Ant

Page semi-protected
Source: Wikipedia, the free encyclopedia.

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

Myrmicinae

Ectatomminae

Heteroponerinae

Formicinae

Dolichoderinae

Aneuretinae

Pseudomyrmecinae

Myrmeciinae

Dorylinae‡

Poneroid

Ponerinae

Agroecomyrmecinae

Paraponerinae

Proceratiinae

Amblyoponinae

Apomyrminae

Leptanillinae

Martialinae

A

phylogeny of the extant ant subfamilies.[2][3][4]

*Cerapachyinae is
paraphyletic

‡ The previous dorylomorph subfamilies - Ecitoninae, Aenictinae, Aenictogitoninae, Cerapachyinae, Leptanilloidinae - were synonymized under
Dorylinae by Brady et al. in 2014[5]

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 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.

Ants gathering food (video)

Ants have colonised almost every landmass on

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

Proto-Germanic *ai-, "off, away" + *mait- "cut").[7][8]

The family name Formicidae is derived from the Latin formīca ("ant")[9] 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.[10][11]

Taxonomy and evolution

Aculeata

Chrysidoidea

 
 

Vespidae

Rhopalosomatidae

 
 

Pompilidae

Tiphiidae

 

Scolioidea

 

Apoidea

Formicidae

Phylogenetic position of the Formicidae[12]

The family Formicidae belongs to the order

Martialinae, are suggested to have diversified from early primitive ants that were likely to have been predators underneath the surface of the soil.[4][19]

Ants fossilised in Baltic amber

During the Cretaceous period, a few species of primitive ants ranged widely on 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).[16][20]

crickets, and other winged insects that do not undergo complete 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.[21] Velvet ants look like large ants, but are wingless female wasps.[22][23]

Distribution and diversity

Region Number of
species [24]
Neotropics 2,162
Nearctic 580
Europe 180
Africa 2,500
Asia 2,080
Melanesia 275
Australia 985
Polynesia 42

Ants have a

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.[28] 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).[29][30][31]

Ants range in size from 0.75 to 52 millimetres (0.030–2.0 in),

Morphology

Diagram of a worker ant (Neoponera verenae)

Ants are distinct in their

synapomorphic character.[39]

Like other arthropods, ants have an

ganglia and branches along the way reaching into the extremities of the appendages.[40]

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

blind.[3] 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.[43]

Two

mandibles, used to carry food, manipulate objects, construct nests, and for defence.[40] In some species, a small pocket (infrabuccal chamber) inside the mouth stores food, so it may be passed to other ants or their larvae.[44]

Mesosoma

Both the

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.[40]

Metasoma

The

stings that are used for subduing prey and defending their nests.[40]

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

bimodal size distribution.[48][49] 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.[50]

Workers cannot mate; however, because of the

haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.[55][56]

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.[57] This may be related to colonisation processes, but further studies are needed to verify this.[57]

Life cycle

Meat eater ant
nest during swarming

The life of an ant starts from an

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.[58]

The larvae grow through a series of four or five

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.[60] Winged male ants, called drones (termed "aner" in old literature[61]), 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.[62]

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. Such age-based task-specialization or polyethism has been suggested as having evolved due to the high casualties involved in foraging and defence, making it an acceptable risk only for ants who are older and likely to die sooner from natural causes.[63][64] In the Brazilian ant Forelius pusillus, the nest entrance is closed from the outside to protect the colony from predatory ant species at sunset each day. About one to eight workers seal the nest entrance from the outside and they have no chance of returning to the nest and are in effect sacrificed.[65] Whether these seemingly suicidal workers are older workers has not been determined.[66]

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.[67] Ant queens are estimated to live 100 times as long as solitary insects of a similar size.[68]

Ants are active all year long in the tropics; however, 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.[69]

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.[70] Secretions from the male accessory glands in some species can plug the female genital opening and prevent females from re-mating.[71] 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.[72]

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.[73] 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.[74]

Most ants are

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.[77] 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.[78]

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,

honeybees
.

Nests, colonies, and supercolonies

The typical ant species has a colony occupying a single nest, housing one or more queens, where the brood is raised. There are however more than 150 species of ants in 49 genera that are known to have colonies consisting of multiple spatially separated nests. These polydomous (as opposed to monodomous) colonies have food and workers moving between the nests.[80] Membership to a colony is identified by the response of worker ants which identify whether another individual belongs to their own colony or not. A signature cocktail of body surface chemicals (also known as cuticular hydrocarbons or CHCs) forms the so-called colony odor which other members can recognize.[81] Some ant species appear to be less discriminating and in the Argentine ant Linepithema humile, workers carried from a colony anywhere in the southern US and Mexico are acceptable within other colonies in the same region. Similarly workers from colonies that have established in Europe are accepted by any other colonies within Europe but not by the colonies in the Americas. The interpretation of these observations has been debated and some have been termed these large populations as supercolonies[82][83] while others have termed the populations as unicolonial.[84]

Behaviour and ecology

Communication

Two Camponotus sericeus workers communicating through touch and pheromones

Ants communicate with each other using pheromones, sounds, and touch.[85] 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.[85][86]

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.[87] 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.[68] Pheromones also are exchanged, mixed with food, and passed by trophallaxis, transferring information within the colony.[88] This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to.[89] In ant species with queen castes, when the dominant queen stops producing a specific pheromone, workers begin to raise new queens in the colony.[90]

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.[91][92]

Defence

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.[93]

The sting of jack jumper ants can be lethal for humans,[94] and an antivenom has been developed for it.[95] Fire ants, Solenopsis spp., are unique in having a venom sac containing piperidine alkaloids.[96] Their stings are painful and can be dangerous to hypersensitive people.[97] Formicine ants secrete a poison from their glands, made mainly of formic acid.[98]

mandibles
wide open

Trap-jaw ants of the genus

Attini, such as Daceton, Orectognathus, and Strumigenys,[99][100] which are viewed as examples of convergent evolution
.

anterior region of its head, spraying a poisonous, corrosive secretion containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies.[101]

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.[102] Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophoresis, the disposal of dead nest-mates.[103] Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana[104] 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.[105]

Nests may be protected from physical threats such as flooding and overheating by elaborate nest architecture.

Camponotus anderseni, which nests in the cavities of wood in mangrove habitats, deals with submergence under water by switching to anaerobic respiration.[109]

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.[110]

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.[111]

Nest construction

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,[77] 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.[112]

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.[113]

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.[114]

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.[115][116]

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.[117][118]

Cultivation of food

Myrmecocystus, honeypot ants, store food to prevent colony famine.

Most ants are generalist predators, scavengers, and indirect herbivores,

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.[122]

Navigation

An ant trail

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[124] and also by evaluating the movement of objects in their visual field (optical flow).[125] Directions are measured using the position of the sun.[126]
They integrate this information to find the shortest route back to their nest.[127] Like all ants, they can also make use of visual landmarks when available[128] as well as olfactory and tactile cues to navigate.[129][130] Some species of ant are able to use the Earth's magnetic field for navigation.[131] The compound eyes of ants have specialised cells that detect polarised light from the Sun, which is used to determine direction.[132][133] These polarization detectors are sensitive in the ultraviolet region of the light spectrum.[134] 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.[135]

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.[136] 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.[137]

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.[138] These rafts may also have a role in allowing ants to colonise islands.[139] 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.[140]

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.[141]

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.[142] 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).[143]

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.[144]

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.[145][146] 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.[147]

A

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.[148]

Relationships with other organisms

to avoid predators.

Ants form

Wasmannian mimicry, a form of mimicry seen only in inquilines.[149][150]

An ant collects honeydew from an aphid

plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect.[151] 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.[152]

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.[158]

dandelion
, but are only rarely known to pollinate flowers.

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.[165] Similar dispersal associations are found with other dolichoderines in the region as well.[166] Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.[167]

Many plant species have seeds that are adapted for dispersal by ants.[168] Seed dispersal by ants or myrmecochory is widespread, and new estimates suggest that nearly 9% of all plant species may have such ant associations.[169][168] Often, seed-dispersing ants perform directed dispersal, depositing the seeds in locations that increase the likelihood of seed survival to reproduction.[170] 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.[171] Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food.[172] Ants can substantially alter rate of decomposition and nutrient cycling in their nest.[173][174] By myrmecochory and modification of soil conditions they substantially alter vegetation and nutrient cycling in surrounding ecosystem.[175]

A

stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.[176]

A meat ant tending a common leafhopper nymph
Bold Jumping Spider (Phidippus audax) with a cutworm (tribe Noctuini) and then lost to ants (Family Formicidae)
Long-jawed orb weaver spiders
were consuming.

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.[123] Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away predatory ant species.[177] The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellent chemical.[178] 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.[179] Stingless bees (Trigona and Melipona) use chemical defences against ants.[123]

Flies in the Old World genus

kleptoparasites, snatching prey or brood from the mandibles of adult ants.[180] 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.[180]

Oecophylla smaragdina killed by a fungus

Fungi in the genera

fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores [181] in a microhabitat that best suits the fungus.[182] Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates.[183]

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.[184]

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.[185]

South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants.[186]

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.[191]

Relationship with humans

biological control
for citrus cultivation in southern China.

Ants perform many ecological roles that are beneficial to humans, including the suppression of

biological control.[192]
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.[193][194][195] The large heads of the dinergates (soldiers) of the leafcutting ant Atta cephalotes are also used by native surgeons in closing wounds.[196]

Some ants have toxic venom and are of medical importance. The species include Paraponera clavata (tocandira) and Dinoponera spp. (false tocandiras) of South America [197] and the Myrmecia ants of Australia.[198]

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.[199][200]

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.[201]

As food

Roasted ants in Colombia
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

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".[205]

In his First Summer in the Sierra,

As pests

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.[206] 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.[206]

In science and technology

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,

Ultraviolet vision was first discovered in ants by Sir John Lubbock in 1881.[207] 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.[208] Ant colonies may be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures.[209] Individuals may be tracked for study by marking them with dots of colours.[210]

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.[211]

As pets

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.[212]

In culture

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

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.[218] Ant bites are used in the initiation ceremonies of some Amazon Indian cultures as a test of endurance.[219][220] 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.[221]

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.

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. A similar German story involving army ants, Leiningen Versus the Ants, was written in 1937 and recreated in movie form as The Naked Jungle in 1954.[223] 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.[224] 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.[225]

In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".[226]

Ants also are quite popular inspiration for many

Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.[227]

See also

References

  1. ^
    S2CID 20729380
    .
  2. . Retrieved 11 February 2024.
  3. ^ (PDF) from the original on 9 October 2022.
  4. ^ .
  5. .
  6. ^ emmet. Merriam-Webster Dictionary
  7. ^ "ant". Merriam-Webster Online Dictionary. Retrieved 6 June 2008.
  8. ^ "Ant. Online Etymology Dictionary". Retrieved 30 May 2009.
  9. .
  10. ^ "Formic". Etymonline.com. Retrieved 30 January 2012.
  11. ^ "Pismire". Etymonline.com. Retrieved 27 August 2020.
  12. ^
    PMID 24094856
    .
  13. .
  14. .
  15. . Retrieved 16 March 2024.
  16. ^ .
  17. ^ .
  18. .
  19. S2CID 83598305. Archived from the original
    (PDF) on 23 April 2013. Retrieved 5 May 2013.
  20. ^ Hölldobler & Wilson (1990), pp. 23–24
  21. PMC 349550. Archived from the original
    (PDF) on 30 May 2010.
  22. ^ "Order Isoptera – Termites". Iowa State University Entomology. 16 February 2004. Archived from the original on 15 June 2008. Retrieved 12 June 2008.
  23. ^ "Family Mutillidae – Velvet ants". Iowa State University Entomology. 16 February 2004. Archived from the original on 30 June 2008. Retrieved 12 June 2008.
  24. ^ Hölldobler & Wilson (1990), p. 4
  25. ^ Jones, Alice S. "Fantastic ants – Did you know?". National Geographic Magazine. Archived from the original on 30 July 2008. Retrieved 5 July 2008.
  26. ^ Thomas, Philip (2007). "Pest Ants in Hawaii". Hawaiian Ecosystems at Risk project (HEAR). Retrieved 6 July 2008.
  27. PMID 36197959
    .
  28. .
  29. .
  30. ^ "How many ants are there for every one person on earth?". info.com. Archived from the original on 13 August 2013. Retrieved 27 July 2013.
  31. S2CID 252381912
    .
  32. ^ Hölldobler & Wilson (1990), p. 589
  33. .
  34. .
  35. ^ a b AntWeb
  36. ^ Agosti D, Majer JD, Alonso JE, Schultz TR, eds. (2000). Ants: Standard methods for measuring and monitoring biodiversity. Smithsonian Institution Press. Retrieved 13 December 2015.
  37. ^ Johnson NF (2007). "Hymenoptera name server". Ohio State University. Archived from the original on 27 January 2016. Retrieved 6 July 2008.
  38. ^ Borror, Triplehorn & Delong (1989), p. 737
  39. PMID 16630727
    .
  40. ^ a b c d Borror, Triplehorn & Delong (1989), pp. 24–71
  41. S2CID 33242108
    .
  42. .
  43. (PDF) from the original on 9 October 2022.
  44. .
  45. ^ Holbrook, Tate (22 September 2009). "Ask a Biologist: Face to Face with Ants". ASU School of Life Sciences. Retrieved 23 January 2018.
  46. .
  47. .
  48. (PDF) from the original on 9 October 2022.
  49. (PDF) from the original on 9 October 2022.
  50. (PDF) on 27 February 2008.
  51. .
  52. ^ Rissing SW (1984). "Replete caste production and allometry of workers in the Honey Ant, Myrmecocystus mexicanus Wesmael (Hymenoptera: Formicidae)". Journal of the Kansas Entomological Society. 57 (2): 347–350.
  53. PMID 12878720
    .
  54. .
  55. .
  56. .
  57. ^ .
  58. .
  59. .
  60. ^ Anderson KE, Linksvayer TA, Smith CR (2008). "The causes and consequences of genetic caste determination in ants (Hymenoptera: Formicidae)". Myrmecol. News. 11: 119–132.
  61. ISSN 1938-2901
    .
  62. ^ Hölldobler & Wilson (1990), pp. 351, 372
  63. .
  64. .
  65. .
  66. .
  67. .
  68. ^ .
  69. ^ Kipyatkov VE (2001). "Seasonal life cycles and the forms of dormancy in ants (Hymenoptera, Formicoidea)". Acta Societatis Zoologicae Bohemicae. 65 (2): 198–217.
  70. S2CID 21948488. Archived from the original
    (PDF) on 27 May 2011. Retrieved 16 April 2009.
  71. .
  72. .
  73. .
  74. .
  75. ^ Taylor RW (2007). "Bloody funny wasps! Speculations on the evolution of eusociality in ants" (PDF). In Snelling RR, Fisher BL, Ward PS (eds.). Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of contributions. Memoirs of the American Entomological Institute, 80. American Entomological Institute. pp. 580–609. Retrieved 13 December 2015.
  76. .
  77. ^ a b c Hölldobler & Wilson (1990), pp. 143–179
  78. PMID 33941829
    .
  79. .
  80. .
  81. .
  82. .
  83. .
  84. .
  85. ^ .
  86. (PDF) from the original on 9 October 2022.
  87. .
  88. .
  89. .
  90. ^ Hölldobler & Wilson (1990), p. 354
  91. PMID 11051518
    .
  92. .
  93. .
  94. .
  95. .
  96. .
  97. .
  98. .
  99. ^ .
  100. (PDF) from the original on 9 October 2022.
  101. .
  102. .
  103. .
  104. .
  105. .
  106. .
  107. .
  108. .
  109. .
  110. .
  111. .
  112. .
  113. ^ Hölldobler & Wilson (1990), p. 573
  114. .
  115. .
  116. .
  117. .
  118. .
  119. .
  120. .
  121. .
  122. (PDF) from the original on 9 October 2022.
  123. ^ .
  124. S2CID 15162376. Archived from the original
    (PDF) on 28 July 2011.
  125. S2CID 4625001. Archived from the original
    (PDF) on 27 July 2011. Retrieved 7 June 2011.
  126. S2CID 4571290. Archived from the original
    (PDF) on 7 July 2011. Retrieved 7 September 2010.
  127. .
  128. .
  129. .
  130. .
  131. .
  132. .
  133. .
  134. .
  135. .
  136. .
  137. S2CID 4368995. Archived from the original
    (PDF) on 16 June 2007.
  138. .
  139. .
  140. .
  141. .
  142. .
  143. , April 9, 2009
  144. .
  145. .
  146. .
  147. ^ Hölldobler & Wilson (1990), pp. 436–448
  148. S2CID 1188960
    .
  149. .
  150. (PDF) from the original on 9 October 2022.
  151. .
  152. .
  153. .
  154. ^ Elfferich NW (1998). "Is the larval and imaginal signalling of Lycaenidae and other Lepidoptera related to communication with ants". Deinsea. 4 (1).
  155. PMID 11729090
    .
  156. .
  157. .
  158. .
  159. .
  160. .
  161. .
  162. .
  163. .
  164. .
  165. .
  166. .
  167. .
  168. ^ .
  169. .
  170. .
  171. ^ Auld TD (1996). "Ecology of the Fabaceae in the Sydney region: fire, ants and the soil seedbank". Cunninghamia. 4 (22).
  172. S2CID 21974767
    .
  173. .
  174. ^ Frouz, J; Jílková, V (2008). "The effect of ants on soil properties and processes (Hymenoptera: Formicidae)". Myrmecological News. 1: 191–199.
  175. . Retrieved 12 July 2021.
  176. .
  177. .
  178. ^ Jeanne RL (1972). "Social biology of the neotropical wasp Mischocyttarus drewseni". Bull. Mus. Comp. Zool. 144: 63–150.
  179. ^ Jeanne R (July 1995). "Foraging in Social Wasps: Agelaia lacks recruitment to food (Hymenoptera: Vespidae)". Journal of the Kansas Entomological Society.
  180. ^ (PDF) from the original on 9 October 2022.
  181. ^ Schaechter E (2000). "Some weird and wonderful fungi". Microbiology Today. 27 (3): 116–117.
  182. S2CID 31283817
    .
  183. JSTOR 3494966. Archived from the original
    (PDF) on 9 March 2021. Retrieved 25 October 2017.
  184. .
  185. .
  186. .
  187. .
  188. ^ Vellely AC (2001). "Foraging at army ant swarms by fifty bird species in the highlands of Costa Rica" (PDF). Ornitologia Neotropical. 12: 271–275. Archived (PDF) from the original on 9 October 2022. Retrieved 8 June 2008.
  189. S2CID 85633598
    .
  190. .
  191. .
  192. ^ Hölldobler & Wilson (1990), pp. 619–629
  193. ^ Gottrup F, Leaper D (2004). "Wound healing: Historical aspects" (PDF). EWMA Journal. 4 (2): 5. Archived from the original (PDF) on 16 June 2007.
  194. .
  195. .
  196. .
  197. .
  198. .
  199. ^ Downes D, Laird SA (1999). "Innovative mechanisms for sharing benefits of biodiversity and related knowledge" (PDF). The Center for International Environmental Law. Archived (PDF) from the original on 23 April 2008. Retrieved 8 June 2008.
  200. S2CID 37728834
    .
  201. .
  202. .
  203. .
  204. ^ Bingham, C.T. (1903). Fauna of British India. Hymenoptera. Volume 2. London: Taylor and Francis. p. 311.
  205. ^ a b Bequaert J (1921). "Insects as food: How they have augmented the food supply of mankind in early and recent times". Natural History Journal. 21: 191–200.
  206. ^ a b "Pest Notes: Ants (Publication 7411)". University of California Agriculture and Natural Resources. 2007. Retrieved 5 June 2008.
  207. .
  208. .
  209. .
  210. .
  211. ^ Dicke E, Byde A, Cliff D, Layzell P (2004). Ispeert AJ, Murata M, Wakamiya N (eds.). "An ant-inspired technique for storage area network design". Proceedings of Biologically Inspired Approaches to Advanced Information Technology: First International Workshop, BioADIT 2004 LNCS 3141: 364–379.
  212. ^ US granted 5803014, Guri A, "Habitat media for ants and other invertebrates", issued 8 September 1998, assigned to Plant Cell Technology Inc 
  213. ^ "The Ant, The Ants". Quran. Vol. Surah 27. pp. 18–19. Archived from the original on 1 January 2007.
  214. ^ Bukhari S. "Beginning of Creation". Sunnah. Vol. 4 Book 54. Archived from the original on 18 August 2000.
  215. ^ See wikisource:Bible (World English)/Proverbs#30:25
  216. ^ Mentioned once in the Quran, Muhammad Asad translates the verse as following: till, when they came upon a valley [full] of ants, and an ant exclaimed: "O you ants! Get into your dwellings, lest Solomon and his hosts crush you without [even] being aware [of you]! (27:18)"
  217. ^ Deen MY (1990). "Islamic Environmental Ethics, Law, and Society" (PDF). In Engel JR, Engel JG (eds.). Ethics of Environment and Development. Bellhaven Press, London. Archived from the original (PDF) on 14 July 2011.
  218. S2CID 162813070
    .
  219. ^ Cesard N, Deturche J, Erikson P (2003). "Les Insectes dans les pratiques médicinales et rituelles d'Amazonie indigène". In Motte-Florac E, Thomas JM (eds.). Les insectes dans la tradition orale (in French). Peeters-Selaf, Paris. pp. 395–406.
  220. S2CID 73875767
    .
  221. Servius, Commentary on Virgil's Aeneid 4.402; Smith 1873, s.v. Myrmex
  222. . Retrieved 13 December 2015.
  223. .
  224. ^ Wilson, EO (25 January 2010). "Trailhead". The New Yorker. pp. 56–62.
  225. PMID 12945958
    .
  226. ^ "1992 Excellence in Software Awards Winners". Software & Information Industry Association. Archived from the original on 11 June 2009. Retrieved 3 April 2008.
  227. S2CID 321326
    .

Cited texts

Further reading

External links

This page is based on the copyrighted Wikipedia article: Ant . Articles is available under the CC BY-SA 3.0 license; additional terms may apply.Privacy Policy