Temporal range: 130–0 Ma
Cretaceous - Recent
Meat eater ant feeding on honey
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Suborder: Apocrita
Superfamily: Vespoidea
Family: Formicidae
Latreille, 1809

Ants are social insects of the family Formicidae (play /fɔrˈmɪsɨd/) and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than 12,500 out of an estimated total of 22,000 species have been classified.[3][4] They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.

Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.[5]

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 most ecosystems, and may form 15–25% of the terrestrial animal biomass.[6] 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.[7]

Ant societies have division of labour, communication between individuals, and an ability to solve complex problems.[8] 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 rituals. Some species are valued in their role as biological pest control agents.[9] However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, establishing themselves in areas where they are accidentally introduced.[10]



The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German āmeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut").[11][12] The family name Formicidae is derived from the Latin formīca ("ant")[13] 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 hypothetized that a Proto-Indo-European word *morwi- was used, cf. Sanskrit vamrah, Latin formīca, Greek myrmex, Old Church Slavonic mraviji, Old Irish moirb, Old Norse maurr.[14]

Taxonomy and evolution

Ants fossilised in Baltic amber.










Phylogenetic position of the Formicidae.[15]

The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.[16][17][18] In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps.[19] Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators underneath the surface of the soil.[2]

During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. 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, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene).[16][20]

Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution.[21] Velvet ants look like large ants, but are wingless female wasps.[22][23]

Distribution and diversity

Region Number of
species [24]
Neotropics 2162
Nearctic 580
Europe 180
Africa 2500
Asia 2080
Melanesia 275
Australia 985
Polynesia 42

Ants 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.[25][26] Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15-20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.[6]

Ants range in size from 0.75 to 52 millimetres (0.030–2.0 in),[27][28] the largest species being the fossil Titanomyrma giganteum, the queen of which was 6 centimetres (2.4 in) long with a wingspan of 15 centimetres (5.9 in).[29] Ants vary in colour; most ants are red or black, but a few species are green and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates 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 AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species.[30] The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.[31][32]


Ants are distinct in their morphology from other insects in having 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. 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 can be formed by one or two nodes (the second alone, or the second and third abdominal segments).[33]

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

Like other insects, 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 like carbon dioxide pass through their exoskeleton through 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 towards 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.[34]

Diagram of a worker ant (Pachycondyla verenae).

An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization.[35] Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also 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.[34] In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.[36]

All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.[34]

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.[34]


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 workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still 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.[37] Weaver ants, for example, have a distinct bimodal size distribution.[38][39] Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights.[40] 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.[41] This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp.[42] 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.[43] The Australian jack jumper ant (Myrmecia pilosula) has only a single pair of chromosomes (males have 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.[44][45]

Development and reproduction

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 larval 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", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa.[46] The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), 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.[47] Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony.[48]

A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.[49][50]

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

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 can 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.[51] The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year.[52] During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males 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. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males.[7] Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak 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. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site,[53] a process akin to swarming in honeybees.

Ants 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[54] and one species, Mycocepurus smithii is known to be all-female.[55]

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, and survive only a few weeks.[56] Ant queens are estimated to live 100 times longer than solitary insects of a similar size.[57]

Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. 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.[58]

Behaviour and ecology


Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps).

Ants communicate with each other using pheromones.[59] These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can 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 finding the best path.[60]

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 further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves.[61] 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.[57] Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony.[62] This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to.[63] In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone.[64]

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.[65][66]


See also Insect defences
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 like formic acid. 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. The sting of Jack jumper ants can be fatal,[67] and an antivenom has been developed.[68] Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids.[69] Their stings are painful and can be dangerous to hypersensitive people.[70]

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.[71] One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 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 backwards to escape a threat.[71] Before the strike, 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 hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys,[71] plus some members of the Dacetini tribe,[72] 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. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies.[73] Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.[74]

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. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates.[75] Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana[76] while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.[77]

Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture.[78][79] 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.[80] Camponotus anderseni which nests in the cavities of wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.[81]


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 leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.[82]

Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their 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 less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.[83]

Nest construction

Leaf nest of weaver ants, Pamalican, Philippines.

Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can 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,[53] 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.[84]

The army ants of South America 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.[85]

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.[86]

Food cultivation

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

Most ants are generalist predators, scavengers and indirect herbivores,[18] but a few have evolved specialised ways of obtaining nutrition. 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. Workers specialise in 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 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 that may harm the fungi.[87]


Foraging ants travel distances of up to 200 metres (700 ft) from their nest[88] and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions 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[89] and also by evaluating the movement of objects in their visual field (optical flow).[90] Directions are measured using the position of the sun.[91] They integrate this information to find the shortest route back to their nest.[92] Like all ants they can also make use of visual landmarks when available[93] and use other olfactory and tactile cues to navigate.[94][95] Some species of ant are able to use the Earth's magnetic field for navigation.[96] Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction.[97][98] These polarization detectors are sensitive in the ultraviolet region of the light spectrum.[99] In some army ant species, a group of foragers that get separated from the main column can sometimes turn back on themselves and form a circular ant mill. The workers may then run around continuously until they die of exhaustion.[100]


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.[101] There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.[102]

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. These rafts may also have a role in allowing ants to colonise islands.[103] Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.[104]

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 its large eyes instead of its chemical senses to find prey.[105]

Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive.[106] Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults).[107]

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

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.[108]

Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts.[109][110] 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 can 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.[111]

A conflict between the sexes of a species is seen in some species of ants with the reproductives 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 where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.[112]

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 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 ants' food stores, or avoiding predators. These inquilines can 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.[113][114]

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.[115] In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On 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 can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.[116]

Myrmecophilous (ant-loving) caterpillars of the 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.[117] 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 ants' nest where it feeds on the ant larvae.[118]

An ant transporting an aphid.

Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. 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.[119]

Ants 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.[120] Although some ants obtain nectar from flowers, pollination by ants is somewhat rare.[121] 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.[122] Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants.[123] In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.

Many tropical tree species have seeds that are dispersed by ants.[124] Seed dispersal by ants or myrmecochory is widespread and new estimates suggest that nearly 9% of all plant species may have such ant associations.[125][126] Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal as the seeds are transported to safety below the ground. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food.[127] 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.[128]

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.[88] Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species.[129] The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical.[130] It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.[88] Army ants forage in a wide roving column attacking any animals in that path that are unable to escape. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers.[131]

Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants.[132] 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.[132]

Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores[133] in a microhabitat that best suits the fungus.[134] Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates.[135] 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, and makes them 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 the further spread of the nematode.[136]

Spiders sometimes feed on ants.

South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants.[137] Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.[138] This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.[139] Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia 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, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.[140]

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.[9] On the other hand, ants can 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.[141][142][143]

Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America[144] and the Myrmecia ants of Australia.[145]

In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds 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 can be collected from one ant-heap.[146][147]

Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.[148]

It has been estimated by E.O. Wilson that the total number of individual ants alive in the world at any one time is between one and ten quadrillion (short scale). According to this estimate, the total biomass of all the ants in the world is approximately equal to the total biomass of the entire human race.[149]

As food

Roasted Ants in Colombia
Ant larvae on sale in Isaan, Thailand.

Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.[150]

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.[151] Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (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".[152]

In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).[152]

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 pests,[10] and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.

Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about 14 litres (3 imp gal; 4 US gal) per mound.[153]

In science and technology

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.[154] Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies.[155] Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures.[156] Individuals may be tracked for study by marking them with colours.[157]

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. 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.[8]

In culture

Aesop's ants: picture by Milo Winter, 1888-1956.

Anthropomorphised ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts.[158][159] In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman (Arabic: سليمان‎) 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][160] In parts of Africa, ants are considered to be the messengers of the gods. 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.[161] Ant bites are used in the initiation ceremonies of some Amazon Indian cultures as a test of endurance.[162][163]

Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad.[164] Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced including Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man. 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.[165]

The Chinese character for ant (simplified Chinese: ; traditional Chinese: ; pinyin: ) is a combination of two logograms that may be interpreted as "insect (simplified Chinese: ; traditional Chinese: ; pinyin: chóng) which behaves properly (simplified Chinese: ; traditional Chinese: ; pinyin: )".[166] The traditional Chinese character (蟻) used in Japanese shares this etymology.[167] In spoken Chinese the ant is usually referred to as mǎyĭ (simplified Chinese: 蚂蚁; traditional Chinese: 螞蟻).

From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility.[168] In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".[169]

Ants are also 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 film Them!, and ants mutated into super intelligence in Phase IV. In 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.[170]

See also


  1. ^ Ward, Philip S (2007). "Phylogeny, classification, and species-level taxonomy of ants (Hymenoptera: Formicidae)" (PDF). Zootaxa 1668: 549–563. 
  2. ^ a b Rabeling C, Brown JM & Verhaagh M (2008). "Newly discovered sister lineage sheds light on early ant evolution". PNAS 105 (39): 14913–7. Bibcode 2008PNAS..10514913R. doi:10.1073/pnas.0806187105. PMC 2567467. PMID 18794530. 
  3. ^ "Hymenoptera name server. Formicidae species count.". Ohio State University. 
  4. ^ La nueva taxonomía de hormigas. Pages 45-48 in Fernández, F. Introducción a las hormigas de la región neotropical.. Instituto Humboldt, Bogotá. 2003. 
  5. ^ Oster GF, Wilson EO (1978). Caste and ecology in the social insects. Princeton University Press, Princeton. pp. 21–22. ISBN 0-691-02361-1. 
  6. ^ a b Schultz TR (2000). "In search of ant ancestors". Proceedings of the National Academy of Sciences 97 (26): 14028–14029. Bibcode 2000PNAS...9714028S. doi:10.1073/pnas.011513798. PMC 34089. PMID 11106367. 
  7. ^ a b Hölldobler & Wilson (1990), p. 471
  8. ^ a b Dicke E, Byde A, Cliff D, Layzell P (2004). "An ant-inspired technique for storage area network design". In A. J. Ispeert, M. Murata & N. Wakamiya. Proceedings of Biologically Inspired Approaches to Advanced Information Technology: First International Workshop, BioADIT 2004 LNCS 3141. pp. 364–379. 
  9. ^ a b Hölldobler & Wilson (1990), pp. 619-629
  10. ^ a b "Pest Notes: Ants (Publication 7411)". University of California Agriculture and Natural Resources. 2007. Retrieved 5 June 2008. 
  11. ^ ""ant". Merriam-Webster Online Dictionary". Merriam-Webster. Retrieved 6 June 2008. 
  12. ^ "Ant. Online Etymology Dictionary". Retrieved 30 May 2009. 
  13. ^ Simpson DP (1979). Cassell's Latin Dictionary (5 ed.). London: Cassell Ltd. ISBN 0-304-52257-0. 
  14. ^ Formic
  15. ^ Brothers DJ (1999). "Phylogeny and evolution of wasps, ants and bees (Hymenoptera, Chrysisoidea, Vespoidea, and Apoidea)". Zoologica Scripta 28: 233–249. doi:10.1046/j.1463-6409.1999.00003.x. 
  16. ^ a b Grimaldi D, Agosti D (2001). "A formicine in New Jersey Cretaceous amber (Hymenoptera: Formicidae) and early evolution of the ants". Proceedings of the National Academy of Sciences 97 (25): 13678–13683. Bibcode 2000PNAS...9713678G. doi:10.1073/pnas.240452097. PMC 17635. PMID 11078527. 
  17. ^ Moreau CS, Bell CD, Vila R, Archibald SB, Pierce NE (2006). "Phylogeny of the ants: Diversification in the Age of Angiosperms". Science 312 (5770): 101–104. Bibcode 2006Sci...312..101M. doi:10.1126/science.1124891. PMID 16601190. 
  18. ^ a b Wilson EO, Hölldobler B (2005). "The rise of the ants: A phylogenetic and ecological explanation". Proceedings of the National Academy of Sciences 102 (21): 7411–7414. Bibcode 2005PNAS..102.7411W. doi:10.1073/pnas.0502264102. PMC 1140440. PMID 15899976. 
  19. ^ Wilson E O, Carpenter FM, Brown WL (1967). "The first Mesozoic ants". Science 157 (3792): 1038–1040. Bibcode 1967Sci...157.1038W. doi:10.1126/science.157.3792.1038. PMID 17770424. 
  20. ^ Hölldobler & Wilson (1990), pp. 23-24
  21. ^ Thorne, Barbara L (1997). "Evolution of eusociality in termites" (PDF). Annu. Rev. Ecol. Syst. 28: 27–53. doi:10.1146/annurev.ecolsys.28.1.27. 
  22. ^ "Order Isoptera - Termites". Iowa State University Entomology. 16 February 2004. Retrieved 12 June 2008. 
  23. ^ "Family Mutillidae - Velvet ants". Iowa State University Entomology. 16 February 2004. Retrieved 12 June 2008. 
  24. ^ Hölldobler & Wilson (1990), p. 4
  25. ^ Jones, Alice S. "Fantastic ants - Did you know?". National Geographic Magazine. Retrieved 5 July 2008. 
  26. ^ Thomas, Philip (2007). "Pest Ants in Hawaii". Hawaiian Ecosystems at Risk project (HEAR). Retrieved 6 July 2008. 
  27. ^ Hölldobler & Wilson (1990), p. 589
  28. ^ Shattuck SO (1999). Australian ants: their biology and identification. Collingwood, Vic: CSIRO. p. 149. ISBN 0-643-06659-4. 
  29. ^ Schaal, Stephan (27 Jan 2006). "Messel". Encyclopedia of Life Sciences. doi:10.1038/npg.els.0004143. ISBN 0470016175. 
  30. ^ Agosti D, Johnson NF (eds.) (2005). "Antbase". American Museum of Natural History. Retrieved 6 July 2008. 
  31. ^ Agosti D, Majer JD, Alonso JE, Schultz TR (eds.) (2000). Ants: Standard methods for measuring and monitoring biodiversity. Smithsonian Institution Press. 
  32. ^ Johnson NF (2007). "Hymenoptera name server". Ohio State University. Retrieved 6 July 2008. 
  33. ^ Borror, Triplehorn & Delong (1989), p. 737
  34. ^ a b c d Borror, Triplehorn & Delong (1989), pp. 24-71
  35. ^ Fent K, Rudiger W (1985). "Ocelli: A celestial compass in the desert ant Cataglyphis". Science 228 (4696): 192–194. Bibcode 1985Sci...228..192F. doi:10.1126/science.228.4696.192. PMID 17779641. 
  36. ^ Eisner T, Happ GM (1962). "The infrabuccal pocket of a formicine ant: a social filtration device". Psyche 69 (3): 107–116. doi:10.1155/1962/25068. 
  37. ^ Wilson EO (1953). "The origin and evolution of polymorphism in ants". Quarterly Review of Biology 28 (2): 136–56. doi:10.1086/399512. 
  38. ^ Weber, NA (1946). "Dimorphism in the African Oecophylla worker and an anomaly (Hym.: Formicidae)" (PDF). Annals of the Entomological Society of America 39: 7–10. 
  39. ^ Edward O. Wilson and Robert W. Taylor (1964). "A Fossil Ant Colony: New Evidence of Social Antiquity" (PDF). Psyche 71 (2): 93–103. doi:10.1155/1964/17612. 
  40. ^ Moffett MW, Tobin JE (1991). "Physical castes in ant workers: a problem for Daceton armigerum and other ants" (PDF). Psyche 98 (4): 283–292. doi:10.1155/1991/30265. Archived from the original on 2008-02-27. 
  41. ^ 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. 
  42. ^ Hughes WOH, Sumner S, Van Borm S, Boomsma JJ (2003). "Worker caste polymorphism has a genetic basis in Acromyrmex leaf-cutting ants". Proceedings of the National Academy of Sciences 100 (16): 9394–9397. Bibcode 2003PNAS..100.9394H. doi:10.1073/pnas.1633701100. PMC 170929. PMID 12878720. 
  43. ^ Rossa KG, Kriegera MJB, Shoemaker DD (2003). "Alternative genetic foundations for a key social polymorphism in fire ants". Genetics 165 (4): 1853–1867. PMC 1462884. PMID 14704171. 
  44. ^ Crosland MWJ, Crozier RH (1986). "Myrmecia pilosula, an ant with only one pair of chromosomes". Science 231 (4743): 1278. Bibcode 1986Sci...231.1278C. doi:10.1126/science.231.4743.1278. PMID 17839565. 
  45. ^ Tsutsui ND, Suarez AV, Spagna JC, Johnston JS (2008). "The evolution of genome size in ants". BMC Evolutionary Biology 8 (64): 64. doi:10.1186/1471-2148-8-64. PMC 2268675. PMID 18302783. Retrieved 2008-06-25. 
  46. ^ Gillott, Cedric (1995). Entomology. Springer. p. 325. ISBN 0-306-44967-6. 
  47. ^ 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. 
  48. ^ Hölldobler & Wilson (1990), pp. 351, 372
  49. ^ Traniello JFA (1989). "Foraging strategies of ants". Annual Review of Entomology 34: 191–210. doi:10.1146/annurev.en.34.010189.001203. 
  50. ^ Sorensen A, Busch TM, Vinson SB (1984). "Behavioral flexibility of temporal sub-castes in the fire ant, Solenopsis invicta, in response to food". Psyche 91 (3–4): 319–332. doi:10.1155/1984/39236. 
  51. ^ Peeters C, Holldobler B (1995). "Reproductive cooperation between queens and their mated workers: the complex life history of an ant with a valuable nest" (PDF). Proceedings of the National Academy of Sciences 92 (24): 10977–10979. Bibcode 1995PNAS...9210977P. doi:10.1073/pnas.92.24.10977. PMC 40553. PMID 11607589. 
  52. ^ Taylor, Richard W. (2007). "Bloody funny wasps! Speculations on the evolution of eusociality in ants". In Snelling, R. R., B. L. Fisher, & P. S. Ward (PDF). Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of contributions. Memoirs of the American Entomological Institute, 80. American Entomological Institute. pp. 580–609. 
  53. ^ a b Hölldobler & Wilson (1990), pp. 143-179
  54. ^ Heinze, Jurgen; Tsuji, Kazuki (1995). "Ant reproductive strategies" (PDF). Res. Popul. Ecol. 37 (2): 135–149. doi:10.1007/BF02515814. 
  55. ^ Himler, Anna G.; Caldera, EJ; Baer, BC; Fernández-Marín, H; Mueller, UG (2009). "No sex in fungus-farming ants or their crops". Proc. R. Soc. B 276 (1667): 2611–6. doi:10.1098/rspb.2009.0313. PMC 2686657. PMID 19369264. 
  56. ^ Keller L (1998). "Queen lifespan and colony characteristics in ants and termites". Insectes Sociaux 45 (3): 235–246. doi:10.1007/s000400050084. 
  57. ^ a b Franks NR, Resh VH, Cardé RT (eds) (2003). Encyclopedia of Insects. San Diego: Academic Press. pp. 29–32. ISBN 0-12-586990-8. 
  58. ^ Kipyatkov VE (2001). "Seasonal life cycles and the forms of dormancy in ants (Hymenoptera, Formicoidea)". Acta Societatis Zoologicae Bohemicae 65 (2): 198–217. 
  59. ^ Jackson DE, Ratnieks FL (August 2006). "Communication in ants". Curr. Biol. 16 (15): R570–R574. doi:10.1016/j.cub.2006.07.015. PMID 16890508. 
  60. ^ Goss S, Aron S, Deneubourg JL, Pasteels JM (1989). "Self-organized shortcuts in the Argentine ant". Naturwissenschaften 76 (12): 579–581. Bibcode 1989NW.....76..579G. doi:10.1007/BF00462870. 
  61. ^ D'Ettorre P, Heinze J (2001). "Sociobiology of slave-making ants". Acta ethologica 3 (2): 67–82. doi:10.1007/s102110100038. 
  62. ^ Detrain C, Deneubourg JL, Pasteels JM (1999). Information processing in social insects. Birkhäuser. pp. 224–227. ISBN 3-7643-5792-4. 
  63. ^ Greene MJ, Gordon DM (2007). "Structural complexity of chemical recognition cues affects the perception of group membership in the ants Linephithema humile and Aphaenogaster cockerelli". Journal of Experimental Biology 210 (Pt 5): 897–905. doi:10.1242/jeb.02706. PMID 17297148. 
  64. ^ Hölldobler & Wilson (1990), p. 354
  65. ^ Hickling R, Brown RL (2000). "Analysis of acoustic communication by ants". Journal of the Acoustical Society of America 108 (4): 1920–1929. Bibcode 2000ASAJ..108.1920H. doi:10.1121/1.1290515. PMID 11051518. 
  66. ^ Roces F, Hölldobler B (1996). "Use of stridulation in foraging leaf-cutting ants: Mechanical support during cutting or short-range recruitment signal?". Behavioral Ecology and Sociobiology 39 (5): 293. doi:10.1007/s002650050292. 
  67. ^ Clarke PS (1986). "The natural history of sensitivity to jack jumper ants (Hymenoptera: Formicidae: Myrmecia pilosula) in Tasmania". Medical Journal of Australia 145 (11–12): 564–566. PMID 3796365. 
  68. ^ Brown SGA, Heddle RJ, Wiese MD, Blackman KE (2005). "Efficacy of ant venom immunotherapy and whole body extracts". Journal of Allergy and Clinical Immunology 116 (2): 464–465. doi:10.1016/j.jaci.2005.04.025. PMID 16083810. 
  69. ^ Obin MS, Vander Meer RK (1985). "Gaster flagging by fire ants (Solenopsis spp.): Functional significance of venom dispersal behavior". Journal of Chemical Ecology 11 (12): 1757–1768. doi:10.1007/BF01012125. 
  70. ^ Stafford CT (1996). "Hypersensitivity to fire ant venom". Annals of allergy, asthma, & immunology 77 (2): 87–99. doi:10.1016/S1081-1206(10)63493-X. 
  71. ^ a b c Patek SN, Baio JE, Fisher BL, Suarez AV (22 August 2006). "Multifunctionality and mechanical origins: Ballistic jaw propulsion in trap-jaw ants". Proceedings of the National Academy of Sciences 103 (34): 12787–12792. Bibcode 2006PNAS..10312787P. doi:10.1073/pnas.0604290103. PMC 1568925. PMID 16924120. 
  72. ^ Gronenberg W (1996). "The trap-jaw mechanism in the Dacetine ant Daceton armigerum and Strumigenys sp" (PDF). The Journal of Experimental Biology 199 (9): 2021–2033. 
  73. ^ Jones, T.H.; Clark, D.A.; Edwards, A.A.; Davidson, D.W.; Spande, T.F. & Snelling, Roy R. (2004). "The Chemistry of Exploding Ants, Camponotus spp. (Cylindricus complex)". Journal of Chemical Ecology 30 (8): 1479–1492. doi:10.1023/B:JOEC.0000042063.01424.28. PMID 15537154. 
  74. ^ Tofilski,Adam; Couvillon, MJ;Evison, SEF; Helantera, H; Robinson, EJH; Ratnieks, FLW (2008). "Preemptive Defensive Self-Sacrifice by Ant Workers" (PDF). The American Naturalist 172 (5): E239–E243. doi:10.1086/591688. PMID 18928332. 
  75. ^ Julian GE, Cahan S (1999). "Undertaking specialization in the desert leaf-cutter ant Acromyrmex versicolor". Animal Behaviour 58 (2): 437–442. doi:10.1006/anbe.1999.1184. PMID 10458895. 
  76. ^ López-Riquelme GO, Malo EA, Cruz-López L, Fanjul-Moles ML (2006). "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. doi:10.1111/j.1365-3032.2006.00526.x. 
  77. ^ Choe, Dong-Hwan;Millar JG; Rust MK (2009). "Chemical signals associated with life inhibit necrophoresis in Argentine ants". Proc. Nat. Acad. Sci. 106 (20): 8251–8255. Bibcode 2009PNAS..106.8251C. doi:10.1073/pnas.0901270106. PMC 2688878. PMID 19416815. 
  78. ^ Tschinkel WR (2004). "The nest architecture of the Florida harvester ant, Pogonomyrmex badius". Journal of Insect Science 4 (21): 1–19. 
  79. ^ Peeters C, Hölldobler B, Moffett M, Musthak Ali TM (1994). ""Wall-papering" and elaborate nest architecture in the ponerine ant Harpegnathos saltator". Insectes Sociaux 41 (2): 211–218. doi:10.1007/BF01240479. 
  80. ^ Maschwitz U, Moog J (2000). "Communal peeing: a new mode of flood control in ants". Naturwissenschaften 87 (12): 563–565. Bibcode 2000NW.....87..563M. doi:10.1007/s001140050780. PMID 11198200. 
  81. ^ Nielsen MG, Christian KA (2007). "The mangrove ant, Camponotus anderseni switches to anaerobic respiration in response to elevated CO2 levels". Journal of Insect Physiology 53 (5): 505–508. doi:10.1016/j.jinsphys.2007.02.002. PMID 17382956. 
  82. ^ Franks NR, Richardson T (2006). "Teaching in tandem-running ants". Nature 439 (7073): 153. Bibcode 2006Natur.439..153F. doi:10.1038/439153a. PMID 16407943. 
  83. ^ Ravary F, Lecoutey E, Kaminski G, Châline N, Jaisson P (2007). "Individual experience alone can generate lasting division of labor in ants". Current Biology 17 (15): 1308–1312. doi:10.1016/j.cub.2007.06.047. PMID 17629482. 
  84. ^ Franks NR, Hooper J, Webb C, Dornhaus A (2005). "Tomb evaders: house-hunting hygiene in ants". Biology Letters 1 (2): 190–192. doi:10.1098/rsbl.2005.0302. PMC 1626204. PMID 17148163. 
  85. ^ Hölldobler & Wilson (1990), p. 573
  86. ^ Robson SK, Kohout RJ (2005). "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. doi:10.1111/j.1440-6055.2005.00462.x. 
  87. ^ Schultz TR (1999). "Ants, plants and antibiotics". Nature 398 (6730): 747–748. Bibcode 1999Natur.398..747S. doi:10.1038/19619. 
  88. ^ a b c Carrol CR, Janzen DH (1973). "Ecology of foraging by ants". Annual Review of Ecology and Systematics 4: 231–257. doi:10.1146/ 
  89. ^ Wittlinger M, Wehner R, Wolf H (2006). "The Ant Odometer: Stepping on Stilts and Stumps". Science 312 (5782): 1965–7. Bibcode 2006Sci...312.1965W. doi:10.1126/science.1126912. PMID 16809544. 
  90. ^ Ronacher B, Werner R (1995). "Desert ants Cataglyphis fortis use self-induced optic flow to measure distances travelled". Journal of Comparative Physiology A. 
  91. ^ Werner R (2003). "Desert ant navigation: how miniature brains solve complex tasks". Journal of Comparative Physiology 189 (8): 579–588. doi:10.1007/s00359-003-0431-1. PMID 12879352. 
  92. ^ Sommer S, Wehner R (2004). "The ant's estimation of distance travelled: experiments with desert ants, Cataglyphis fortis". Journal of Comparative Physiology 190 (1): 1–6. doi:10.1007/s00359-003-0465-4. PMID 14614570. 
  93. ^ Åkesson S, Wehner R (2002). "Visual navigation in desert ants Cataglyphis fortis: are snapshots coupled to a celestial system of reference?" (PDF). Journal of Experimental Biology 205 (14): 1971–1978. 
  94. ^ Steck K, Hansson B, Knaden M (2009). "Smells like home: Desert ants, Cataglyphis fortis, use olfactory landmarks to pinpoint the nest" (PDF). Frontiers in Zoology. 
  95. ^ Seidl T, Wehner R (2006). "Visual and tactile learning of ground structures in desert ants" (PDF). Journal of Experimental Biology 209 (Pt 17): 3336–44. doi:10.1242/jeb.02364. PMID 16916970. 
  96. ^ Banks AN, Srygley RB (2003). "Orientation by magnetic field in leaf-cutter ants, Atta colombica (Hymenoptera: Formicidae)". Ethology 109 (10): 835–846. doi:10.1046/j.0179-1613.2003.00927.x. 
  97. ^ Fukushi T (15 June 2001). "Homing in wood ants, Formica japonica: use of the skyline panorama". Journal of Experimental Biology 204 (12): 2063–2072. PMID 11441048. 
  98. ^ Wehner R, Menzel R (1969). "Homing in the ant Cataglyphis bicolor". Science 164 (3876): 192–194. Bibcode 1969Sci...164..192W. doi:10.1126/science.164.3876.192. PMID 5774195. 
  99. ^ Chapman, Reginald Frederick (1998). The Insects: Structure and Function (4 ed.). Cambridge University Press. p. 600. ISBN 0-521-57890-6. 
  100. ^ Delsuc F (2003). "Army Ants Trapped by Their Evolutionary History". PLoS Biol. 1 (2): E37. doi:10.1371/journal.pbio.0000037. PMC 261877. PMID 14624241. 
  101. ^ Baroni-Urbani C, Boyan GS, Blarer A, Billen J, Musthak Ali TM (1994). "A novel mechanism for jumping in the Indian ant Harpegnathos saltator (Jerdon) (Formicidae, Ponerinae)". Experientia 50: 63–71. doi:10.1007/BF01992052. 
  102. ^ Yanoviak SP, Dudley R, Kaspari M (2005). "Directed aerial descent in canopy ants" (PDF). Nature 433 (7026): 624–626. Bibcode 2005Natur.433..624Y. doi:10.1038/nature03254. PMID 15703745. 
  103. ^ 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. 
  104. ^ Clay RE, Andersen AN (1996). "Ant fauna of a mangrove community in the Australian seasonal tropics, with particular reference to zonation". Australian Journal of Zoology 44 (5): 521–533. doi:10.1071/ZO9960521. 
  105. ^ Crosland MWJ, Crozier RH, Jefferson E (1988). "Aspects of the biology of the primitive ant genus Myrmecia F. (Hymenoptera: Formicidae)". Australian Journal of Entomology 27 (4): 305–309. doi:10.1111/j.1440-6055.1988.tb01179.x. 
  106. ^ Diehl E, Junqueira LK, Berti-Filho E (2005). "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. doi:10.1590/S1519-69842005000300008. 
  107. ^ Achenbach, A; Foitzik, Susanne (2009). "First evidence for slave rebellion: enslaved ant workers systematically kill the brood of their social parasite Protomognathus americanus". Evolution 63 (4): 1068–1075. doi:10.1111/j.1558-5646.2009.00591.x. PMID 19243573.  See also New Scientist, 2009 April 9
  108. ^ Henderson G, Andersen JF, Phillips JK, Jeanne RL (2005). "Internest aggression and identification of possible nestmate discrimination pheromones in polygynous ant Formica montana". Journal of Chemical Ecology 16 (7): 2217–2228. doi:10.1007/BF01026932. 
  109. ^ 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. 
  110. ^ Taylor RW (1968). "The Australian workerless inquiline ant, Strumigenys xenos Brown (Hymenoptera-Formicidae) recorded from New Zealand". New Zealand Entomologist 4 (1): 47–49. 
  111. ^ Hölldobler & Wilson (1990), pp. 436—448
  112. ^ Fournier, D, Estoup A, Orivel J, Foucaud J, Jourdan H, Le Breton J, Keller L (2005). "Clonal reproduction by males and females in the little fire ant". Nature 435 (7046): 1230–1234. Bibcode 2005Natur.435.1230F. doi:10.1038/nature03705. PMID 15988525. 
  113. ^ Reiskind J (1977). "Ant-mimicry in Panamanian clubionid and salticid spiders (Araneae: Clubionidae, Salticidae)". Biotropica 9 (1): 1–8. doi:10.2307/2387854. JSTOR 2387854. 
  114. ^ Cushing PE (1997). "Myrmecomorphy and myrmecophily in spiders: A Review" (PDF). The Florida Entomologist 80 (2): 165–193. doi:10.2307/3495552. JSTOR 3495552. 
  115. ^ Styrsky JD, Eubanks MD (January 2007). "Ecological consequences of interactions between ants and honeydew-producing insects". Proc. Biol. Sci. 274 (1607): 151–164. doi:10.1098/rspb.2006.3701. PMC 1685857. PMID 17148245. 
  116. ^ Jahn GC, Beardsley JW (1996). "Effects of Pheidole megacephala (Hymenoptera: Formicidae) on survival and dispersal of Dysmicoccus neobrevipes (Homoptera: Pseudococcidae)". Journal of Economic Entomology 89: 1124–1129. 
  117. ^ DeVries PJ (1992). "Singing caterpillars, ants and symbiosis". Scientific American 267 (4): 76. doi:10.1038/scientificamerican1092-76. 
  118. ^ Pierce NE, Braby MF, Heath A (2002). "The ecology and evolution of ant association in the Lycaenidae (Lepidoptera)". Annual Review of Entomology 47: 733–771. doi:10.1146/annurev.ento.47.091201.145257. PMID 11729090. 
  119. ^ Dejean A, Solano PJ, Ayroles J, Corbara B, Orivel J (2005). "Arboreal ants build traps to capture prey". Nature 434 (7036): 973. Bibcode 2005Natur.434..973D. doi:10.1038/434973a. PMID 15846335. 
  120. ^ Frederickson ME, Gordon DM (2007). "The devil to pay: a cost of mutualism with Myrmelachista schumanni ants in 'devil's gardens' is increased herbivory on Duroia hirsuta trees" (PDF). Proceedings of the Royal Society B 274 (1613): 1117–1123. doi:10.1098/rspb.2006.0415. PMC 2124481. PMID 17301016. 
  121. ^ Beattie, Andrew J.; Christine Turnbull, R. B. Knox and 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. 
  122. ^ Katayama N, Suzuki N (2004). "Role of extrafloral nectaries of Vicia faba in attraction of ants and herbivore exclusion by ants". Entomological Science 7 (2): 119–124. doi:10.1111/j.1479-8298.2004.00057.x. 
  123. ^ Fischer RC, Wanek W, Richter A, Mayer V (2003). "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. doi:10.1046/j.1365-2745.2003.00747.x. 
  124. ^ Hanzawa FM, Beattie AJ, Culver DC (1988). "Directed dispersal: demographic analysis of an ant-seed mutualism". American Naturalist 131 (1): 1–13. doi:10.1086/284769. 
  125. ^ 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. 
  126. ^ Lengyel, Szabolcs; Aaron D. Gove, Andrew M. Latimer, Jonathan D. Majer, Robert R. Dunn (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. 
  127. ^ Fischer RC, Ölzant SM, Wanek W, Mayer V (2005). "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. doi:10.1007/s00040-004-0773-x. 
  128. ^ Hughes L, Westoby M (1 January 1992). "Capitula on stick insect eggs and elaiosomes on seeds: convergent adaptations for burial by ants". Functional Ecology 6 (6): 642–648. doi:10.2307/2389958. ISSN 0269-8463. JSTOR 2389958. 
  129. ^ Quinet Y, Tekule N & de Biseau JC (2005). "Behavioural Interactions Between Crematogaster brevispinosa rochai Forel (Hymenoptera: Formicidae) and Two Nasutitermes Species (Isoptera: Termitidae)". Journal of Insect Behavior 18 (1): 1–17. doi:10.1007/s10905-005-9343-y. 
  130. ^ Jeanne, RL (1972). "Social biology of the neotropical wasp Mischocyttarus drewseni". Bull. Mus. Comp. Zool. 144: 63–150. 
  131. ^ Willis, E. & Y. Oniki (1978). "Birds and Army Ants". Annual Review of Ecology and Systematics 9: 243–263. doi:10.1146/ 
  132. ^ a b Sivinski J, Marshall S, Petersson E (1999). "Kleptoparasitism and phoresy in the Diptera" (PDF). Florida Entomologist 82 (2): 179–197. doi:10.2307/3496570. JSTOR 3496570. 
  133. ^ Schaechter E (2000). "Some weird and wonderful fungi". Microbiology Today 27 (3): 116–117. 
  134. ^ Sandra B. Andersen, Sylvia Gerritsma, Kalsum M. Yusah, David Mayntz, Nigel L. Hywel‐Jones, Johan Billen, Jacobus J. Boomsma, and David P. Hughes (2009). "The Life of a Dead Ant: The Expression of an Adaptive Extended Phenotype". The American Naturalist 174 (3): 424–433. doi:10.1086/603640. PMID 19627240. 
  135. ^ Wojcik DP (1989). "Behavioral interactions between ants and their parasites". The Florida Entomologist 72 (1): 43–51. doi:10.2307/3494966. JSTOR 3494966. 
  136. ^ Poinar G Jr., Yanoviak SP (2008). "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" (PDF). Systematic Parasitology 69 (2): 145–153. doi:10.1007/s11230-007-9125-3. PMID 18038201. 
  137. ^ 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. 
  138. ^ Vellely AC (2001). "Foraging at army ant swarms by fifty bird species in the highlands of Costa Rica" (PDF). Ornitologia Neotropical 12: 271–275. Retrieved 8 June 2008. 
  139. ^ 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. 
  140. ^ Swenson JE, Jansson A, Riig R, Sandegren R (1999). "Bears and ants: myrmecophagy by brown bears in central Scandinavia". Canadian Journal of Zoology 77 (4): 551–561. doi:10.1139/cjz-77-4-551. 
  141. ^ Gottrup F, Leaper D (2004). "Wound healing: Historical aspects" (PDF). EWMA Journal 4 (2): 5. Archived from the original on 2007-06-16. 
  142. ^ Gudger EW (1925). "Stitching wounds with the mandibles of ants and beetles". Journal of the American Medical Association 84: 1861–1864. 
  143. ^ Sapolsky, Robert M. (2001). A Primate's Memoir: A Neuroscientist's Unconventional Life Among the Baboons. Simon and Schuster. p. 156. ISBN 0-7432-0241-4. 
  144. ^ Haddad Jr. V, Cardoso JLC, Moraes RHP (2005). "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)" (PDF). Revista do Instituto de Medicina Tropical de São Paulo 47 (4): 235–238. doi:10.1590/S0036-46652005000400012. 
  145. ^ McGain F, Winkel KD (2002). "Ant sting mortality in Australia". Toxicon 40 (8): 1095–1100. doi:10.1016/S0041-0101(02)00097-1. PMID 12165310. 
  146. ^ Downes D, Laird SA (1999). "Innovative mechanisms for sharing benefits of biodiversity and related knowledge" (PDF). The Center for International Environmental Law. Retrieved 8 June 2008. 
  147. ^ Cheney RH, Scholtz E (1963). "Rooibos tea, a South African contribution to world beverages". Economic Botany 17 (3): 186–194. doi:10.1007/BF02859435. 
  148. ^ Chapman, RE; Bourke, Andrew FG (2001). "The influence of sociality on the conservation biology of social insects" (PDF). Ecology Letters 4 (6): 650–662. doi:10.1046/j.1461-0248.2001.00253.x. 
  149. ^ Holldobler, B & E. O. Wilson (2009). The Superorganism: The Beauty, Elegance, and Strangeness of Insect Societies. New York: W. W. Norton. p. 5. ISBN 0-393-06704-1. 
  150. ^ DeFoliart GR (1999). "Insects as food: Why the western attitude is important". Annual Review of Entomology 44: 21–50. doi:10.1146/annurev.ento.44.1.21. PMID 9990715. 
  151. ^ Bingham CT (1903). Fauna of British India. Hymenoptera Volume 3. p. 311. 
  152. ^ a b Bequaert J (1921). "Insects as food: How they have augmented the food supply of mankind in early and recent times". Natural History Journal 21: 191–200. 
  153. ^ "Two step method for fire ant control". Oklahoma State University. Retrieved 5 June 2008. 
  154. ^ Lubbock, J. (1881). "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. doi:10.1111/j.1096-3642.1882.tb02275.x. 
  155. ^ Stadler B, Dixon, AFG (2008). Mutualism: Ants and their insect partners. Cambridge University Press. ISBN 139780521860352. 
  156. ^ Kennedy CH (1951). "Myrmecological technique. IV. Collecting ants by rearing pupae". The Ohio Journal of Science 51 (1): 17–20. 
  157. ^ Wojcik DP, Burges RJ, Blanton CM, Focks DA (2000). "An improved and quantified technique for marking individual fire ants (Hymenoptera: Formicidae)" (PDF). The Florida Entomologist 83 (1): 74–78. doi:10.2307/3496231. JSTOR 3496231. 
  158. ^ Quran 27:18–19. 
  159. ^ Sahih Bukhari Vol 4, Book 54, Number 536. 
  160. ^ Deen, Mawil Y. Izzi (1990). "Islamic Environmental Ethics, Law, and Society". In Engel JR & JG Engel (PDF). Ethics of Environment and Development. Bellhaven Press, London. 
  161. ^ 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. 
  162. ^ (French) Cesard N, Deturche J, Erikson P (2003). "Les Insectes dans les pratiques médicinales et rituelles d’Amazonie indigène". In Motte-Florac, E. & J. M. C. Thomas. Les insectes dans la tradition orale. Peeters-Selaf, Paris. pp. 395–406. 
  163. ^ Schmidt RJ (1985). "The super-nettles: a dermatologist's guide to ants in the plants". International Journal of Dermatology 24 (4): 204–210. doi:10.1111/j.1365-4362.1985.tb05760.x. PMID 3891647. 
  164. ^ Twain, Mark (1880). "22 The Black Forest and Its Treasures". A Tramp Abroad. New York: Oxford University Press. ISBN 0-19-510137-5. 
  165. ^ Wilson, EO (25 January 2010). "Trailhead". The New Yorker. pp. 56–62. 
  166. ^ Harbaugh, Rick (1998). Chinese Characters: A Genealogy and Dictionary. Yale University Press. ISBN 0-9660750-0-5. 
  167. ^ Hearn L (1904). Kwaidan: Stories and studies Of strange things. Tuttle publishing (2005 reprint). p. 223. ISBN 0-8048-3662-0. 
  168. ^ Guri, Assaf (8 September 1998). Habitat media for ants and other invertebrates (US Patent 5803014). United States Patent and Trademark Office 
  169. ^ "1992 Excellence in Software Awards Winners". Software & Information Industry Association. Retrieved 3 April 2008. 
  170. ^ Sharkey AJC (2006). "Robots, insects and swarm intelligence". Artificial Intelligence Review 26 (4): 255–268. doi:10.1007/s10462-007-9057-y. 

Cited texts

  • Borror DJ, Triplehorn CA, Delong DM (1989). Introduction to the Study of Insects, 6th Edition. Saunders College Publishing. ISBN 0-03-025397-7. 
  • Hölldobler B, Wilson EO (1990). The Ants. Harvard University Press. ISBN 0-674-04075-9. 

Further reading

  • Bolton, Barry (1995). A New General Catalogue of the Ants of the World. Harvard University Press. ISBN 978-0-674-61514-4. 
  • Hölldobler B, Wilson EO (1998). Journey to the Ants: A Story of Scientific Exploration. Belknap Press. ISBN 0-674-48526-2. 
  • Hölldobler B, Wilson EO (1990). The Ants. Harvard University Press. ISBN 978-3-540-52092-4. 
  • Hölldobler B, Wilson EO (2009). The Superorganism: The Beauty, Elegance and Strangeness of Insect Societies. Norton & Co.. ISBN 978-0-393-06704-0. 

External links

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  • ant-s —     ant s     English meaning: forward, before, outer side     Deutsche Übersetzung: “Vorderseite, Stirn”     Material: O.Ind. ánta ḥ “ end, border, edge “ (therefrom antya ḥ “ the last “); Alb. (*ánta) ana ‘side, end”. Gk. gen. sg. κάταντες ( …   Proto-Indo-European etymological dictionary

  • Ant — Cette page d’homonymie répertorie les différents sujets et articles partageant un même nom. {{{image}}}   Sigles d une seule lettre   Sigles de deux lettres > Sigles de trois lettres …   Wikipédia en Français

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  • ant- — ↑ anti (1). * * * ant , Ant : ↑ anti , Anti . * * * ant , Ant : ↑anti , ↑Anti (1). an|ti , An|ti , (vor Vokalen u. gelegentlich vor h:) ant , Ant [griech. antí]: 1. bedeutet in Bildungen mit Substantiven od. Adjektiven gegen[über] …   Universal-Lexikon

  • -ant — ant, ante ❖ ♦ Suffixe d adjectifs, de même origine (lat. antem, terminaison à l accusatif du p. prés. en ans) que le morphème du p. prés. du verbe, ant, invar. en genre et en nombre. || Ex. : une fusée éclairante (adj.); la lampe éclairant la… …   Encyclopédie Universelle

  • Ant — (с англ. муравей), Ants и аббревиатура ANT может относится к: ANT является собственническим протоколом беспроводной сенсорной сети позволяющий работать устройства на частоте 2.4 ГГц промышленного, научного и медицинского распределения… …   Википедия

  • -ant — [ant], der; en, en <Suffix>: besagt, dass das im Basiswort Genannte von der so bezeichneten männlichen Person ausgeführt, ausgeübt wird: aktivische Bedeutung; Basiswort ist meist ein Verb auf »ieren«> /Ggs. and/: a) <verbale Basis> …   Universal-Lexikon

  • ant... — ant...1, Ant...1 〈Vorsilbe〉 = ante..., Ante... ant...2, Ant...2 〈Vorsilbe〉 = anti..., Anti... * * * ant...   [griechisch], anti …   Universal-Lexikon

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