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Formicidae

    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] Sphecomyrma was possibly a ground forager, while Haidomyrmex and Haidomyrmodes, related genera in subfamily Sphecomyrminae, are reconstructed as active arboreal predators.[13] Older ants in the genus Sphecomyrmodes have been found in 99 million year-old amber from Myanmar.[14][15] A 2006 study suggested that ants arose tens of millions of years earlier than previously thought, up to 168 million years ago.[1] After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.[16][1][17][18] 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][19]

An ant collects honeydew from an aphid.

Aphids and other hemipteran insects secrete a sweet liquid called 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.[134] 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.[135]

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.[136] 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.[137] Other ca

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.[136] 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.[137] 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.[138] A number of specialized bacteria have been found as endosymbionts in ant guts. Some of the dominant bacteria belong to the order Rhizobiales whose members are known for being nitrogen-fixing symbionts in legumes but the species found in ant lack the ability to fix nitrogen.[139][140] 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.[141]

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.[142] 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.[143] 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.[144][145] 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.[146] 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.[147] 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.[148] Similar dispersal associations are found with other dolichoderines in the region as well.[149] Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.[150]

Many plant species have seeds that are adapted for dispersal by ants.[151] Seed dispersal by ants or myrmecochory is widespread, and new estimates suggest that nearly 9% of all plant species may have such ant associations.[152][151] Often, seed-dispersing ants perform directed dispersal, depositing the seeds in locations that increase the likelihood of seed survival to reproduction.[153] Some plants in arid, fire-prone sys

Many plant species have seeds that are adapted for dispersal by ants.[151] Seed dispersal by ants or myrmecochory is widespread, and new estimates suggest that nearly 9% of all plant species may have such ant associations.[152][151] Often, seed-dispersing ants perform directed dispersal, depositing the seeds in locations that increase the likelihood of seed survival to reproduction.[153] 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.[154] Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food.[155]

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.[156]

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.[106] Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away predatory ant species.[157] The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellent chemical.[158] 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.[159] Stingless bees (Trigona and Melipona) use chemical defences against ants.[106]

Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants.[160] Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared

Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants.[160] 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.[160]

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 [161] in a microhabitat that best suits the fungus.[162] Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates.[163]

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.[164]

South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants.[165]

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.[166][167] This behaviour was once considered 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.[166][167] 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.[168] Birds that follow ants eat many prey insects and thus decrease the foraging success of ants.[169] 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.[170]

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.[171] 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.[172][173][174] The large heads of the dinergates (soldiers) of the leafcutting ant Atta cephalotes are also used by native surgeons in closing wounds.[175]

Some ants have toxic venom and are of 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.[172][173][174] The large heads of the dinergates (soldiers) of the leafcutting ant Atta cephalotes are also used by native surgeons in closing wounds.[175]