour is poorly developed compared to other species Each individual hunts alone using her large eyes instead of chemical senses to find prey 133 Some species attack and take over neighbouring ant colonies Extreme specialists among these slave raiding ants such as the Amazon ants are incapable of feeding themselves and need captured workers to survive 134 Captured workers of enslaved Temnothorax species have evolved a counter strategy destroying just the female pupae of the slave making Temnothorax americanus but sparing the males who do not take part in slave raiding as adults 135 A worker Harpegnathos saltator a jumping ant engaged in battle with a rival colony s queen on top Ants identify kin and nestmates through their scent which comes from hydrocarbon laced secretions that coat their exoskeletons If an ant is separated from its original colony it will eventually lose the colony scent Any ant that enters a colony without a matching scent will be attacked 136 Parasitic ant species enter the colonies of host ants and establish themselves as social parasites species such as Strumigenys xenos are entirely parasitic and do not have workers but instead rely on the food gathered by their Strumigenys perplexa hosts 137 138 This form of parasitism is seen across many ant genera but the parasitic ant is usually a species that is closely related to its host A variety of methods are employed to enter the nest of the host ant A parasitic queen may enter the host nest before the first brood has hatched establishing herself prior to development of a colony scent Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest Some simply fight their way into the nest 139 A conflict between the sexes of a species is seen in some species of ants with these reproducers apparently competing to produce offspring that are as closely related to them as possible The most extreme form involves the production of clonal offspring An extreme of sexual conflict is seen in Wasmannia auropunctata where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process whereby a diploid egg loses its maternal contribution to produce haploid males who are clones of the father 140 Relationships with other organisms The spider Myrmarachne plataleoides female shown mimics weaver ants to avoid predators Ants form symbiotic associations with a range of species including other ant species other insects plants and fungi They also are preyed on by many animals and even certain fungi Some arthropod species spend part of their lives within ant nests either preying on ants their larvae and eggs consuming the food stores of the ants or avoiding predators These inquilines may bear a close resemblance to ants The nature of this ant mimicry myrmecomorphy varies with some cases involving Batesian mimicry where the mimic reduces the risk of predation Others show Wasmannian mimicry a form of mimicry seen only in inquilines 141 142 An ant collects honeydew from an aphid Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap The sugars in honeydew are a high energy food source which many ant species collect 143 In some cases the aphids secrete the honeydew in response to ants tapping them with their antennae The ants in turn keep predators away from the aphids and will move them from one feeding location to another When migrating to a new area many colonies will take the aphids with them to ensure a continued supply of honeydew Ants also tend mealybugs to harvest their honeydew Mealybugs may become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies 144 Myrmecophilous ant loving caterpillars of the butterfly family Lycaenidae e g blues coppers or hairstreaks are herded by the ants led to feeding areas in the daytime and brought inside the ants nest at night The caterpillars have a gland which secretes honeydew when the ants massage them Some caterpillars produce vibrations and sounds that are perceived by the ants 145 A similar adaptation can be seen in Grizzled skipper butterflies that emit vibrations by expanding their wings in order to communicate with ants which are natural predators of these butterflies 146 Other caterpillars have evolved from ant loving to ant eating these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae The caterpillar is then taken into the ant nest where it feeds on the ant larvae 147 A number of specialized bacteria have been found as endosymbionts in ant guts Some of the dominant bacteria belong to the order Hyphomicrobiales whose members are known for being nitrogen fixing symbionts in legumes but the species found in ant lack the ability to fix nitrogen 148 149 Fungus growing ants that make up the tribe Attini including leafcutter ants cultivate certain species of fungus in the genera Leucoagaricus or Leucocoprinus of the family Agaricaceae In this ant fungus mutualism both species depend on each other for survival The ant Allomerus decemarticulatus has evolved a three way association with the host plant Hirtella physophora Chrysobalanaceae and a sticky fungus which is used to trap their insect prey 150 Ants may obtain nectar from flowers such as the dandelion but are only rarely known to pollinate flowers Lemon ants make devil s gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees Duroia hirsuta This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees 151 Although some ants obtain nectar from flowers pollination by ants is somewhat rare one example being of the pollination of the orchid Leporella fimbriata which induces male Myrmecia urens to pseudocopulate with the flowers transferring pollen in the process 152 One theory that has been proposed for the rarity of pollination is that the secretions of the metapleural gland inactivate and reduce the viability of pollen 153 154 Some plants have special nectar exuding structures extrafloral nectaries that provide food for ants which in turn protect the plant from more damaging herbivorous insects 155 Species such as the bullhorn acacia Acacia cornigera in Central America have hollow thorns that house colonies of stinging ants Pseudomyrmex ferruginea who defend the tree against insects browsing mammals and epiphytic vines Isotopic labelling studies suggest that plants also obtain nitrogen from the ants 156 In return the ants obtain food from protein and lipid rich Beltian bodies In Fiji Philidris nagasau Dolichoderinae are known to selectively grow species of epiphytic Squamellaria Rubiaceae which produce large domatia inside which the ant colonies nest The ants plant the seeds and the domatia of young seedling are immediately occupied and the ant faeces in them contribute to rapid growth 157 Similar dispersal associations are found with other dolichoderines in the region as well 158 Another example of this type of ectosymbiosis comes from the Macaranga tree which has stems adapted to house colonies of Crematogaster ants 159 Many plant species have seeds that are adapted for dispersal by ants 160 Seed dispersal by ants or myrmecochory is widespread and new estimates suggest that nearly 9 of all plant species may have such ant associations 161 160 Often seed dispersing ants perform directed dispersal depositing the seeds in locations that increase the likelihood of seed survival to reproduction 162 Some plants in arid fire prone systems are particularly dependent on ants for their survival and dispersal as the seeds are transported to safety below the ground 163 Many ant dispersed seeds have special external structures elaiosomes that are sought after by ants as food 164 Ants can substantially alter rate of decomposition and nutrient cycling in their nest 165 166 By myrmecochory and modification of soil conditions they substantially alter vegetation and nutrient cycling in surrounding ecosystem 167 A convergence possibly a form of mimicry is seen in the eggs of stick insects They have an edible elaiosome like structure and are taken into the ant nest where the young hatch 168 A meat ant tending a common leafhopper nymph Most ants are predatory and some prey on and obtain food from other social insects including other ants Some species specialise in preying on termites Megaponera and Termitopone while a few Cerapachyinae prey on other ants 115 Some termites including Nasutitermes corniger form associations with certain ant species to keep away predatory ant species 169 The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant repellent chemical 170 It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants Other wasps such as A multipicta defend against ants by blasting them off the nest with bursts of wing buzzing 171 Stingless bees Trigona and Melipona use chemical defences against ants 115 Flies in the Old World genus Bengalia Calliphoridae prey on ants and are kleptoparasites snatching prey or brood from the mandibles of adult ants 172 Wingless and legless females of the Malaysian phorid fly Vestigipoda myrmolarvoidea live in the nests of ants of the genus Aenictus and are cared for by the ants 172 Oecophylla smaragdina killed by a fungus Fungi in the genera Cordyceps and Ophiocordyceps infect ants Ants react to their infection by climbing up plants and sinking their mandibles into plant tissue The fungus kills the ants grows on their remains and produces a fruiting body It appears that the fungus alters the behaviour of the ant to help disperse its spores 173 in a microhabitat that best suits the fungus 174 Strepsipteran parasites also manipulate their ant host to climb grass stems to help the parasite find mates 175 A nematode Myrmeconema neotropicum that infects canopy ants Cephalotes atratus causes the black coloured gasters of workers to turn red The parasite also alters the behaviour of the ant causing them to carry their gasters high The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten The droppings of the bird are collected by other ants and fed to their young leading to further spread of the nematode 176 Spiders Like this Menemerus jumping spider sometimes feed on ants A study of Temnothorax nylanderi colonies in Germany found that workers parasitized by the tapeworm Anomotaenia brevis ants are intermediate hosts the definitive hosts are woodpeckers lived much longer than unparasitized workers and had a reduced mortality rate comparable to that of the queens of the same species which live for as long as two decades 177 South American poison dart frogs in the genus Dendrobates feed mainly on ants and the toxins in their skin may come from the ants 178 Army ants forage in a wide roving column attacking any animals in that path that are unable to escape In Central and South America Eciton burchellii is the swarming ant most commonly attended by ant following birds such as antbirds and woodcreepers 179 180 This behaviour was once considered mutualistic but later studies found the birds to be parasitic Direct kleptoparasitism birds stealing food from the ants grasp is rare and has been noted in Inca doves which pick seeds at nest entrances as they are being transported by species of Pogonomyrmex 181 Birds that follow ants eat many prey insects and thus decrease the foraging success of ants 182 Birds indulge in a peculiar behaviour called anting that as yet is not fully understood Here birds rest on ant nests or pick and drop ants onto their wings and feathers this may be a means to remove ectoparasites from the birds Anteaters aardvarks pangolins echidnas and numbats have special adaptations for living on a diet of ants These adaptations include long sticky tongues to capture ants and strong claws to break into ant nests Brown bears Ursus arctos have been found to feed on ants About 12 16 and 4 of their faecal volume in spring summer and autumn respectively is composed of ants 183 Relationship with humans Weaver ants are used as a biological control for citrus cultivation in southern China Ants perform many ecological roles that are beneficial to humans including the suppression of pest populations and aeration of the soil The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control 184 On the other hand ants may become nuisances when they invade buildings or cause economic losses In some parts of the world mainly Africa and South America large ants especially army ants are used as surgical sutures The wound is pressed together and ants are applied along it The ant seizes the edges of the wound in its mandibles and locks in place The body is then cut off and the head and mandibles remain in place to close the wound 185 186 187 The large heads of the dinergates soldiers of the leafcutting ant Atta cephalotes are also used by native surgeons in closing wounds 188 Some ants have toxic venom and are of medical importance The species include Paraponera clavata tocandira and Dinoponera spp false tocandiras of South America 189 and the Myrmecia ants of Australia 190 In South Africa ants are used to help harvest the seeds of rooibos Aspalathus linearis a plant used to make a herbal tea The plant disperses its seeds widely making manual collection difficult Black ants collect and store these and other seeds in their nest where humans can gather them en masse Up to half a pound 200 g of seeds may be collected from one ant heap 191 192 Although most ants survive attempts by humans to eradicate them a few are highly endangered These tend to be island species that have evolved specialized traits and risk being displaced by introduced ant species Examples include the critically endangered Sri Lankan relict ant Aneuretus simoni and Adetomyrma venatrix of Madagascar 193 As food See also Entomophagy Roasted ants in Colombia Ant larvae for sale in Isaan Thailand Ants and their larvae are eaten in different parts of the world The eggs of two species of ants are used in Mexican escamoles They are considered a form of insect caviar and can sell for as much as US 50 per kg going up to US 200 per kg as of 2006 because they are seasonal and hard to find 194 In the Colombian department of Santander hormigas culonas roughly interpreted as large bottomed ants Atta laevigata are toasted alive and eaten 195 In areas of India and throughout Burma and Thailand a paste of the green weaver ant Oecophylla smaragdina is served as a condiment with curry 196 Weaver ant eggs and larvae as well as the ants may be used in a Thai salad yam Thai ya in a dish called yam khai mot daeng Thai yaikhmdaedng or red ant egg salad a dish that comes from the Issan or north eastern region of Thailand Saville Kent in the Naturalist in Australia wrote Beauty in the case of the green ant is more than skin deep Their attractive almost sweetmeat like translucency possibly invited the first essays at their consumption by the human species Mashed up in water after the manner of lemon squash these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland and is even appreciated by many European palates 197 In his First Summer in the Sierra John Muir notes that the Digger Indians of California ate the tickling acid gasters of the large jet black carpenter ants The Mexican Indians eat the repletes or living honey pots of the honey ant Myrmecocystus 197 As pests See also Ants of medical importance The tiny pharaoh ant is a major pest in hospitals and office blocks it can make nests between sheets of paper Some ant species are considered as pests primarily those that occur in human habitations where their presence is often problematic For example the presence of ants would be undesirable in sterile places such as hospitals or kitchens Some species or genera commonly categorized as pests include the Argentine ant immigrant pavement ant yellow crazy ant banded sugar ant pharaoh ant red wood ant black carpenter ant odorous house ant red imported fire ant and European fire ant Some ants will raid stored food some will seek water sources others may damage indoor structures some may damage agricultural crops directly or by aiding sucking pests Some will sting or bite 198 The adaptive nature of ant colonies make it nearly impossible to eliminate entire colonies and most pest management practices aim to control local populations and tend to be temporary solutions Ant populations are managed by a combination of approaches that make use of chemical biological and physical methods Chemical methods include the use of insecticidal bait which is gathered by ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis Management is based on the species and techniques may vary according to the location and circumstance 198 In science and technology See also Myrmecology Biomimetics and Ant colony optimization algorithms Camponotus nearcticus workers travelling between two formicaria through connector tubing Observed by humans since the dawn of history the behaviour of ants has been documented and the subject of early writings and fables passed from one century to another Those using scientific methods myrmecologists study ants in the laboratory and in their natural conditions Their complex and variable social structures have made ants ideal model organisms Ultraviolet vision was first discovered in ants by Sir John Lubbock in 1881 199 Studies on ants have tested hypotheses in ecology and sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies 200 Ant colonies may be studied by rearing or temporarily maintaining them in formicaria specially constructed glass framed enclosures 201 Individuals may be tracked for study by marking them with dots of colours 202 The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault tolerant systems for solving problems for example Ant colony optimization and Ant robotics This area of biomimetics has led to studies of ant locomotion search engines that make use of foraging trails fault tolerant storage and networking algorithms 203 As pets Main article Ant keeping From the late 1950s through the late 1970s ant farms were popular educational children s toys in the United States Some later commercial versions use transparent gel instead of soil allowing greater visibility at the cost of stressing the ants with unnatural light 204 In culture Aesop s ants illustration by Milo Winter 1888 1956 Anthropomorphised ants have often been used in fables and children s stories to represent industriousness and cooperative effort They also are mentioned in religious texts 205 206 In the Book of Proverbs in the Bible ants are held up as a good example of hard work and cooperation 207 Aesop did the same in his fable The Ant and the Grasshopper In the Quran Sulayman is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army Quran 27 18 208 209 In parts of Africa ants are considered to be the messengers of the deities Some Native American mythology such as the Hopi mythology considers ants as the very first animals Ant bites are often said to have curative properties The sting of some species of Pseudomyrmex is claimed to give fever relief 210 Ant bites are used in the initiation ceremonies of some Amazon Indian cultures as a test of endurance 211 212 In Greek mythology the goddess Athena turned the maiden Myrmex into an ant when the latter claimed to have invented the plough when in fact it was Athena s own invention 213 An ant pictured in the coat of arms of Multia a town in Finland Ant society has always fascinated humans and has been written about both humorously and seriously Mark Twain wrote about ants in his 1880 book A Tramp Abroad 214 Some modern authors have used the example of the ants to comment on the relationship between society and the individual Examples are Robert Frost in his poem Departmental and T H White in his fantasy novel The Once and Future King The plot in French entomologist and writer Bernard Werber s Les Fourmis science fiction trilogy is divided between the worlds of ants and humans ants and their behaviour is described using contemporary scientific knowledge H G Wells wrote about intelligent ants destroying human settlements in Brazil and threatening human civilization in his 1905 science fiction short story The Empire of the Ants In more recent times animated cartoons and 3 D animated films featuring ants have been produced including Antz A Bug s Life The Ant Bully The Ant and the Aardvark Ferdy the Ant and Atom Ant Renowned myrmecologist E O Wilson wrote a short story Trailhead in 2010 for The New Yorker magazine which describes the life and death of an ant queen and the rise and fall of her colony from an ants point of view 215 The French neuroanatomist psychiatrist and eugenicist Auguste Forel believed that ant societies were models for human society He published a five volume work from 1921 to 1923 that examined ant biology and society 216 In the early 1990s the video game SimAnt which simulated an ant colony won the 1992 Codie award for Best Simulation Program 217 Ants also are quite popular inspiration for many science fiction insectoids such as the Formics of Ender s Game the Bugs of Starship Troopers the giant ants in the films Them and Empire of the Ants Marvel Comics super hero Ant Man and ants mutated into super intelligence in Phase IV In computer strategy games ant based species often benefit from increased production rates due to their single minded focus such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II These characters are often credited with a hive mind a common misconception about ant colonies 218 See alsoMain article Outline of ants Ant venom Glossary of ant terms International Union for the Study 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January 2010 Trailhead The New Yorker pp 56 62 Parent A August 2003 Auguste Forel on ants and neurology The Canadian Journal of Neurological Sciences 30 3 284 91 doi 10 1017 s0317167100002754 PMID 12945958 1992 Excellence in Software Awards Winners Software amp Information Industry Association Archived from the original on 2009 06 11 Retrieved 3 April 2008 Sharkey AJC 2006 Robots insects and swarm intelligence Artificial Intelligence Review 26 4 255 268 doi 10 1007 s10462 007 9057 y S2CID 321326 Cited texts Borror DJ Triplehorn CA Delong DM 1989 Introduction to the Study of Insects 6th Edition Saunders College Publishing ISBN 978 0 03 025397 3 Holldobler B Wilson EO 1990 The Ants Harvard University Press ISBN 978 0 674 04075 5 Further readingMike Snider 8 Jan 2022 When the trees where these ants live were damaged they made some DIY home repairs USA Today Bolton Barry 1995 A New General Catalogue of the Ants of the World Harvard University Press ISBN 978 0 674 61514 4 Holldobler B Wilson EO 1998 Journey to the Ants A Story of Scientific Exploration Belknap Press ISBN 978 0 674 48526 6 Holldobler B Wilson EO 2009 The Superorganism The Beauty Elegance and Strangeness of Insect Societies Norton amp Co ISBN 978 0 393 06704 0 External links Wikiquote has quotations related to Ant Wikimedia Commons has media related to Formicidae Wikispecies has information related to Formicidae AntWeb from The California Academy of Sciences AntWiki Bringing Ants to the World Ant Species Fact Sheets from the National Pest Management Association on Argentine Carpenter Pharaoh Odorous and other ant species Ant Genera of the World distribution maps The super nettles A dermatologist s guide to ants in the plants Retrieved from https en wikipedia org w index php title Ant amp oldid 1153015424, wikipedia, wiki, book, books, library,

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