From HandWiki
Short description: Family of insects

Temporal range: 100–0 Ma[1]
Late Albian – Present
Fire ants 01.jpg
Fire ants
Scientific classification e
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Infraorder: Aculeata
Superfamily: Formicoidea
Family: Formicidae
Latreille, 1809
Type species
Formica rufa
Cladogram of






















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 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. File:Blackants-bredcrust-tokyo-may2015.webm

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.


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 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.[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 12 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]


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]


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]


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]


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]


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. 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.[61][62] 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.[63] Whether these seemingly suicidal workers are older workers has not been determined.[64]

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.[65] Ant queens are estimated to live 100 times as long as solitary insects of a similar size.[66]

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.[67]

Alate male ant, Prenolepis imparis


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.[68] Secretions from the male accessory glands in some species can plug the female genital opening and prevent females from re-mating.[69] 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.[70]

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.[71] 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.[72]

Most ants are univoltine, producing a new generation each year.[73] 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.[74] 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.[75] 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.[76]

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,[77] 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,[75] a process akin to swarming in 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.[78] 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.[79] 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[80][81] while others have termed the poulations as unicolonial.[82]

Behaviour and ecology


Two Camponotus sericeus workers communicating through touch and pheromones

Ants communicate with each other using pheromones, sounds, and touch.[83] 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.[83][84]

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.[85] 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.[66] Pheromones also are exchanged, mixed with food, and passed by trophallaxis, transferring information within the colony.[86] This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to.[87] In ant species with queen castes, when the dominant queen stops producing a specific pheromone, workers begin to raise new queens in the colony.[88]

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.[89][90]


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.[91]

The sting of jack jumper ants can be lethal for humans,[92] and an antivenom has been developed for it.[93] Fire ants, Solenopsis spp., are unique in having a venom sac containing piperidine alkaloids.[94] Their stings are painful and can be dangerous to hypersensitive people.[95] Formicine ants secrete a poison from their glands, made mainly of formic acid.[96]

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.[97] 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.[97] 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,[97][98] 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.[99]

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.[100] Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophoresis, the disposal of dead nest-mates.[101] Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana[102] 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.[103]

Nests may be protected from physical threats such as flooding and overheating by elaborate nest architecture.[104][105] 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.[106] Camponotus anderseni, which nests in the cavities of wood in mangrove habitats, deals with submergence under water by switching to anaerobic respiration.[107]


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.[108]

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.[109]

Nest construction

Main page: Biology: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,[75] 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.[110]

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.[111]

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.[112]

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.[113][114]

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.[115][116]

Cultivation of food

Main page: Biology: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[117] to upgrade the nutritional value of the food they collect[118] and allow them to survive in nitrogen poor regions, such as rainforest canopies.[119] 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.[120]


An ant trail

Foraging ants travel distances of up to 200 metres (700 ft) from their nest [121] 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[122] and also by evaluating the movement of objects in their visual field (optical flow).[123] Directions are measured using the position of the sun.[124] They integrate this information to find the shortest route back to their nest.[125] Like all ants, they can also make use of visual landmarks when available[126] as well as olfactory and tactile cues to navigate.[127][128] Some species of ant are able to use the Earth's magnetic field for navigation.[129] The compound eyes of ants have specialised cells that detect polarised light from the Sun, which is used to determine direction.[130][131] These polarization detectors are sensitive in the ultraviolet region of the light spectrum.[132] 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.[133]


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.[134] 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.[135]

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.[136] These rafts may also have a role in allowing ants to colonise islands.[137] Polyrhachis sokolova, a species of ant found in Australia n 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.[138]

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.[139]

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.[140] 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).[141]

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.[142]

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.[143][144] 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.[145]

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.[146]

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.[147][148]

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.[149] 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.[150]

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.[151] 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.[152] 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.[153] 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.[154][155] 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.[156]

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.[157] 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.[158] 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.[159][160] 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.[161] 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.[162] 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.[163] Similar dispersal associations are found with other dolichoderines in the region as well.[164] Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.[165]

Many plant species have seeds that are adapted for dispersal by ants.[166] Seed dispersal by ants or myrmecochory is widespread, and new estimates suggest that nearly 9% of all plant species may have such ant associations.[167][166] Often, seed-dispersing ants perform directed dispersal, depositing the seeds in locations that increase the likelihood of seed survival to reproduction.[168] 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.[169] Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food.[170] Ants can substantially alter rate of decomposition and nutrient cycling in their nest.[171][172] By myrmecochory and modification of soil conditions they substantially alter vegetation and nutrient cycling in surrounding ecosystem.[173]

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.[174]

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.[121] Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away predatory ant species.[175] The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellent chemical.[176] 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.[177] Stingless bees (Trigona and Melipona) use chemical defences against ants.[121]

Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants.[178] 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.[178]

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 [179] in a microhabitat that best suits the fungus.[180] Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates.[181]

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.[182]

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.[183]

South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants.[184]

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.[185][186] 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.[187] Birds that follow ants eat many prey insects and thus decrease the foraging success of ants.[188] 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.[189]

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.[190] 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.[191][192][193] The large heads of the dinergates (soldiers) of the leafcutting ant Atta cephalotes are also used by native surgeons in closing wounds.[194]

Some ants have toxic venom and are of medical importance. The species include Paraponera clavata (tocandira) and Dinoponera spp. (false tocandiras) of South America [195] and the Myrmecia ants of Australia.[196]

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.[197][198]

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.[199]

As food

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.[200] In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.[201] 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.[202] 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".[203]

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).[203]

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.[204] 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.[204]

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, 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.[205] 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.[206] Ant colonies may be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures.[207] Individuals may be tracked for study by marking them with dots of colours.[208]

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.[209]

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.[210]

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.[211][212] In the Book of Proverbs in the Bible, ants are held up as a good example of hard work and cooperation.[213] 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],[214][215] 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.[216] Ant bites are used in the initiation ceremonies of some Amazon Indian cultures as a test of endurance.[217][218] 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.[219]

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.[220] 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 are 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.[221] 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.[222]

In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".[223]

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.[224]

See also

Main page: Biology:Outline of ants


  1. 1.0 1.1 "Phylogeny of the ants: diversification in the age of angiosperms". Science 312 (5770): 101–104. April 2006. doi:10.1126/science.1124891. PMID 16601190. Bibcode2006Sci...312..101M. 
  2. 2.0 2.1 "Phylogeny, classification, and species-level taxonomy of ants (Hymenoptera: Formicidae)". Zootaxa 1668: 549–563. 2007. doi:10.11646/zootaxa.1668.1.26. 
  3. 3.0 3.1 "Newly discovered sister lineage sheds light on early ant evolution". Proceedings of the National Academy of Sciences of the United States of America 105 (39): 14913–14917. September 2008. doi:10.1073/pnas.0806187105. PMID 18794530. Bibcode2008PNAS..10514913R. 
  4. "The rise of army ants and their relatives: diversification of specialized predatory doryline ants". BMC Evolutionary Biology 14 (93): 93. May 2014. doi:10.1186/1471-2148-14-93. PMID 24886136. 
  5. emmet. Merriam-Webster Dictionary
  6. "ant". Merriam-Webster Online Dictionary. 
  7. "Ant. Online Etymology Dictionary". 
  8. Simpson DP (1979). Cassell's Latin Dictionary (5th ed.). London: Cassell. ISBN 978-0-304-52257-6. 
  9. "Formic". 
  10. "Pismire". 
  11. 11.0 11.1 "Phylogenomics resolves evolutionary relationships among ants, bees, and wasps". Current Biology 23 (20): 2058–2062. October 2013. doi:10.1016/j.cub.2013.08.050. PMID 24094856. 
  12. "The first mesozoic ants". Science 157 (3792): 1038–1040. September 1967. doi:10.1126/science.157.3792.1038. PMID 17770424. Bibcode1967Sci...157.1038W. 
  13. Boudinot, Brendon E; Richter, Adrian; Katzke, Julian; Chaul, Júlio C M; Keller, Roberto A; Economo, Evan P; Beutel, Rolf Georg; Yamamoto, Shûhei (2022-07-29). "Evidence for the evolution of eusociality in stem ants and a systematic revision of † Gerontoformica (Hymenoptera: Formicidae)" (in en). Zoological Journal of the Linnean Society 195 (4): 1355–1389. doi:10.1093/zoolinnean/zlab097. ISSN 0024-4082. 
  14. 14.0 14.1 "A formicine in New Jersey cretaceous amber (Hymenoptera: formicidae) and early evolution of the ants". Proceedings of the National Academy of Sciences of the United States of America 97 (25): 13678–13683. December 2000. doi:10.1073/pnas.240452097. PMID 11078527. Bibcode2000PNAS...9713678G. 
  15. 15.0 15.1 "The rise of the ants: a phylogenetic and ecological explanation". Proceedings of the National Academy of Sciences of the United States of America 102 (21): 7411–7414. May 2005. doi:10.1073/pnas.0502264102. PMID 15899976. Bibcode2005PNAS..102.7411W. 
  16. "Ants and the fossil record". Annual Review of Entomology 58: 609–730. 2013. doi:10.1146/annurev-ento-120710-100600. PMID 23317048. 
  17. "Rediscovery of the bizarre Cretaceous ant Haidomyrmex Dlussky (Hymenoptera: Formicidae), with two new species". American Museum Novitates (3755): 1–16. 2012. doi:10.1206/3755.2. Retrieved 2013-05-05. 
  18. Hölldobler & Wilson (1990), pp. 23–24
  19. "Evolution of eusociality in termites". Annu. Rev. Ecol. Syst. 28 (5): 27–53. 1997. doi:10.1146/annurev.ecolsys.28.1.27. PMC 349550. 
  20. "Order Isoptera – Termites". Iowa State University Entomology. 16 February 2004. 
  21. "Family Mutillidae – Velvet ants". Iowa State University Entomology. 16 February 2004. 
  22. Hölldobler & Wilson (1990), p. 4
  23. Jones, Alice S. "Fantastic ants – Did you know?". National Geographic Magazine. 
  24. Thomas, Philip (2007). "Pest Ants in Hawaii". Hawaiian Ecosystems at Risk project (HEAR). 
  25. Fayle, Tom M.; Klimes, Petr (2022-10-18). "Improving estimates of global ant biomass and abundance" (in en). Proceedings of the National Academy of Sciences 119 (42): e2214825119. doi:10.1073/pnas.2214825119. ISSN 0027-8424. PMID 36197959. Bibcode2022PNAS..11914825F. 
  26. The Superorganism: The Beauty, Elegance, and Strangeness of Insect Societies. New York: W.W. Norton. 2009. p. 5. ISBN 978-0-393-06704-0. 
  27. "In search of ant ancestors". Proceedings of the National Academy of Sciences of the United States of America 97 (26): 14028–14029. December 2000. doi:10.1073/pnas.011513798. PMID 11106367. Bibcode2000PNAS...9714028S. 
  28. "How many ants are there for every one person on earth?". 
  29. Schultheiss, Patrick; Nooten, Sabine S.; Wang, Runxi; Wong, Mark K. L.; Brassard, François; Guénard, Benoit (2022-10-04). "The abundance, biomass, and distribution of ants on Earth" (in en). Proceedings of the National Academy of Sciences 119 (40): e2201550119. doi:10.1073/pnas.2201550119. ISSN 0027-8424. PMID 36122199. Bibcode2022PNAS..11901550S. 
  30. Hölldobler & Wilson (1990), p. 589
  31. Shattuck SO (1999). Australian ants: their biology and identification. Collingwood, Vic: CSIRO. p. 149. ISBN 978-0-643-06659-5. 
  32. Schaal, Stephan (27 January 2006). "Messel". Encyclopedia of Life Sciences. doi:10.1038/npg.els.0004143. ISBN 978-0-470-01617-6. 
  33. 33.0 33.1 AntWeb
  34. Ants: Standard methods for measuring and monitoring biodiversity. Smithsonian Institution Press. 2000. Retrieved 2015-12-13. 
  35. "Hymenoptera name server". Ohio State University. 2007. 
  36. Borror, Triplehorn & Delong (1989), p. 737
  37. Ouellette, Gary D.; Fisher, Brian L.; Girman, Derek J. (2006). "Molecular systematics of basal subfamilies of ants using 28S rRNA (Hymenoptera: Formicidae)" (in en). Molecular Phylogenetics and Evolution 40 (2): 359–369. doi:10.1016/j.ympev.2006.03.017. ISSN 1055-7903. PMID 16630727. 
  38. 38.0 38.1 38.2 38.3 Borror, Triplehorn & Delong (1989), pp. 24–71
  39. "Oceili: a celestial compass in the desert ant cataglyphis". Science 228 (4696): 192–194. April 1985. doi:10.1126/science.228.4696.192. PMID 17779641. Bibcode1985Sci...228..192F. 
  40. "Miniaturisation decreases visual navigational competence in ants". The Journal of Experimental Biology 221 (Pt 7): jeb177238. April 2018. doi:10.1242/jeb.177238. PMID 29487158. 
  41. "Attack behaviour and distance perception in the Australian bulldog ant Myrmecia nigriceps". Journal of Experimental Biology 119 (1): 115–131. 1985. doi:10.1242/jeb.119.1.115. 
  42. "The infrabuccal pocket of a formicine ant: a social filtration device". Psyche: A Journal of Entomology 69 (3): 107–116. 1962. doi:10.1155/1962/25068. 
  43. Holbrook, Tate (22 September 2009). "Ask a Biologist: Face to Face with Ants". ASU School of Life Sciences. 
  44. Singh, Rajendra (2006). Elements of Entomology. Rastogi Publications. p. 284. ISBN 978-8171336777. 
  45. "The origin and evolution of polymorphism in ants". The Quarterly Review of Biology 28 (2): 136–156. June 1953. doi:10.1086/399512. PMID 13074471. 
  46. Weber, NA (1946). "Dimorphism in the African Oecophylla worker and an anomaly (Hym.: Formicidae)". Annals of the Entomological Society of America 39: 7–10. doi:10.1093/aesa/39.1.7. 
  47. Wilson, Edward O.; Taylor, Robert W. (1964). "A Fossil Ant Colony: New Evidence of Social Antiquity". Psyche: A Journal of Entomology 71 (2): 93–103. doi:10.1155/1964/17612. 
  48. "Physical castes in ant workers: a problem for Daceton armigerum and other ants". Psyche: A Journal of Entomology 98 (4): 283–292. 1991. doi:10.1155/1991/30265. 
  49. Børgesen LW (2000). "Nutritional function of replete workers in the pharaoh's ant, Monomorium pharaonis (L.)". Insectes Sociaux 47 (2): 141–146. doi:10.1007/PL00001692. 
  50. Rissing, Steven W (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. 
  51. "Worker caste polymorphism has a genetic basis in Acromyrmex leaf-cutting ants". Proceedings of the National Academy of Sciences of the United States of America 100 (16): 9394–9397. August 2003. doi:10.1073/pnas.1633701100. PMID 12878720. Bibcode2003PNAS..100.9394H. 
  52. "Alternative genetic foundations for a key social polymorphism in fire ants". Genetics 165 (4): 1853–1867. December 2003. doi:10.1093/genetics/165.4.1853. PMID 14704171. 
  53. "Myrmecia pilosula, an ant with only one Pair of chromosomes". Science 231 (4743): 1278. March 1986. doi:10.1126/science.231.4743.1278. PMID 17839565. Bibcode1986Sci...231.1278C. 
  54. "The evolution of genome size in ants". BMC Evolutionary Biology 8 (64): 64. February 2008. doi:10.1186/1471-2148-8-64. PMID 18302783. 
  55. 55.0 55.1 Moura, Mariana Neves; Cardoso, Danon Clemes; Cristiano, Maykon Passos (2021). "The tight genome size of ants: diversity and evolution under ancestral state reconstruction and base composition". Zoological Journal of the Linnean Society 193 (1): 124–144. doi:10.1093/zoolinnean/zlaa135. ISSN 0024-4082. 
  56. The Ants. Harvard University Press. 1990. p. 291. ISBN 978-0-674-04075-5. 
  57. Gillott, Cedric (1995). Entomology. Springer. p. 325. ISBN 978-0-306-44967-3. 
  58. Anderson, Kirk E.; Linksvayer, Timothy A.; Smith, Chris R. (2008). "The causes and consequences of genetic caste determination in ants (Hymenoptera: Formicidae)". Myrmecol. News 11: 119–132. 
  59. "A Glossary of Terms and Phrases Used in the Study of Social Insects" (in en). Annals of the Entomological Society of America 44 (3): 473–484. 1951. doi:10.1093/aesa/44.3.473. ISSN 1938-2901. 
  60. Hölldobler & Wilson (1990), pp. 351, 372
  61. Traniello JFA (1989). "Foraging strategies of ants". Annual Review of Entomology 34: 191–210. doi:10.1146/annurev.en.34.010189.001203. 
  62. "Behavioral flexibility of temporal sub-castes in the fire ant, Solenopsis invicta, in response to food". Psyche: A Journal of Entomology 91 (3–4): 319–332. 1984. doi:10.1155/1984/39236. 
  63. Tofilski, Adam; Couvillon, Margaret J.; Evison, Sophie E. F.; Helanterä, Heikki; Robinson, Elva J. H.; Ratnieks, Francis L. W. (2008). "Preemptive Defensive Self-Sacrifice by Ant Workers" (in en). The American Naturalist 172 (5): E239–E243. doi:10.1086/591688. ISSN 0003-0147. PMID 18928332. 
  64. Shorter, J. R.; Rueppell, O. (2012). "A review on self-destructive defense behaviors in social insects" (in en). Insectes Sociaux 59 (1): 1–10. doi:10.1007/s00040-011-0210-x. ISSN 0020-1812. 
  65. Keller L (1998). "Queen lifespan and colony characteristics in ants and termites". Insectes Sociaux 45 (3): 235–246. doi:10.1007/s000400050084. 
  66. 66.0 66.1 Encyclopedia of Insects. San Diego: Academic Press. 2003. pp. 29–32. ISBN 978-0-12-586990-4. 
  67. Kipyatkov VE (2001). "Seasonal life cycles and the forms of dormancy in ants (Hymenoptera, Formicoidea)". Acta Societatis Zoologicae Bohemicae 65 (2): 198–217. 
  68. "Ant reproductive strategies". Res. Popul. Ecol. 37 (2): 135–149. 1995. doi:10.1007/BF02515814. Retrieved 2009-04-16. 
  69. "Evidence for mating plugs in the fire ant Solenopsis invicta". Insectes Sociaux 50 (4): 401–402. 2003. doi:10.1007/s00040-003-0697-x. 
  70. "Reproductive cooperation between queens and their mated workers: the complex life history of an ant with a valuable nest". Proceedings of the National Academy of Sciences of the United States of America 92 (24): 10977–10979. November 1995. doi:10.1073/pnas.92.24.10977. PMID 11607589. Bibcode1995PNAS...9210977P. 
  71. "Sexual competition during colony reproduction in army ants". Biological Journal of the Linnean Society 30 (3): 229–243. 1987. doi:10.1111/j.1095-8312.1987.tb00298.x. 
  72. "Pheromonal manipulation of workers by a fighting male to kill his rival males in the ant Cardiocondyla wroughtonii". Naturwissenschaften 79 (6): 274–276. 1992. doi:10.1007/BF01175395. Bibcode1992NW.....79..274Y. 
  73. "Bloody funny wasps! Speculations on the evolution of eusociality in ants". 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. 2007. pp. 580–609. Retrieved 2015-12-13. 
  74. "The organization of a nuptial flight of the ant Pheidole sitarches Wheeler". Psyche: A Journal of Entomology 64 (2): 46–50. 1957. doi:10.1155/1957/68319. 
  75. 75.0 75.1 75.2 Hölldobler & Wilson (1990), pp. 143–179
  76. "Worker ants promote outbreeding by transporting young queens to alien nests". Communications Biology 4 (1): 515. May 2021. doi:10.1038/s42003-021-02016-1. PMID 33941829. 
  77. The Behavioural Ecology of Ants. Springer Science & Business Media. 2013. p. 41. ISBN 978-9400931237. 
  78. Cook, Zoe; Franks, Daniel W.; Robinson, Elva J.H. (2013). "Exploration versus exploitation in polydomous ant colonies" (in en). Journal of Theoretical Biology 323: 49–56. doi:10.1016/j.jtbi.2013.01.022. PMID 23380232. Bibcode2013JThBi.323...49C. 
  79. Bos, Nick; d’Ettorre, Patrizia (2012). "Recognition of Social Identity in Ants". Frontiers in Psychology 3: 83. doi:10.3389/fpsyg.2012.00083. ISSN 1664-1078. PMID 22461777. 
  80. Moffett, Mark W. (2012). "Supercolonies of billions in an invasive ant: What is a society?" (in en). Behavioral Ecology 23 (5): 925–933. doi:10.1093/beheco/ars043. ISSN 1465-7279. 
  81. Van Wilgenburg, Ellen; Torres, Candice W.; Tsutsui, Neil D. (2010). "The global expansion of a single ant supercolony" (in en). Evolutionary Applications 3 (2): 136–143. doi:10.1111/j.1752-4571.2009.00114.x. ISSN 1752-4571. PMID 25567914. 
  82. Helanterä, Heikki; Strassmann, Joan E.; Carrillo, Juli; Queller, David C. (2009). "Unicolonial ants: where do they come from, what are they and where are they going?" (in en). Trends in Ecology & Evolution 24 (6): 341–349. doi:10.1016/j.tree.2009.01.013. PMID 19328589. 
  83. 83.0 83.1 "Communication in ants". Current Biology 16 (15): R570–R574. August 2006. doi:10.1016/j.cub.2006.07.015. PMID 16890508. 
  84. "Self-organized shortcuts in the Argentine ant". Naturwissenschaften 76 (12): 579–581. 1989. doi:10.1007/BF00462870. Bibcode1989NW.....76..579G. 
  85. "Sociobiology of slave-making ants". Acta Ethologica 3 (2): 67–82. 2001. doi:10.1007/s102110100038. 
  86. Information processing in social insects. Birkhäuser. 1999. pp. 224–227. ISBN 978-3-7643-5792-4. 
  87. "Structural complexity of chemical recognition cues affects the perception of group membership in the ants Linephithema humile and Aphaenogaster cockerelli". The Journal of Experimental Biology 210 (Pt 5): 897–905. March 2007. doi:10.1242/jeb.02706. PMID 17297148. 
  88. Hölldobler & Wilson (1990), p. 354
  89. "Analysis of acoustic communication by ants". The Journal of the Acoustical Society of America 108 (4): 1920–1929. October 2000. doi:10.1121/1.1290515. PMID 11051518. Bibcode2000ASAJ..108.1920H. 
  90. "Use of stridulation in foraging leaf-cutting ants: Mechanical support during cutting or short-range recruitment signal?". Behavioral Ecology and Sociobiology 39 (5): 293–299. 1996. doi:10.1007/s002650050292. 
  91. "Hemolytic activities of stinging insect venoms". Archives of Insect Biochemistry and Physiology 1 (2): 155–160. 1983. doi:10.1002/arch.940010205. 
  92. "The natural history of sensitivity to jack jumper ants (Hymenoptera formicidae Myrmecia pilosula) in Tasmania". The Medical Journal of Australia 145 (11–12): 564–566. 1986. doi:10.5694/j.1326-5377.1986.tb139498.x. PMID 3796365. 
  93. "Efficacy of ant venom immunotherapy and whole body extracts". The Journal of Allergy and Clinical Immunology 116 (2): 464–465; author reply 465–466. August 2005. doi:10.1016/j.jaci.2005.04.025. PMID 16083810. 
  94. "Gaster flagging by fire ants (Solenopsis spp.): Functional significance of venom dispersal behavior". Journal of Chemical Ecology 11 (12): 1757–1768. December 1985. doi:10.1007/BF01012125. PMID 24311339. 
  95. "Hypersensitivity to fire ant venom". Annals of Allergy, Asthma & Immunology 77 (2): 87–95; quiz 96–99. August 1996. doi:10.1016/S1081-1206(10)63493-X. PMID 8760773. 
  96. "Hexadecanol and hexadecyl formate in the venom gland of formicine ants". Philosophical Transactions of the Royal Society B 341 (1296): 177–180. 1993. doi:10.1098/rstb.1993.0101. 
  97. 97.0 97.1 97.2 "Multifunctionality and mechanical origins: ballistic jaw propulsion in trap-jaw ants". Proceedings of the National Academy of Sciences of the United States of America 103 (34): 12787–12792. August 2006. doi:10.1073/pnas.0604290103. PMID 16924120. Bibcode2006PNAS..10312787P. 
  98. "The trap-jaw mechanism in the dacetine ants Daceton armigerum and Strumigenys sp.". The Journal of Experimental Biology 199 (Pt 9): 2021–2033. 1996. doi:10.1242/jeb.199.9.2021. PMID 9319931. 
  99. "The chemistry of exploding ants, Camponotus spp. (cylindricus complex)". Journal of Chemical Ecology 30 (8): 1479–1492. August 2004. doi:10.1023/B:JOEC.0000042063.01424.28. PMID 15537154. 
  100. Yek, Sze Huei; Mueller, Ulrich G. (2011). "The metapleural gland of ants" (in en). Biological Reviews 86 (4): 774–791. doi:10.1111/j.1469-185X.2010.00170.x. PMID 21504532. 
  101. "Undertaking specialization in the desert leaf-cutter ant Acromyrmex versicolor". Animal Behaviour 58 (2): 437–442. August 1999. doi:10.1006/anbe.1999.1184. PMID 10458895. 
  102. "Antennal olfactory sensitivity in response to task-related odours of three castes of the ant Atta mexicana (hymenoptera: formicidae)". Physiological Entomology 31 (4): 353–360. 2006. doi:10.1111/j.1365-3032.2006.00526.x. 
  103. "Chemical signals associated with life inhibit necrophoresis in Argentine ants". Proceedings of the National Academy of Sciences of the United States of America 106 (20): 8251–8255. May 2009. doi:10.1073/pnas.0901270106. PMID 19416815. Bibcode2009PNAS..106.8251C. 
  104. "The nest architecture of the Florida harvester ant, Pogonomyrmex badius". Journal of Insect Science 4 (21): 21. 2004. doi:10.1093/jis/4.1.21. PMID 15861237. 
  105. ""Wall-papering" and elaborate nest architecture in the ponerine ant Harpegnathos saltator". Insectes Sociaux 41 (2): 211–218. 1994. doi:10.1007/BF01240479. 
  106. "Communal peeing: a new mode of flood control in ants". Die Naturwissenschaften 87 (12): 563–565. December 2000. doi:10.1007/s001140050780. PMID 11198200. Bibcode2000NW.....87..563M. 
  107. "The mangrove ant, Camponotus anderseni, switches to anaerobic respiration in response to elevated CO2 levels". Journal of Insect Physiology 53 (5): 505–508. May 2007. doi:10.1016/j.jinsphys.2007.02.002. PMID 17382956. 
  108. "Teaching in tandem-running ants". Nature 439 (7073): 153. January 2006. doi:10.1038/439153a. PMID 16407943. Bibcode2006Natur.439..153F. 
  109. "Individual experience alone can generate lasting division of labor in ants". Current Biology 17 (15): 1308–1312. August 2007. doi:10.1016/j.cub.2007.06.047. PMID 17629482. 
  110. "Tomb evaders: house-hunting hygiene in ants". Biology Letters 1 (2): 190–192. June 2005. doi:10.1098/rsbl.2005.0302. PMID 17148163. 
  111. Hölldobler & Wilson (1990), p. 573
  112. "Evolution of nest-weaving behaviour in arboreal nesting ants of the genus Polyrhachis Fr. Smith (Hymenoptera: Formicidae)". Australian Journal of Entomology 44 (2): 164–169. 2005. doi:10.1111/j.1440-6055.2005.00462.x. 
  113. "The Effect of Nest Moisture on Daily Temperature Regime in the Nests of Formica polyctena Wood Ants". Insectes Sociaux 47 (3): 229–235. 2000. doi:10.1007/PL00001708. 
  114. Kadochová, Štěpánka; Frouz, Jan (2013). "Thermoregulation strategies in ants in comparison to other social insects, with a focus on red wood ants ( Formica rufa group)". F1000Research 2: 280. doi:10.12688/f1000research.2-280.v2. ISSN 2046-1402. PMID 24715967. 
  115. "Impact of human dwellings on the distribution of the exotic Argentine ant: a case study in the Doñana National Park, Spain". Biological Conservation 115 (2): 279–289. 2004. doi:10.1016/S0006-3207(03)00147-2. 
  116. "Nest-site limitation and nesting resources of ants (Hymenoptera: Formicidae) in urban green spaces". Environmental Entomology 38 (3): 600–607. June 2009. doi:10.1603/022.038.0311. PMID 19508768. 
  117. "Highly similar microbial communities are shared among related and trophically similar ant species". Molecular Ecology 21 (9): 2282–2296. May 2012. doi:10.1111/j.1365-294x.2011.05464.x. PMID 22276952. 
  118. "Nutritional upgrading for omnivorous carpenter ants by the endosymbiont Blochmannia". BMC Biology 5: 48. October 2007. doi:10.1186/1741-7007-5-48. PMID 17971224. 
  119. "Bacterial gut symbionts are tightly linked with the evolution of herbivory in ants". Proceedings of the National Academy of Sciences of the United States of America 106 (50): 21236–21241. December 2009. doi:10.1073/pnas.0907926106. PMID 19948964. Bibcode2009PNAS..10621236R. 
  120. Schultz TR (1999). "Ants, plants and antibiotics". Nature 398 (6730): 747–748. doi:10.1038/19619. Bibcode1999Natur.398..747S. 
  121. 121.0 121.1 121.2 "Ecology of foraging by ants". Annual Review of Ecology and Systematics 4: 231–257. 1973. doi:10.1146/ 
  122. "The ant odometer: stepping on stilts and stumps". Science 312 (5782): 1965–1967. June 2006. doi:10.1126/science.1126912. PMID 16809544. Bibcode2006Sci...312.1965W. 
  123. "Desert ants Cataglyphis fortis use self-induced optic flow to measure distances travelled". Journal of Comparative Physiology A 177. 1995. doi:10.1007/BF00243395. Retrieved 2011-06-07. 
  124. "Desert ant navigation: how miniature brains solve complex tasks". Journal of Comparative Physiology A 189 (8): 579–588. August 2003. doi:10.1007/s00359-003-0431-1. PMID 12879352. Retrieved 2010-09-07. 
  125. "The ant's estimation of distance travelled: experiments with desert ants, Cataglyphis fortis". Journal of Comparative Physiology A 190 (1): 1–6. January 2004. doi:10.1007/s00359-003-0465-4. PMID 14614570. 
  126. "Visual navigation in desert ants Cataglyphis fortis: are snapshots coupled to a celestial system of reference?". Journal of Experimental Biology 205 (14): 1971–1978. 2002. doi:10.1242/jeb.205.14.1971. PMID 12089203. 
  127. "Smells like home: Desert ants, Cataglyphis fortis, use olfactory landmarks to pinpoint the nest". Frontiers in Zoology 6: 5. February 2009. doi:10.1186/1742-9994-6-5. PMID 19250516. 
  128. "Visual and tactile learning of ground structures in desert ants". The Journal of Experimental Biology 209 (Pt 17): 3336–3344. September 2006. doi:10.1242/jeb.02364. PMID 16916970. 
  129. "Orientation by magnetic field in leaf-cutter ants, Atta colombica (Hymenoptera: Formicidae)". Ethology 109 (10): 835–846. 2003. doi:10.1046/j.0179-1613.2003.00927.x. 
  130. "Homing in wood ants, Formica japonica: use of the skyline panorama". The Journal of Experimental Biology 204 (Pt 12): 2063–2072. June 2001. doi:10.1242/jeb.204.12.2063. PMID 11441048. 
  131. "Homing in the ant Cataglyphis bicolor". Science 164 (3876): 192–194. April 1969. doi:10.1126/science.164.3876.192. PMID 5774195. Bibcode1969Sci...164..192W. 
  132. Chapman, Reginald Frederick (1998). The Insects: Structure and Function (4th ed.). Cambridge University Press. pp. 600. ISBN 978-0-521-57890-5. 
  133. "Army ants trapped by their evolutionary history". PLOS Biology 1 (2): E37. November 2003. doi:10.1371/journal.pbio.0000037. PMID 14624241. 
  134. "A novel mechanism for jumping in the Indian ant Harpegnathos saltator (Jerdon) (Formicidae, Ponerinae)". Experientia 50: 63–71. 1994. doi:10.1007/BF01992052. 
  135. "Directed aerial descent in canopy ants". Nature 433 (7026): 624–626. February 2005. doi:10.1038/nature03254. PMID 15703745. Bibcode2005Natur.433..624Y. 
  136. "Fire ants self-assemble into waterproof rafts to survive floods". Proceedings of the National Academy of Sciences of the United States of America 108 (19): 7669–7673. May 2011. doi:10.1073/pnas.1016658108. PMID 21518911. Bibcode2011PNAS..108.7669M. 
  137. Morrison LW (1998). "A review of Bahamian ant (Hymenoptera: Formicidae) biogeography". Journal of Biogeography 25 (3): 561–571. doi:10.1046/j.1365-2699.1998.2530561.x. 
  138. "Ant fauna of a mangrove community in the Australian seasonal tropics, with particular reference to zonation". Australian Journal of Zoology 44 (5): 521–533. 1996. doi:10.1071/ZO9960521. 
  139. "Aspects of the biology of the primitive ant genus Myrmecia F. (Hymenoptera: Formicidae)". Australian Journal of Entomology 27 (4): 305–309. 1988. doi:10.1111/j.1440-6055.1988.tb01179.x. 
  140. "Ant and termite mound coinhabitants in the wetlands of Santo Antonio da Patrulha, Rio Grande do Sul, Brazil". Brazilian Journal of Biology 65 (3): 431–437. August 2005. doi:10.1590/S1519-69842005000300008. PMID 16341421. 
  141. "First evidence for slave rebellion: enslaved ant workers systematically kill the brood of their social parasite protomognathus americanus". Evolution; International Journal of Organic Evolution 63 (4): 1068–1075. April 2009. doi:10.1111/j.1558-5646.2009.00591.x. PMID 19243573.  See also New Scientist, April 9, 2009
  142. "Internest aggression and identification of possible nestmate discrimination pheromones in polygynous antFormica montana". Journal of Chemical Ecology 16 (7): 2217–2228. July 1990. doi:10.1007/BF01026932. PMID 24264088. 
  143. Ward PS (1996). "A new workerless social parasite in the ant genus Pseudomyrmex (Hymenoptera: Formicidae), with a discussion of the origin of social parasitism in ants". Systematic Entomology 21 (3): 253–263. doi:10.1046/j.1365-3113.1996.d01-12.x. 
  144. Taylor RW (1968). "The Australian workerless inquiline ant, Strumigenys xenos Brown (Hymenoptera-Formicidae) recorded from New Zealand". New Zealand Entomologist 4 (1): 47–49. doi:10.1080/00779962.1968.9722888. 
  145. Hölldobler & Wilson (1990), pp. 436–448
  146. "Clonal reproduction by males and females in the little fire ant". Nature 435 (7046): 1230–1234. June 2005. doi:10.1038/nature03705. PMID 15988525. Bibcode2005Natur.435.1230F. 
  147. Reiskind J (1977). "Ant-mimicry in Panamanian clubionid and salticid spiders (Araneae: Clubionidae, Salticidae)". Biotropica 9 (1): 1–8. doi:10.2307/2387854. 
  148. Cushing PE (1997). "Myrmecomorphy and myrmecophily in spiders: A Review". The Florida Entomologist 80 (2): 165–193. doi:10.2307/3495552. 
  149. "Ecological consequences of interactions between ants and honeydew-producing insects". Proceedings. Biological Sciences 274 (1607): 151–164. January 2007. doi:10.1098/rspb.2006.3701. PMID 17148245. 
  150. "Effects of Pheidole megacephala (Hymenoptera: Formicidae) on survival and dispersal of Dysmicoccus neobrevipes (Homoptera: Pseudococcidae)". Journal of Economic Entomology 89 (5): 1124–1129. 1996. doi:10.1093/jee/89.5.1124. 
  151. DeVries PJ (1992). "Singing caterpillars, ants and symbiosis". Scientific American 267 (4): 76–82. doi:10.1038/scientificamerican1092-76. Bibcode1992SciAm.267d..76D. 
  152. Elfferich, Nico W. (1998). "Is the larval and imaginal signalling of Lycaenidae and other Lepidoptera related to communication with ants". Deinsea 4 (1). 
  153. "The ecology and evolution of ant association in the Lycaenidae (Lepidoptera)". Annual Review of Entomology 47: 733–771. 2002. doi:10.1146/annurev.ento.47.091201.145257. PMID 11729090. 
  154. "Surveying the microbiome of ants: comparing 454 pyrosequencing with traditional methods to uncover bacterial diversity". Applied and Environmental Microbiology 79 (2): 525–534. January 2013. doi:10.1128/AEM.03107-12. PMID 23124239. Bibcode2013ApEnM..79..525K. 
  155. "The genome of Rhizobiales bacteria in predatory ants reveals urease gene functions but no genes for nitrogen fixation" (in En). Scientific Reports 6 (1): 39197. December 2016. doi:10.1038/srep39197. PMID 27976703. Bibcode2016NatSR...639197N. 
  156. "Insect behaviour: arboreal ants build traps to capture prey". Nature 434 (7036): 973. April 2005. doi:10.1038/434973a. PMID 15846335. Bibcode2005Natur.434..973D. 
  157. "The devil to pay: a cost of mutualism with Myrmelachista schumanni ants in 'devil's gardens' is increased herbivory on Duroia hirsuta trees". Proceedings. Biological Sciences 274 (1613): 1117–23. April 2007. doi:10.1098/rspb.2006.0415. PMID 17301016. 
  158. "Pseudocopulation of an orchid by male ants: a test of two hypotheses accounting for the rarity of ant pollination". Oecologia 73 (4): 522–524. October 1987. doi:10.1007/BF00379410. PMID 28311968. Bibcode1987Oecol..73..522P. 
  159. Beattie, Andrew J.; Turnbull, Christine; R. B., Knox; E. G., Williams (1984). "Ant Inhibition of Pollen Function: A Possible Reason Why Ant Pollination is Rare". American Journal of Botany 71 (3): 421–426. doi:10.2307/2443499. 
  160. New, Tim R. (2017). "Classic Themes: Ants, Plants and Fungi" (in en). Mutualisms and Insect Conservation. Springer International Publishing. pp. 63–103. doi:10.1007/978-3-319-58292-4_4. ISBN 9783319582917. 
  161. "Role of extrafloral nectaries of Vicia faba in attraction of ants and herbivore exclusion by ants". Entomological Science 7 (2): 119–124. 2004. doi:10.1111/j.1479-8298.2004.00057.x. 
  162. "Do ants feed plants? A 15N labelling study of nitrogen fluxes from ants to plants in the mutualism of Pheidole and Piper". Journal of Ecology 91: 126–134. 2003. doi:10.1046/j.1365-2745.2003.00747.x. 
  163. "Obligate plant farming by a specialized ant". Nature Plants 2 (12): 16181. November 2016. doi:10.1038/nplants.2016.181. PMID 27869787. 
  164. "The assembly of ant-farmed gardens: mutualism specialization following host broadening". Proceedings. Biological Sciences 284 (1850): 20161759. March 2017. doi:10.1098/rspb.2016.1759. PMID 28298344. 
  165. "Studies of a South East Asian ant-plant association: protection of Macaranga trees by Crematogaster borneensis". Oecologia 79 (4): 463–470. June 1989. doi:10.1007/bf00378662. PMID 28313479. Bibcode1989Oecol..79..463F. 
  166. 166.0 166.1 Lengyel, Szabolcs; Gove, Aaron D.; Latimer, Andrew M.; Majer, Jonathan D.; Dunn, Robert R. (2010). "Convergent evolution of seed dispersal by ants, and phylogeny and biogeography in flowering plants: A global survey". Perspectives in Plant Ecology, Evolution and Systematics 12: 43–55. doi:10.1016/j.ppees.2009.08.001. 
  167. Giladi I (2006). "Choosing benefits or partners: a review of the evidence for the evolution of myrmecochory". Oikos 112 (3): 481–492. doi:10.1111/j.0030-1299.2006.14258.x. 
  168. "Directed dispersal: demographic analysis of an ant-seed mutualism". American Naturalist 131 (1): 1–13. 1988. doi:10.1086/284769. 
  169. "Ecology of the Fabaceae in the Sydney region: fire, ants and the soil seedbank". Cunninghamia 4 (22). 1996. 
  170. "The fate of Corydalis cava elaiosomes within an ant colony of Myrmica rubra: elaiosomes are preferentially fed to larvae". Insectes Sociaux 52 (1): 55–62. 2005. doi:10.1007/s00040-004-0773-x. 
  171. Frouz, Jan (1997). "The effect of wood ants (Formica polyctena Foerst) on the transformation of phosphorus in a spruce plantation". Pedobiologia 41: 437–447. 
  172. Frouz, J; Jílková, V (2008). "The effect of ants on soil properties and processes (Hymenoptera: Formicidae)". Myrmecological News 1: 191–199. 
  173. Frouz, Jan; Jílková, Veronika; Sorvari, Jouni (2016). "Contribution of wood ants to nutrient cycling and ecosystem function". in Robinson, Elva J. H.; Stockan, Jenni A.. Wood Ant Ecology and Conservation. Ecology, Biodiversity and Conservation. Cambridge: Cambridge University Press. pp. 207–220. ISBN 978-1-107-04833-1. Retrieved 2021-07-12. 
  174. "Capitula on stick insect eggs and elaiosomes on seeds: convergent adaptations for burial by ants". Functional Ecology 6 (6): 642–648. 1992. doi:10.2307/2389958. 
  175. "Behavioural Interactions Between Crematogaster brevispinosa rochai Forel (Hymenoptera: Formicidae) and Two Nasutitermes Species (Isoptera: Termitidae)". Journal of Insect Behavior 18 (1): 1–17. 2005. doi:10.1007/s10905-005-9343-y. 
  176. "Social biology of the neotropical wasp Mischocyttarus drewseni". Bull. Mus. Comp. Zool. 144: 63–150. 1972. 
  177. Jeanne, Robert (July 1995). "Foraging in Social Wasps: Agelaia lacks recruitment to food (Hymenoptera: Vespidae)". Journal of the Kansas Entomological Society. 
  178. 178.0 178.1 "Kleptoparasitism and phoresy in the Diptera". Florida Entomologist 82 (2): 179–197. 1999. doi:10.2307/3496570. 
  179. Schaechter E (2000). "Some weird and wonderful fungi". Microbiology Today 27 (3): 116–117. 
  180. "The life of a dead ant: the expression of an adaptive extended phenotype". The American Naturalist 174 (3): 424–433. September 2009. doi:10.1086/603640. PMID 19627240. 
  181. Wojcik DP (1989). "Behavioral interactions between ants and their parasites" (PDF). The Florida Entomologist 72 (1): 43–451. doi:10.2307/3494966. Retrieved 2017-10-25. 
  182. "Myrmeconema neotropicum n. g., n. sp., a new tetradonematid nematode parasitising South American populations of Cephalotes atratus (Hymenoptera: Formicidae), with the discovery of an apparent parasite-induced host morph". Systematic Parasitology 69 (2): 145–153. 2008. doi:10.1007/s11230-007-9125-3. PMID 18038201. 
  183. "Parasite Presence Induces Gene Expression Changes in an Ant Host Related to Immunity and Longevity". Genes 12 (1): 202118. January 2021. doi:10.1098/rsos.202118. PMID 34017599. 
  184. Caldwell JP (1996). "The evolution of myrmecophagy and its correlates in poison frogs (Family Dendrobatidae)". Journal of Zoology 240 (1): 75–101. doi:10.1111/j.1469-7998.1996.tb05487.x. 
  185. "Birds and Army Ants". Annual Review of Ecology and Systematics 9: 243–263. 1978. doi:10.1146/ 
  186. Vellely AC (2001). "Foraging at army ant swarms by fifty bird species in the highlands of Costa Rica". Ornitologia Neotropical 12: 271–275. Retrieved 8 June 2008. 
  187. Inzunza, Ernesto Ruelas; Martínez-Leyva, J. Eduardo; Valenzuela-González, Jorge E. (2015). "Doves kleptoparasitize ants". The Southwestern Naturalist 60 (1): 103–106. doi:10.1894/msh-03.1. 
  188. Wrege PH; Wikelski, Martin; Mandel, James T.; Rassweiler, Thomas; Couzin, Iain D. (2005). "Antbirds parasitize foraging army ants". Ecology 86 (3): 555–559. doi:10.1890/04-1133. 
  189. "Bears and ants: myrmecophagy by brown bears in central Scandinavia". Canadian Journal of Zoology 77 (4): 551–561. 1999. doi:10.1139/z99-004. 
  190. Hölldobler & Wilson (1990), pp. 619–629
  191. "Wound healing: Historical aspects". EWMA Journal 4 (2): 5. 2004. 
  192. Gudger EW (1925). "Stitching wounds with the mandibles of ants and beetles". Journal of the American Medical Association 84 (24): 1861–1864. doi:10.1001/jama.1925.02660500069048. 
  193. Sapolsky, Robert M. (2001). A Primate's Memoir: A Neuroscientist's Unconventional Life Among the Baboons. Simon and Schuster. pp. 156. ISBN 978-0-7432-0241-1. 
  194. Wheeler, William M. (1910). Ants: Their Structure, Development and Behavior. Columbia University Biological Series. 9. Columbia University Press. pp. 10. doi:10.5962/bhl.title.1937. ISBN 978-0-231-00121-2. OCLC 560205. 
  195. "Description of an injury in a human caused by a false tocandira (Dinoponera gigantea, Perty, 1833) with a revision on folkloric, pharmacological and clinical aspects of the giant ants of the genera Paraponera and Dinoponera (sub-family Ponerinae)". Revista do Instituto de Medicina Tropical de Sao Paulo 47 (4): 235–238. 2005. doi:10.1590/S0036-46652005000400012. PMID 16138209. 
  196. "Ant sting mortality in Australia". Toxicon 40 (8): 1095–1100. August 2002. doi:10.1016/S0041-0101(02)00097-1. PMID 12165310. 
  197. "Innovative mechanisms for sharing benefits of biodiversity and related knowledge". The Center for International Environmental Law. 1999. 
  198. "Rooibos tea, a South African contribution to world beverages". Economic Botany 17 (3): 186–194. 1963. doi:10.1007/BF02859435. 
  199. "The influence of sociality on the conservation biology of social insects". Ecology Letters 4 (6): 650–662. 2001. doi:10.1046/j.1461-0248.2001.00253.x. 
  200. Cruz-Labana, J. D.; Tarango-Arámbula, L. A.; Alcántara-Carbajal, J. L.; Pimentel-López, J.; Ugalde-Lezama, S.; Ramírez-Valverde, G.; Méndez-Gallegos, S. J. (2014). "Habitat use by the "Escamolera" ant (Liometopum apiculatum Mayr) in central Mexico" (in en). Agrociencia 48 (6): 569–582. ISSN 1405-3195. 
  201. "Insects as food: why the western attitude is important". Annual Review of Entomology 44: 21–50. 1999. doi:10.1146/annurev.ento.44.1.21. PMID 9990715. 
  202. Bingham CT (1903). Fauna of British India. Hymenoptera Volume 3. p. 311. 
  203. 203.0 203.1 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. 
  204. 204.0 204.1 "Pest Notes: Ants (Publication 7411)". University of California Agriculture and Natural Resources. 2007. 
  205. "Observations on ants, bees, and wasps. IX. Color of flowers as an attraction to bees: Experiments and considerations thereon". J. Linn. Soc. Lond. (Zool.) 16 (90): 110–112. 1881. doi:10.1111/j.1096-3642.1882.tb02275.x. 
  206. Mutualism: Ants and their insect partners. Cambridge University Press. 2008. ISBN 978-0-521-86035-2. 
  207. Kennedy CH (1951). "Myrmecological technique. IV. Collecting ants by rearing pupae". The Ohio Journal of Science 51 (1): 17–20. 
  208. "An improved and quantified technique for marking individual fire ants (Hymenoptera: Formicidae)". The Florida Entomologist 83 (1): 74–78. 2000. doi:10.2307/3496231. 
  209. "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. 2004. 
  210. Guri A, "Habitat media for ants and other invertebrates", US patent granted 5803014, issued 8 September 1998, assigned to Plant Cell Technology Inc
  211. "The Ant, The Ants". Quran. Surah 27. pp. 18–19. 
  212. Bukhari, Sahih. "Beginning of Creation". Sunnah. 4 Book 54. 
  213. See wikisource:Bible (World English)/Proverbs#30:25
  214. Mentioned once in theTemplate:Quran ref, 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)"
  215. Deen, Mawil Y. Izzi (1990). "Islamic Environmental Ethics, Law, and Society". Ethics of Environment and Development. Bellhaven Press, London. 
  216. Balee WL (2000). "Antiquity of traditional ethnobiological knowledge in Amazonia: The Tupi-Guarani family and time". Ethnohistory 47 (2): 399–422. doi:10.1215/00141801-47-2-399. 
  217. "Les Insectes dans les pratiques médicinales et rituelles d'Amazonie indigène" (in fr). Les insectes dans la tradition orale. Peeters-Selaf, Paris. 2003. pp. 395–406. 
  218. "The super-nettles. A dermatologist's guide to ants in the plants". International Journal of Dermatology 24 (4): 204–210. May 1985. doi:10.1111/j.1365-4362.1985.tb05760.x. PMID 3891647. 
  219. Servius, Commentary on Virgil's Aeneid 4.402; Smith 1873, s.v. Myrmex
  220. Twain, Mark (1880). "22 The Black Forest and Its Treasures". A Tramp Abroad. New York: Oxford University Press. ISBN 978-0-19-510137-9. Retrieved 2015-12-13. 
  221. Wilson, EO (25 January 2010). "Trailhead". The New Yorker: 56–62. 
  222. "Auguste Forel on ants and neurology". The Canadian Journal of Neurological Sciences 30 (3): 284–291. August 2003. doi:10.1017/s0317167100002754. PMID 12945958. 
  223. "1992 Excellence in Software Awards Winners". Software & Information Industry Association. 
  224. Sharkey AJC (2006). "Robots, insects and swarm intelligence". Artificial Intelligence Review 26 (4): 255–268. doi:10.1007/s10462-007-9057-y. 

Cited texts

Further reading

External links

Wikidata ☰ Q7386 entry