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1 Etymology
The word ant is derived from ante, emete of Middle English which are derived from mette of Old English, and is
related to the dialectal Dutch emt and the Old High German meiza, hence the modern German Ameise. All of
these words come from West Germanic *maitij, and the
original meaning of the word was the biter (from ProtoGermanic *ai-, o, away + *mait- cut).[12][13] The
family name Formicidae is derived from the Latin formca (ant)[14] from which the words in other Romance languages, such as the Portuguese formiga, Italian formica,
Spanish hormiga, Romanian furnic, and French fourmi
are derived. It has been hypothesised that a Proto-IndoEuropean word *morwi- was used, cf. Sanskrit vamrah, Latin formca, Greek mrmx, Old Church
Slavonic mraviji, Old Irish moirb, Old Norse maurr,
Dutch mier.[15]
4 MORPHOLOGY
4 Morphology
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 rst 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
Termites, although sometimes called 'white ants, are not segments).[37]
ants. They belong to the order Isoptera. Termites are
more closely related to cockroaches and mantids. Termites are eusocial, but dier greatly in the genetics of
reproduction. The similarity of their social structure
to that of ants is attributed to convergent evolution.[25]
Velvet ants look like large ants, but are wingless female
wasps.[26][27]
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 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 uids. 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
4.4
Metasoma
4.4 Metasoma
the appendages.[38]
ALITRUNK
HEAD
Scutellum
Pronotum
Occiput
Compound eye
Metanotum
Mesonotum
PETIOLE
Propodeal spiracle
Metathoracic
spiracle
Mesothoracic
spiracle
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 modied
into stings that are used for subduing prey and defending
their nests.[38]
GASTER
Metapleural gland
Bulla
Orifice
Propodeum
Postpetiole (III)
Petiole (II)
(IV)
Funiculus
Scape
Frontal lobes
Tergite
(V)
Anepisternum
Sternite
Antennal fossa
Clypeus
Mandibles
(VI)
Katepisternum
Coxa
(VII)
Trochanter
Ventral process
Sting
Femur
4.5 Polymorphism
Tibia
Tarsal claw
Tibial spurs
Tarsus
4.1
Head
4.2
Legs
4.3
Wings
In the colonies of a few ant species, there are physical castesworkers 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 eective in ghting, 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.[42] Weaver ants, for example, have a distinct bimodal size distribution.[43][44]
Some other species show continuous variation in the
size of workers. The smallest and largest workers in
Pheidologeton diversus show nearly a 500-fold dierence
in their dry-weights.[45] 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.[46]
For instance, these replete workers develop in the North
American honeypot ant Myrmecocystus mexicanus. Rissing found that usually the largest workers in the colony
develop into repletes, and if repletes are removed from
the colony other workers become repletes, demonstrating
the exibility of this particular polymorphism.[47] 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 dierences
between worker castes have been noted in Acromyrmex
sp.[48] These polymorphisms are caused by relatively
small genetic changes; dierences in a single gene of
Solenopsis invicta can decide whether the colony will have
single or multiple queens.[49] 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.[50][51]
6.1
Communication
Ants mating
6
6.1
permit slow and ne movements for other tasks. Trapjaws also are seen in the following genera: Anochetus,
Orectognathus, and Strumigenys,[76] plus some members
of the Dacetini tribe,[77] which are viewed as examples
Some ants produce sounds by stridulation, using the of convergent evolution.
gaster segments and their mandibles. Sounds may be A Malaysian species of ant in the Camponotus cylindriused to communicate with colony members or with other cus group has enlarged mandibular glands that extend into
species.[70][71]
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.[78]
6.2 Defence
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
Ants attack and defend themselves by biting and, in many outside each evening.[79]
species, by stinging, often injecting or spraying chemicals, such as formic acid in the case of formicine ants,
alkaloids and piperidines in re ants, and a variety of protein components in other ants. 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.
See also Insect defences
Ant mound holes prevent water from entering the nest during rain
6.3 Learning
Many animals can learn behaviours by imitation, but
ants may be the only group apart from mammals where
interactive teaching has been observed. A knowledge-
6.5
Cultivation of food
6.4
Nest construction
7
by attaching leaves together, rst 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.[91]
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.[92]
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 articial structures to house their colonies and
nest structures.[93][94]
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,[59] 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 rst sign of
threats.[89]
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.[90]
Weaver ant (Oecophylla spp.) workers build nests in trees
type of leaf is found to be toxic to the fungus, the colony they go to trapped pockets of air in the submerged nests
will no longer collect it. The ants feed on structures pro- to breathe.[115]
duced 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 6.8 Cooperation and competition
fungi.[98]
6.6
Navigation
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 nd prey.[116]
6.9
6.9
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.[126] 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 con-
10
tinued 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.[127]
Myrmecophilous (ant-loving) caterpillars of the buttery
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.[128] 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.[129] 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.[130]
Ants may obtain nectar from owers such as the dandelion but
are only rarely known to pollinate owers.
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.[99] Some termites, including Nasutitermes
corniger, form associations with certain ant species to
keep away predatory ant species.[140] The tropical wasp
Mischocyttarus drewseni coats the pedicel of its nest with
an ant-repellent chemical.[141] 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 o
the nest with bursts of wing buzzing.[142] Stingless bees
(Trigona and Melipona) use chemical defences against
ants.[99] Certain species of ants have the power to drive
certain wasps, such as Polybia occidentalis to extinction
if they attack more than once and the wasps cannot keep
up with rebuilding their nest.
Flies in the Old World genus Bengalia (Calliphoridae)
prey on ants and are kleptoparasites, snatching prey or
brood from the mandibles of adult ants.[143] Wingless and
legless females of the Malaysian phorid y (Vestigipoda
myrmolarvoidea) live in the nests of ants of the genus
Aenictus and are cared for by the ants.[143]
11
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 [144] in a
microhabitat that best suits the fungus.[145] Strepsipteran
parasites also manipulate their ant host to climb grass
stems, to help the parasite nd mates.[146]
12
7.1
As food
7.2 As pests
See also: Ants of medical importance
Some ant species are considered as pests. The presence
The tiny pharaoh ant is a major pest in hospitals and oce blocks;
it can make nests between sheets of paper
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$40 per pound
($90/kg) because they are seasonal and hard to nd. In
the Colombian department of Santander, hormigas cu-
7.4
In culture
13
7.3
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 scientic 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 rst discovered in ants by Sir John Lubbock
in 1881.[166] 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.[167] Ant
colonies may be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass
framed enclosures.[168] Individuals may be tracked for
study by marking them with dots of colours.[169]
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.[9]
7.4
In culture
the Hopi mythology, considers ants as the very rst animals. Ant bites are often said to have curative properties.
The sting of some species of Pseudomyrmex is claimed to
give fever relief.[173] Ant bites are used in the initiation
ceremonies of some Amazon Indian cultures as a test of
endurance.[174][175]
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.[176] 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-ction trilogy is divided between the worlds of
ants and humans; ants and their behaviour is described
using contemporary scientic knowledge. H.G. Wells
wrote about intelligent ants destroying human settlements
in Brazil and threatening human civilization in his 1905
science-ction short story, The Empire of the Ants. In
more recent times, animated cartoons and 3-D animated
lms featuring ants have been produced including Antz, A
Bugs Life, The Ant Bully, The Ant and the Aardvark, 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 antqueen and the rise and fall of her colony, from an ants
point of view.[177]
From the late 1950s through the late 1970s, ant
farms were popular educational childrens toys in the
United States. Later versions use transparent gel instead of soil, allowing greater visibility.[178] In the early
1990s, the video game SimAnt, which simulated an ant
colony, won the 1992 Codie award for Best Simulation
Program.[179]
14
Ants also are quite popular inspiration for many sciencection insectoids, such as the Formics of Enders Game,
the Bugs of Starship Troopers, the giant ants in the lms
[8]
Them! and Empire of the Ants, Marvel Comics' super
hero Ant-Man, and ants mutated into super-intelligence [9]
in Phase IV. In computer strategy games, ant-based
species often benet 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.[180]
[10]
See also
doi:10.1073/pnas.011513798.
11106367.
REFERENCES
PMC 34089.
PMID
Ant robotics
Ant sting
International Union for the Study of Social Insects
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EXTERNAL LINKS
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sharing benets of biodiversity and related knowledge
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digne. In Motte-Florac, E. & J. M. C. Thomas. Les
Archived from the original on 23 April 2008. Retrieved
insectes dans la tradition orale (in French). Peeters-Selaf,
8 June 2008.
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doi:10.1046/j.1461-0248.2001.00253.x.
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The Beauty, Elegance, and Strangeness of Insect Societies.
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New York: W. W. Norton. p. 5. ISBN 0-393-06704-1.
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wasps. IX. Color of owers as an attraction to bees:
Experiments and considerations thereon. J. Linn. Soc.
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[167] Stadler B, Dixon, AFG (2008). Mutualism: Ants and their
insect partners. Cambridge University Press. ISBN 9780-521-86035-2.
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Collecting ants by rearing pupae. The Ohio Journal of
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[170] 27:1819. Quran.
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family and time. Ethnohistory 47 (2): 399422.
doi:10.1215/00141801-47-2-399.
10 Further reading
Bolton, Barry (1995). A New General Catalogue of
the Ants of the World. Harvard University Press.
ISBN 978-0-674-61514-4.
Hlldobler B, Wilson EO (1998). Journey to the
Ants: A Story of Scientic Exploration. Belknap
Press. ISBN 0-674-48526-2.
Hlldobler B, Wilson EO (1990). The Ants. Harvard University Press. ISBN 978-3-540-52092-4.
Hlldobler B, Wilson EO (2009). The Superorganism: The Beauty, Elegance and Strangeness of Insect
Societies. Norton & Co. ISBN 978-0-393-06704-0.
11 External links
AntWeb from The California Academy of Sciences
21
AntBase a taxonomic database with literature
sources
AntWiki Bringing Ants to the World
Ant Species Fact Sheets from the National Pest
Management Association on Argentine, Carpenter,
Pharaoh, Odorous, and other ant species
Ant Genera of the World distribution maps
The super-nettles. A dermatologists guide to antsin-the-plants
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