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ant
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{{Other uses|Ant (disambiguation)}}{{pp-vandalism|small=yes}}{{pp-move-indef}}{hide}automatic taxobox
| name = Ants
| fossil_range = {{fossilrange|140|0|earliest=168|refs={edih} Berriasian â€“ Present
| image = Fire ants 01.jpg
| image_caption = A group of fire ants.
| taxon = Formicidae
| authority = Latreille, 1809
| type_species = Formica rufa
| type_species_authority = Linnaeus, 1761
| subdivision_ranks = Subfamilies
| subdivision =
{hide}hidden|headerstyle=background:#D3D3A4; text-size:90%;|Cladogram of subfamilies|{{clade|style=font-size:75%;line-height:75%
|1={{clade
|1=Martialinae
|2=Leptanillinae
|3=Amblyoponinae
|4=Paraponerinae
|5=Agroecomyrmecinae
|6=Ponerinae
|7=Proceratiinae
|8={{clade
|1={{clade
|1={{clade
|1=Ecitoninae‡
|2=Aenictinae‡
|3={{clade
|1=Dorylini‡
|2=Aenictogitoninae‡
{edih}
}}
|2=Cerapachyinae‡*|state2=double
|3=Leptanilloidinae‡
}}
|2={{clade
|1={{clade
|1={{clade
|1=Dolichoderinae
|2=Aneuretinae
}}
|2={{clade
|1=Pseudomyrmecinae
|2=Myrmeciinae
}}
}}
|2={{clade
|1={{clade
|1=Ectatomminae
|2=Heteroponerinae
}}
|2=Myrmicinae
|3=Formicinae
}}
}}
}}
}}
}}
A phylogeny of the extant ant subfamilies.JOURNAL, Ward, Philip S, Phylogeny, classification, and species-level taxonomy of ants (Hymenoptera: Formicidae), Zootaxa, 1668, 2007, 549–563,weblink PDF, JOURNAL, Rabeling C, Brown JM, Verhaagh M, Newly discovered sister lineage sheds light on early ant evolution, Proceedings of the National Academy of Sciences of the United States of America, 105, 39, 14913–7, September 2008, 18794530, 2567467, 10.1073/pnas.0806187105, 2008PNAS..10514913R, ‡ The previous dorylomorph subfamilies were synonymized under Dorylinae by Brady et al. in 2014JOURNAL, Brady SG, Fisher BL, Schultz TR, Ward PS, The rise of army ants and their relatives: diversification of specialized predatory doryline ants, BMC Evolutionary Biology, 14, 93, May 2014, 24886136, 4021219, 10.1186/1471-2148-14-93, }}}}Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the Cretaceous period, about 140 million years ago, and diversified after the rise of flowering plants. More than 12,500 of an estimated total of 22,000 species have been classified.WEB,weblinkweblink" title="web.archive.org/web/20160127143602weblink">weblink yes, 2016-01-27, Hymenoptera name server. Formicidae species count., Ohio State University, BOOK, Agosti D, Johnson NF, La nueva taxonomía de hormigas, 45–48, Fernández, F., Introducción a las hormigas de la región neotropical, Instituto Humboldt, Bogotá, 2003,weblink 2015-12-13, They are easily identified by their elbowed antennae and the distinctive node-like structure that forms their slender waists.Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies that may occupy large territories and consist of millions of individuals. Larger colonies consist of various castes of sterile, wingless females, most of which are workers (ergates), as well as soldiers (dinergates) and other specialised groups.BOOK, Fisher, Brian L., Bolton, Barry, vanc, Ants of Africa and Madagascar: A Guide to the Genera, 26 July 2016, University of California Press, 9780520290891, 24, BOOK, Singh, Rajendra, vanc, Elements of Entomology, 2006, Rastogi Publications, 9788171336777, 284, Nearly all ant colonies also have some fertile males called "drones" (aner) and one or more fertile females called "queens" (gynes). The colonies are described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.BOOK, Oster GF, Wilson EO, 1978, Caste and ecology in the social insects, Princeton University Press, Princeton, 21–22, 978-0-691-02361-8, BOOK, Flannery, Tim, vanc, A Natural History of the Planet, 2011, Grove/Atlantic, Inc., 978-0-8021-9560-9, 79, (File:Blackants-bredcrust-tokyo-may2015.webm|thumb|(video) Ants gathering food)(File:Ant in Slow Motion.webm|thumb|Ant in Slow Motion)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. 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.Hölldobler & Wilson (1990), p. 471Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study. Many human cultures make use of ants in cuisine, medication, and rituals. Some species are valued in their role as biological pest control agents. Their ability to exploit resources may bring ants into conflict with humans, however, as they can damage crops and invade buildings. Some species, such as the red imported fire ant (Solenopsis invicta), are regarded as invasive species, establishing themselves in areas where they have been introduced accidentally.

Etymology

The word ant and its chiefly dialectal form emmetemmet. Merriam-Webster Dictionary come from ', ' of Middle English, which come from ' of Old English, and these are all related to the dialectal Dutch ' and the Old High German ', from which comes the modern German '. All of these words come from West Germanic *, and the original meaning of the word was "the biter" (from Proto-Germanic ', "off, away" + ' "cut").WEB,weblink ant, Merriam-Webster Online Dictionary, 6 June 2008, WEB,weblink Ant. Online Etymology Dictionary, 30 May 2009, The family name Formicidae is derived from the Latin ' ("ant")BOOK, Simpson DP, Cassell's Latin Dictionary, Cassell, 1979, 5, London, 978-0-304-52257-6, from which the words in other Romance languages, such as the Portuguese ', Italian ', Spanish ', Romanian ', and French ' are derived. It has been hypothesised that a Proto-Indo-European word *morwi- was used, cf. Sanskrit vamrah, Latin formīca, Greek μύρμηξ mýrmēx, Old Church Slavonic mraviji, Old Irish moirb, Old Norse maurr, Dutch mier.WEB,weblink Formic, Etymonline.com, 2012-01-30,

Taxonomy and evolution

missing image!
- Ants in amber.jpg -
Ants fossilised in Baltic amber
{{cladogram|! Region !! Number ofspecies Hölldobler & Wilson (1990), p. 4
JOURNAL = CURRENT BIOLOGY ISSUE = 20 DATE = OCTOBER 2013 DOI = 10.1016/J.CUB.2013.08.050, 1996CBio....6.1213A, style=font-size:75%;line-height:75%
|label1=Aculeata
|1={{clade
|1=Chrysidoidea
|label2= 
|2={{clade
|label1= 
|1={{clade
|1=Vespidae
|2=Rhopalosomatidae
{edih}
|label2= 
|2={{clade
|label1= 
|1={{clade
|1=Pompilidae
|2=Tiphiidae
}}
|label2= 
|2={{clade
|1=Scolioidea
|label2= 
|2={{clade
|1=Apoidea
|2=Formicidae
}}
}}
}}
}}
}}
}}}}The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees, and wasps. Ants evolved from a lineage within the stinging wasps, and a 2013 study suggests that they are a sister group of the Apoidea. In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma) that lived in the Cretaceous period. The specimen, trapped in amber dating back to around 92 million years ago, has features found in some wasps, but not found in modern ants.JOURNAL, Wilson EO, Carpenter FM, Brown WL, The first mesozoic ants, Science, 157, 3792, 1038–40, September 1967, 17770424, 10.1126/science.157.3792.1038, E. O. Wilson, 1967Sci...157.1038W, Sphecomyrma was possibly a ground forager, while Haidomyrmex and Haidomyrmodes, related genera in subfamily Sphecomyrminae, are reconstructed as active arboreal predators.JOURNAL, Perrichot V, Nel A, Néraudeau D, Lacau S, Guyot T, New fossil ants in French Cretaceous amber (Hymenoptera: Formicidae), Die Naturwissenschaften, 95, 2, 91–7, February 2008, 17828384, 10.1007/s00114-007-0302-7, 2008NW.....95...91P, Older ants in the genus Sphecomyrmodes have been found in 99 million year-old amber from Myanmar.JOURNAL, Barden P, Grimaldi D, A diverse ant fauna from the mid-cretaceous of Myanmar (Hymenoptera: Formicidae), PLOS One, 9, 4, e93627, 2014, 24699881, 3974876, 10.1371/journal.pone.0093627, 2014PLoSO...993627B, JOURNAL, Perrichot V, Lacau S, Néraudeau D, Nel A, Fossil evidence for the early ant evolution, Die Naturwissenschaften, 95, 2, 85–90, February 2008, 17891532, 10.1007/s00114-007-0301-8,weblink 2008NW.....95...85P, A 2006 study suggested that ants arose tens of millions of years earlier than previously thought, up to 168 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.JOURNAL, Grimaldi D, Agosti D, A formicine in New Jersey cretaceous amber (Hymenoptera: formicidae) and early evolution of the ants, Proceedings of the National Academy of Sciences of the United States of America, 97, 25, 13678–83, December 2000, 11078527, 17635, 10.1073/pnas.240452097, 2000PNAS...9713678G, JOURNAL, Moreau CS, Bell CD, Vila R, Archibald SB, Pierce NE, Phylogeny of the ants: diversification in the age of angiosperms, Science, 312, 5770, 101–4, April 2006, 16601190, 10.1126/science.1124891, 2006Sci...312..101M, JOURNAL, Wilson EO, Hölldobler B, The rise of the ants: a phylogenetic and ecological explanation, Proceedings of the National Academy of Sciences of the United States of America, 102, 21, 7411–4, May 2005, 15899976, 1140440, 10.1073/pnas.0502264102, 2005PNAS..102.7411W, JOURNAL, LaPolla JS, Dlussky GM, Perrichot V, Ants and the fossil record, Annual Review of Entomology, 58, 609–30, 2013, 23317048, 10.1146/annurev-ento-120710-100600, Some groups, such as the Leptanillinae and Martialinae, are suggested to have diversified from early primitive ants that were likely to have been predators underneath the surface of the soil.JOURNAL, Barden P, Grimaldi D, 2012, Rediscovery of the bizarre Cretaceous ant Haidomyrmex Dlussky (Hymenoptera: Formicidae), with two new species, American Museum Novitates, 3755, 1–16, 10.1206/3755.2,weblink 3755, 2246/6368, During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian supercontinent (the Northern Hemisphere). They were scarce in comparison to the populations of other insects, representing only about 1% of the entire 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, around one in 10 genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene).Hölldobler & Wilson (1990), pp. 23–24Termites live in colonies and are sometimes called ‘white ants’, but termites are not ants. They are the sub-order Isoptera, and together with cockroaches they form the order Blattodea. Blattodeans are related to mantids, crickets, and other winged insects that do not undergo full metamorphosis. Like ants, termites are eusocial, with sterile workers, but they differ greatly in the genetics of reproduction. The similarity of their social structure to that of ants is attributed to convergent evolution.JOURNAL, Thorne, Barbara L, vanc, 1997, Evolution of eusociality in termites, Annu. Rev. Ecol. Syst., 28, 27–53,weblink PDF, 10.1146/annurev.ecolsys.28.1.27, 349550, yes,weblink" title="web.archive.org/web/20100530162505weblink">weblink 2010-05-30, Velvet ants look like large ants, but are wingless female wasps.WEB,weblink Order Isoptera – Termites, 12 June 2008, Iowa State University Entomology
archive-url=weblink" title="web.archive.org/web/20080615140449weblink">weblink deadurl= no, WEB,weblinkaccess-date=12 June 2008, Iowa State University Entomology archive-url=weblink" title="web.archive.org/web/20080630045231weblink">weblink deadurl= no,

Distribution and diversity{| class"wikitable sortable" style"float:right; margin:10px;"

Neotropic ecozone>Neotropics align="right"| 2162
Nearctic ecozone>Nearctic align="right"| 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.WEB,weblink Fantastic ants – Did you know?, National Geographic Magazine, Jones, Alice S archive-url=weblink" title="web.archive.org/web/20080730071158weblink">weblink deadurl= no, HTTP://WWW.HEAR.ORG/ANTS/ >TITLE=PEST ANTS IN HAWAII, 2007, Thomas, Philip, Hawaiian Ecosystems at Risk project (HEAR)ecological niches and exploit many different food resources as direct or indirect herbivores, predators and scavengers. Most ant species are omnivorous Generalist and specialist species>generalists, but a few are specialist feeders. Their ecological dominance is demonstrated by their biomass (ecology): ants are estimated to contribute 15–20 % (on average and nearly 25% in the tropics) of terrestrial animal biomass, exceeding that of the vertebrates.SCHULTZ TR JOURNAL = PROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICA ISSUE = 26 DATE = DECEMBER 2000 PMC = 34089 BIBCODE = 2000PNAS...9714028S, Ants range in size from {{convert|0.75|to(-)|52|mm|sigfig=2}},Hölldobler & Wilson (1990), p. 589BOOK, Shattuck SO, Australian ants: their biology and identification, 1999, CSIRO, Collingwood, Vic, 978-0-643-06659-5, 149, the largest species being the fossil Titanomyrma giganteum, the queen of which was {{convert|6|cm|in}} long with a wingspan of {{convert|15|cm|in}}.JOURNAL, Encyclopedia of Life Sciences, Schaal, Stephan, 27 January 2006, 10.1038/npg.els.0004143, Messel, 978-0-470-01617-6, 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 the potential existence of about 22,000) (see the article List of ant genera), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species.WEB, Agosti D, Johnson NF, 2005, Antbase,weblink American Museum of Natural History, 6 July 2008,weblink" title="web.archive.org/web/20080811000511weblink">weblink 11 August 2008, no, The relative ease with which ants may be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.BOOK, Agosti D, Majer JD, Alonso JE, Schultz TR, 2000, Ants: Standard methods for measuring and monitoring biodiversity, Smithsonian Institution, Smithsonian Institution Press,weblink 2015-12-13, WEB,weblinkweblink" title="web.archive.org/web/20160127143848weblink">weblink yes, 27 January 2016, Hymenoptera name server, Ohio State University, Johnson NF, 2007, 6 July 2008,

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 first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole may be formed by one or two nodes (the second alone, or the second and third abdominal segments).Borror, Triplehorn & Delong (1989), p. 737File:Bullant head detail.jpg|thumb|Bull antBull antLike 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 fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several missing image!
- Scheme ant worker anatomy-en.svg">thumb|450px|left|Diagram of a worker ant (Pachycondyla verenae)

Head

An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ant eyes are good for acute movement detection, but do not offer a high resolution image. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization.JOURNAL, Fent K, Wehner R, Oceili: a celestial compass in the desert ant cataglyphis, Science, 228, 4696, 192–4, April 1985, 17779641, 10.1126/science.228.4696.192, 1985Sci...228..192F, Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. However, some ants, such as Australia's bulldog ant, have excellent vision and are capable of discriminating the distance and size of objects moving nearly a metre away.JOURNAL, J. Exp. Biol., 119, 115–131, 1985, Attack behaviour and distance perception in the Australian bulldog ant Myrmecia nigriceps, Eriksson, E. Sture, vanc,weblink 1, Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents, and vibrations; they also are used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species, a small pocket (infrabuccal chamber) inside the mouth stores food, so it may be passed to other ants or their larvae.JOURNAL, Eisner T, Happ GM, The infrabuccal pocket of a formicine ant: a social filtration device, Psyche, 69, 107–116, 1962,weblink 10.1155/1962/25068, 3,

Mesosoma

Both the legs and wings of the ant are attached to the mesosoma ("thorax"). The legs terminate in a hooked claw which allows them to hook on and climb surfaces.WEB, Holbrook, Tate, Ask a Biologist: Face to Face with Ants, ASU School of Life Sciences,weblink 2018-01-23, Only reproductive ants, queens, and males, have wings. Queens shed their wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. In a few species, wingless queens (ergatoids) and males occur.

Metasoma

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.

Polymorphism

Atta.cephalotes.gamut.selection.jpg
-
In the colonies of a few ant species, there are physical castes—workers in distinct size-classes, called minor, median, and major ergates. Often, the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. These are known as macrergates while smaller workers are known as micrergates. Although formally known as dinergates, such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they still are workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species, the median workers are absent, creating a sharp divide between the minors and majors.JOURNAL, Wilson EO, The origin and evolution of polymorphism in ants, The Quarterly Review of Biology, 28, 2, 136–56, June 1953, 13074471, 10.1086/399512, Weaver ants, for example, have a distinct bimodal size distribution.JOURNAL, Weber, NA, 1946, Dimorphism in the African Oecophylla worker and an anomaly (Hym.: Formicidae), Annals of the Entomological Society of America, 39, 7–10,weblink PDF, JOURNAL, Edward O., Wilson, Robert W., Taylor, vanc, 1964, A Fossil Ant Colony: New Evidence of Social Antiquity, Psyche, 71, 93–103,weblink PDF, 10.1155/1964/17612, 2, 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.JOURNAL, Moffett MW, Tobin JE, 1991, Physical castes in ant workers: a problem for Daceton armigerum and other ants, Psyche, 98, 283–292,weblinkweblink" title="web.archive.org/web/20080227015919weblink">weblink 2008-02-27, PDF, 10.1155/1991/30265, 4, 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.JOURNAL, Børgesen LW, 2000, Nutritional function of replete workers in the pharaoh's ant, Monomorium pharaonis (L.), Insectes Sociaux, 47, 2, 141–146, 10.1007/PL00001692, For instance, these replete workers develop in the North American honeypot ant Myrmecocystus mexicanus. Usually the largest workers in the colony develop into repletes; and, if repletes are removed from the colony, other workers become repletes, demonstrating the flexibility of this particular polymorphism.JOURNAL, Rissing, Steven W, vanc, 1984, Replete Caste Production and Allometry of Workers in the Honey Ant, Myrmecocystus mexicanus Wesmael (Hymenoptera: Formicidae), Journal of the Kansas Entomological Society, 57, 2, 347–350, This polymorphism in morphology and behaviour of workers initially was thought to be determined by environmental factors such as nutrition and hormones that led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp.JOURNAL, Hughes WO, Sumner S, Van Borm S, Boomsma JJ, Worker caste polymorphism has a genetic basis in Acromyrmex leaf-cutting ants, Proceedings of the National Academy of Sciences of the United States of America, 100, 16, 9394–7, August 2003, 12878720, 170929, 10.1073/pnas.1633701100, 2003PNAS..100.9394H, 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.JOURNAL, Ross KG, Krieger MJ, Shoemaker DD, Alternative genetic foundations for a key social polymorphism in fire ants, Genetics, 165, 4, 1853–67, December 2003, 14704171, 1462884, 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.JOURNAL, Crosland MW, Crozier RH, Myrmecia pilosula, an Ant with Only One Pair of Chromosomes, Science, 231, 4743, 1278, March 1986, 17839565, 10.1126/science.231.4743.1278, 1986Sci...231.1278C, JOURNAL, Tsutsui ND, Suarez AV, Spagna JC, Johnston JS, The evolution of genome size in ants, BMC Evolutionary Biology, 8, 64, 64, February 2008, 18302783, 2268675, 10.1186/1471-2148-8-64,weblinkweblink" title="web.archive.org/web/20080627234811weblink">weblink no, 27 June 2008,

Life cycle

missing image!
- Meat eater ant nest swarming02.jpg -
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female diploid; if not, it will be male haploid. Ants develop by complete metamorphosis with the larva stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach". Larvae, especially in the later stages, may also be provided solid food, such as trophic eggs, pieces of prey, and seeds brought by workers.The larvae grow through a series of four or five moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa.BOOK, Gillott, Cedric, 1995, Entomology, Springer, 978-0-306-44967-3, 325, The differentiation into queens and workers (which are both female), and different castes of workers, is influenced in some species by the nutrition the larvae obtain. Genetic influences and the control of gene expression by the developmental environment are complex and the determination of caste continues to be a subject of research.JOURNAL, The causes and consequences of genetic caste determination in ants (Hymenoptera: Formicidae), Kirk E., Anderson, Timothy A., Linksvayer, Chris R., Smith, vanc, Myrmecol. News, 11, 119–132, 2008,weblink Winged male ants, called drones, emerge from pupae along with the usually winged breeding females. Some species, such as army ants, have wingless queens. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so often, are moved around among the various brood chambers within the colony.Hölldobler & Wilson (1990), pp. 351, 372A new ergate spends the first few days of its adult life caring for the queen and young. She then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants who are older and are likely to die soon of natural causes.JOURNAL, Annual Review of Entomology, 1989, 34, 191–210, Foraging strategies of ants, Traniello JFA, 10.1146/annurev.en.34.010189.001203, JOURNAL, Sorensen A, Busch TM, Vinson SB, Behavioral flexibility of temporal sub-castes in the fire ant, Solenopsis invicta, in response to food, Psyche, 91, 319–332, 1984,weblink 10.1155/1984/39236, 3–4, Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, being quite short-lived and surviving for only a few weeks.JOURNAL, Keller L, 1998, Queen lifespan and colony characteristics in ants and termites, Insectes Sociaux, 45, 235–246, 10.1007/s000400050084, 3, Ant queens are estimated to live 100 times as long as solitary insects of a similar size.BOOK, Franks NR, Resh VH, Cardé RT, 2003, Encyclopedia of Insects, 29–32, 978-0-12-586990-4, Academic Press, San Diego, Ants are active all year long in the tropics, but, in cooler regions, they survive the winter in a state of dormancy known as hibernation. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.JOURNAL, Kipyatkov VE, 2001, Seasonal life cycles and the forms of dormancy in ants (Hymenoptera, Formicoidea), Acta Societatis Zoologicae Bohemicae, 65, 2, 198–217,

Reproduction

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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.JOURNAL, Heinze, Jurgen, Tsuji, Kazuki, vanc, 1995, Ant reproductive strategies, Res. Popul. Ecol., 37, 2, 135–149,weblink PDF, 10.1007/BF02515814, Secretions from the male accessory glands in some species can plug the female genital opening and prevent females from re-mating.JOURNAL, 10.1007/s00040-003-0697-x, Evidence for mating plugs in the fire ant Solenopsis invicta, Insectes Sociaux, 50, 4, 401–402, 2003, Mikheyev AS, Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens, while others may exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right.JOURNAL, Peeters C, Hölldobler B, Reproductive cooperation between queens and their mated workers: the complex life history of an ant with a valuable nest, Proceedings of the National Academy of Sciences of the United States of America, 92, 24, 10977–9, November 1995, 11607589, 40553, 10.1073/pnas.92.24.10977, 1995PNAS...9210977P, Drones can also mate with existing queens by entering a foreign colony. When the drone is initially attacked by the workers, it releases a mating pheromone. If recognized as a mate, it will be carried to the queen to mate.JOURNAL, Franks NR, Hölldobler B, 10.1111/j.1095-8312.1987.tb00298.x, Sexual competition during colony reproduction in army ants, Biological Journal of the Linnean Society, 30, 3, 229–243, 1987, Males may also patrol the nest and fight others by grabbing them with their mandibles, piercing their exoskeleton and then marking them with a pheromone. The marked male is interpreted as an invader by worker ants and is killed.JOURNAL, 10.1007/BF01175395, Pheromonal manipulation of workers by a fighting male to kill his rival males in the ant Cardiocondyla wroughtonii, Naturwissenschaften, 79, 6, 274–276, 1992, Yamauchi K, Kawase N, 1992NW.....79..274Y,
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Fertilised meat-eater ant queen beginning to dig a new colony
Most ants are univoltine, producing a new generation each year.BOOK, Taylor, Richard W., vanc, 2007, Bloody funny wasps! Speculations on the evolution of eusociality in ants, 580–609, Snelling RR, Fisher BL, Ward PS, Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson â€“ 50 years of contributions. Memoirs of the American Entomological Institute, 80, American Entomological Institute,weblink PDF, 2015-12-13, During the species-specific breeding period, new reproductives, females, and winged males leave the colony in what is called a nuptial flight. The nuptial flight usually takes place in the late spring or early summer when the weather is hot and humid. Heat makes flying easier and freshly fallen rain makes the ground softer for mated queens to dig nests.JOURNAL, Wilson EO, 1957, The organization of a nuptial flight of the ant Pheidole sitarches Wheeler, Psyche,weblink 64, 2, 46–50, 10.1155/1957/68319, Males typically take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Males will mount females in the air, but the actual mating process usually takes place on the ground. Females of some species mate with just one male but in others they may mate with as many as ten or more different males, storing the sperm in their spermathecae.Hölldobler & Wilson (1990), pp. 143–179Mated 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 can selectively fertilise future eggs with the sperm stored to produce diploid workers or lay unfertilized haploid eggs to produce drones. The first workers to hatch are known as nanitics,BOOK, Sudd JH, Franks NR, The Behavioural Ecology of Ants, 9 March 2013, Springer Science & Business Media, 9789400931237, 41, and are weaker and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food, and care for the other eggs. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, a process akin to swarming in honeybees.

Behaviour and ecology

Communication

File:Camponotus sericeus Senegal.jpg|thumb|Two Camponotus sericeusCamponotus sericeusAnts communicate with each other using pheromones, sounds, and touch.JOURNAL, Jackson DE, Ratnieks FL, Communication in ants, Current Biology, 16, 15, R570–4, August 2006, 16890508, 10.1016/j.cub.2006.07.015,weblink 1996CBio....6.1213A, The use of pheromones as chemical signals is more developed in ants, such as the red harvester ant, 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 may be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually identifying the best path.JOURNAL, Goss S, Aron S, Deneubourg JL, Pasteels JM, 1989, Self-organized shortcuts in the Argentine ant, Naturwissenschaften, 76, 579–581, 10.1007/BF00462870, 12,weblink 1989NW.....76..579G, Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from farther away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves.JOURNAL, D'Ettorre P, Heinze J, Acta Ethologica, 2001, 3, 67–82, Sociobiology of slave-making ants,weblink 10.1007/s102110100038, 2, 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. Pheromones also are exchanged, mixed with food, and passed by trophallaxis, transferring information within the colony.BOOK, Detrain C, Deneubourg JL, Pasteels JM, 1999, Information processing in social insects, Birkhäuser, 978-3-7643-5792-4, 224–227, This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to.JOURNAL, Greene MJ, Gordon DM, Structural complexity of chemical recognition cues affects the perception of group membership in the ants Linephithema humile and Aphaenogaster cockerelli, The Journal of Experimental Biology, 210, Pt 5, 897–905, March 2007, 17297148, 10.1242/jeb.02706, In ant species with queen castes, when the dominant queen stops producing a specific pheromone, workers begin to raise new queens in the colony.Hölldobler & Wilson (1990), p. 354Some 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.JOURNAL, Hickling R, Brown RL, Analysis of acoustic communication by ants, The Journal of the Acoustical Society of America, 108, 4, 1920–9, October 2000, 11051518, 10.1121/1.1290515, 2000ASAJ..108.1920H, JOURNAL, 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, 293–299, 10.1007/s002650050292, 5,

Defence

{{see also|Defense in insects|l1=Insect defences}}File:Plectroctena sp ants.jpg|thumb|A PlectroctenaPlectroctenaAnts attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals, such as formic acid in the case of formicine ants, alkaloids and piperidines in fire 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.The sting of jack jumper ants can be fatal,JOURNAL, Clarke PS, The natural history of sensitivity to jack jumper ants (Hymenoptera formicidae Myrmecia pilosula) in Tasmania, The Medical Journal of Australia, 145, 11–12, 564–6, 1986, 3796365, and an antivenom has been developed for it.JOURNAL, Brown SG, Heddle RJ, Wiese MD, Blackman KE, Efficacy of ant venom immunotherapy and whole body extracts, The Journal of Allergy and Clinical Immunology, 116, 2, 464–5; author reply 465–6, August 2005, 16083810, 10.1016/j.jaci.2005.04.025, Fire ants, Solenopsis spp., are unique in having a venom sac containing piperidine alkaloids.JOURNAL, Obin MS, Vander Meer RK, Gaster flagging by fire ants (Solenopsis spp.): Functional significance of venom dispersal behavior, Journal of Chemical Ecology, 11, 12, 1757–68, December 1985, 24311339, 10.1007/BF01012125, Their stings are painful and can be dangerous to hypersensitive people.JOURNAL, Stafford CT, Hypersensitivity to fire ant venom, Annals of Allergy, Asthma & Immunology, 77, 2, 87–95; quiz 96–9, August 1996, 8760773, 10.1016/S1081-1206(10)63493-X,
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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.JOURNAL, Patek SN, Baio JE, Fisher BL, Suarez AV, Multifunctionality and mechanical origins: ballistic jaw propulsion in trap-jaw ants, Proceedings of the National Academy of Sciences of the United States of America, 103, 34, 12787–92, August 2006, 16924120, 1568925, 10.1073/pnas.0604290103, 2006PNAS..10312787P, One study of Odontomachus bauri recorded peak speeds of between {{convert|126|and|230|km/h|mph|abbr=on}}, with the jaws closing within 130 microseconds on average.The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backward to escape a threat. Before striking, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory organs resembling hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws also are seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe,JOURNAL, The trap-jaw mechanism in the Dacetine ant Daceton armigerum and Strumigenys sp, Gronenberg W, 1996, The Journal of Experimental Biology, 199, 9, 2021–2033,weblink PDF, which are viewed as examples of convergent evolution.A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. If combat takes a turn for the worse, a worker may perform a final act of suicidal altruism by rupturing the membrane of its gaster, causing the content of its mandibular glands to burst from the anterior region of its head, spraying a poisonous, corrosive secretion containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies.JOURNAL, Jones TH, Clark DA, Edwards AA, Davidson DW, Spande TF, Snelling RR, The chemistry of exploding ants, Camponotus spp. (cylindricus complex), Journal of Chemical Ecology, 30, 8, 1479–92, August 2004, 15537154, 10.1023/B:JOEC.0000042063.01424.28, 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.JOURNAL, Tofilski A, Couvillon MJ, Evison SE, Helanterä H, Robinson EJ, Ratnieks FL, Preemptive defensive self-sacrifice by ant workers, The American Naturalist, 172, 5, E239–43, November 2008, 18928332, 10.1086/591688,weblink PDF,
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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 (wikt:undertaker|undertaking) or necrophory, the disposal of dead nest-mates.JOURNAL, Julian GE, Cahan S, Undertaking specialization in the desert leaf-cutter ant Acromyrmex versicolor, Animal Behaviour, 58, 2, 437–442, August 1999, 10458895, 10.1006/anbe.1999.1184, Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicanaJOURNAL, 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, 10.1111/j.1365-3032.2006.00526.x, while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates to trigger similar behaviour.JOURNAL, Choe DH, Millar JG, Rust MK, Chemical signals associated with life inhibit necrophoresis in Argentine ants, Proceedings of the National Academy of Sciences of the United States of America, 106, 20, 8251–5, May 2009, 19416815, 2688878, 10.1073/pnas.0901270106, 2009PNAS..106.8251C, Nests may be protected from physical threats such as flooding and overheating by elaborate nest architecture.JOURNAL, Tschinkel WR, The nest architecture of the Florida harvester ant, Pogonomyrmex badius, Journal of Insect Science, 4, 21, 21, 2004, 15861237, 528881, JOURNAL, 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, 211–218, 10.1007/BF01240479, 2, 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.JOURNAL, Maschwitz U, Moog J, Communal peeing: a new mode of flood control in ants, Die Naturwissenschaften, 87, 12, 563–5, December 2000, 11198200, 10.1007/s001140050780, 2000NW.....87..563M, Camponotus anderseni, which nests in the cavities of wood in mangrove habitats, deals with submergence under water by switching to anaerobic respiration.JOURNAL, Nielsen MG, Christian KA, The mangrove ant, Camponotus anderseni, switches to anaerobic respiration in response to elevated CO2 levels, Journal of Insect Physiology, 53, 5, 505–8, May 2007, 17382956, 10.1016/j.jinsphys.2007.02.002,

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 knowledgeable forager of Temnothorax albipennis will lead a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. The leader is acutely sensitive to the progress of the follower and slows down when the follower lags and speeds up when the follower gets too close.JOURNAL, Franks NR, Richardson T, Teaching in tandem-running ants, Nature, 439, 7073, 153, January 2006, 16407943, 10.1038/439153a, 2006Natur.439..153F, Controlled experiments with colonies of Cerapachys biroi suggest that an individual may choose nest roles based on her previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out fewer and fewer times. A month later, the successful foragers continued in their role while the others had moved to specialise in brood care.JOURNAL, Ravary F, Lecoutey E, Kaminski G, Châline N, Jaisson P, Individual experience alone can generate lasting division of labor in ants, Current Biology, 17, 15, 1308–12, August 2007, 17629482, 10.1016/j.cub.2007.06.047, 1996CBio....6.1213A,

Nest construction

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Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ant species, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests may be found in the ground, under stones or logs, inside logs, hollow stems, or even acorns. The materials used for construction include soil and plant matter, 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.JOURNAL, Franks NR, Hooper J, Webb C, Dornhaus A, Tomb evaders: house-hunting hygiene in ants, Biology Letters, 1, 2, 190–2, June 2005, 17148163, 1626204, 10.1098/rsbl.2005.0302, 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.Hölldobler & Wilson (1990), p. 573Weaver 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.JOURNAL, 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, 10.1111/j.1440-6055.2005.00462.x, 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.JOURNAL, Frouz J, The Effect of Nest Moisture on Daily Temperature Regime in the Nests of Formica polyctena Wood Ants, Insectes Sociaux, 47, 3, 2000, 229–235, 10.1007/PL00001708, Some ant species, such as those that use natural cavities, can be opportunistic and make use of the controlled micro-climate provided inside human dwellings and other artificial structures to house their colonies and nest structures.JOURNAL, Biological Conservation, 115, 2, 2004, 279–289, Impact of human dwellings on the distribution of the exotic Argentine ant: a case study in the Doñana National Park, Spain, Soledad, Carpintero, Joaquin, Reyes-López, Luis Arias, de Reynac, vanc, 10.1016/S0006-3207(03)00147-2, JOURNAL, Nest-site Limitation and Nesting Resources of Ants (Hymenoptera: Formicidae) in Urban Green Spaces, Russell, Friedrich, Stacy M., Philpott, vanc, Environmental Entomology, 38, 3, 600–607, 2009, 10.1603/022.038.0311,

Cultivation of food

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Myrmecocystus, honeypot ants, store food to prevent colony famine
Most ants are generalist predators, scavengers, and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. It is believed that many ant species that engage in indirect herbivory rely on specialized symbiosis with their gut microbes JOURNAL, Anderson KE, Russell JA, Moreau CS, Kautz S, Sullam KE, Hu Y, Basinger U, Mott BM, Buck N, Wheeler DE, Highly similar microbial communities are shared among related and trophically similar ant species, Molecular Ecology, 21, 9, 2282–96, May 2012, 22276952, 10.1111/j.1365-294x.2011.05464.x, to upgrade the nutritional value of the food they collect JOURNAL, Feldhaar H, Straka J, Krischke M, Berthold K, Stoll S, Mueller MJ, Gross R, Nutritional upgrading for omnivorous carpenter ants by the endosymbiont Blochmannia, BMC Biology, 5, 48, October 2007, 17971224, 2206011, 10.1186/1741-7007-5-48, and allow them to survive in nitrogen poor regions, such as rainforest canopies.JOURNAL, Russell JA, Moreau CS, Goldman-Huertas B, Fujiwara M, Lohman DJ, Pierce NE, Bacterial gut symbionts are tightly linked with the evolution of herbivory in ants, Proceedings of the National Academy of Sciences of the United States of America, 106, 50, 21236–41, December 2009, 19948964, 2785723, 10.1073/pnas.0907926106, 2009PNAS..10621236R, Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Ergates specialise in related tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is found to be toxic to the fungus, the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria introduced into the nest that may harm the fungi.JOURNAL, Nature, 1999, 398, Ants, plants and antibiotics, Schultz TR, 747–748, 10.1038/19619, 6730,weblink 1999Natur.398..747S,

Navigation

File:Ant trail.jpg|thumb|An ant trailtrailForaging ants travel distances of up to {{convert|200|m|sigfig=1}} from their nest JOURNAL, Ecology of foraging by ants, Carrol CR, Janzen DH, Annual Review of Ecology and Systematics, 1973, 4, 231–257, 10.1146/annurev.es.04.110173.001311, and scent trails allow them to find their way back even in the dark. In hot and arid regions, day-foraging ants face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants of the genus Cataglyphis such as the Sahara desert ant navigate by keeping track of direction as well as distance travelled. Distances travelled are measured using an internal pedometer that keeps count of the steps takenJOURNAL, Wittlinger M, Wehner R, Wolf H, The ant odometer: stepping on stilts and stumps, Science, 312, 5782, 1965–7, June 2006, 16809544, 10.1126/science.1126912,weblinkweblink" title="web.archive.org/web/20110728042723weblink">weblink yes, 2006Sci...312.1965W, 2011-07-28, and also by evaluating the movement of objects in their visual field (optical flow).JOURNAL, Desert ants Cataglyphis fortis use self-induced optic flow to measure distances travelled, Ronacher B, Werner R, Journal of Comparative Physiology A, 1995,weblink 10.1007/BF00243395, 177, Directions are measured using the position of the sun.JOURNAL, Wehner R, Desert ant navigation: how miniature brains solve complex tasks, Journal of Comparative Physiology A, 189, 8, 579–88, August 2003, 12879352, 10.1007/s00359-003-0431-1,weblink They integrate this information to find the shortest route back to their nest.JOURNAL, Sommer S, Wehner R, The ant's estimation of distance travelled: experiments with desert ants, Cataglyphis fortis, Journal of Comparative Physiology A, 190, 1, 1–6, January 2004, 14614570, 10.1007/s00359-003-0465-4,weblink Like all ants, they can also make use of visual landmarks when availableJOURNAL, Visual navigation in desert ants Cataglyphis fortis: are snapshots coupled to a celestial system of reference?, Åkesson S, Wehner R, Journal of Experimental Biology, 2002, 205, 1971–1978,weblink PDF, 14, as well as olfactory and tactile cues to navigate.JOURNAL, Steck K, Hansson BS, Knaden M, Smells like home: Desert ants, Cataglyphis fortis, use olfactory landmarks to pinpoint the nest, Frontiers in Zoology, 6, 5, February 2009, 19250516, 2651142, 10.1186/1742-9994-6-5,weblink PDF, JOURNAL, Seidl T, Wehner R, Visual and tactile learning of ground structures in desert ants, The Journal of Experimental Biology, 209, Pt 17, 3336–44, September 2006, 16916970, 10.1242/jeb.02364,weblink PDF, Some species of ant are able to use the Earth's magnetic field for navigation.JOURNAL
, Orientation by magnetic field in leaf-cutter ants, ''Atta colombica (Hymenoptera: Formicidae)
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, The compound eyes of ants have specialised cells that detect polarised light from the Sun, which is used to determine direction.JOURNAL, Fukushi T, Homing in wood ants, Formica japonica: use of the skyline panorama, The Journal of Experimental Biology, 204, Pt 12, 2063–72, June 2001, 11441048,weblink JOURNAL, Wehner R, Menzel R, Homing in the ant Cataglyphis bicolor, Science, 164, 3876, 192–4, April 1969, 5774195, 10.1126/science.164.3876.192, 1969Sci...164..192W, These polarization detectors are sensitive in the ultraviolet region of the light spectrum.BOOK, The Insects: Structure and Function, Reginald Frederick, Chapman, vanc, 1998, 4, Cambridge University Press, 978-0-521-57890-5, 600, In some army ant species, a group of foragers who become separated from the main column may sometimes turn back on themselves and form a circular ant mill. The workers may then run around continuously until they die of exhaustion.JOURNAL, Delsuc F, Army ants trapped by their evolutionary history, PLoS Biology, 1, 2, E37, November 2003, 14624241, 261877, 10.1371/journal.pbio.0000037,

Locomotion

The female worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs.JOURNAL, 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, 10.1007/BF01992052, 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.JOURNAL, Yanoviak SP, Dudley R, Kaspari M, Directed aerial descent in canopy ants, Nature, 433, 7026, 624–6, February 2005, 15703745, 10.1038/nature03254,weblink yes, PDF, 2005Natur.433..624Y,weblink" title="wayback.archive-it.org/all/20070616090223weblink">weblink 2007-06-16, 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.JOURNAL, Mlot NJ, Tovey CA, Hu DL, Fire ants self-assemble into waterproof rafts to survive floods, Proceedings of the National Academy of Sciences of the United States of America, 108, 19, 7669–73, May 2011, 21518911, 3093451, 10.1073/pnas.1016658108, 2011PNAS..108.7669M, These rafts may also have a role in allowing ants to colonise islands.JOURNAL, Morrison LW, 1998, A review of Bahamian ant (Hymenoptera: Formicidae) biogeography, Journal of Biogeography, 25, 3, 561–571, 10.1046/j.1365-2699.1998.2530561.x, Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they go to trapped pockets of air in the submerged nests to breathe.JOURNAL, 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, 521–533, 10.1071/ZO9960521, 5,

Cooperation and competition

File:Ants eating cicada, jjron 22.11.2009.jpg|thumb|right|Meat-eater ants feeding on a (cicada]]: social ants cooperate and collectively gather food)Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal of ants. Like virtually all ants, they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using her large eyes instead of chemical senses to find prey.JOURNAL, 10.1111/j.1440-6055.1988.tb01179.x, Aspects of the biology of the primitive ant genus Myrmecia F. (Hymenoptera: Formicidae), 1988, Australian Journal of Entomology, 27, 305–309, Crosland MW, Crozier RH, Jefferson E, 4, 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 or slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive.JOURNAL, 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, 10.1590/S1519-69842005000300008, Captured workers of enslaved Temnothorax species have evolved a counter strategy, destroying just the female pupae of the slave-making Temnothorax americanus, but sparing the males (who don't take part in slave-raiding as adults).JOURNAL, Achenbach A, Foitzik S, First evidence for slave rebellion: enslaved ant workers systematically kill the brood of their social parasite protomognathus americanus, Evolution; International Journal of Organic Evolution, 63, 4, 1068–75, April 2009, 19243573, 10.1111/j.1558-5646.2009.00591.x, See also New Scientist, April 9, 2009
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upright|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.JOURNAL, Henderson G, Andersen JF, Phillips JK, Jeanne RL, Internest aggression and identification of possible nestmate discrimination pheromones in polygynous antFormica montana, Journal of Chemical Ecology, 16, 7, 2217–28, July 1990, 24264088, 10.1007/BF01026932, Also, the reason why two separate colonies of ants will attack each other even if they are of the same species is because the genes responsible for pheromone production are different between them. The Argentine ant, however, does not have this characteristic, due to lack of genetic diversity, and has become a global pest because of it.Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species such as Strumigenys xenos are entirely parasitic and do not have workers, but instead, rely on the food gathered by their Strumigenys perplexa hosts.JOURNAL, 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, 253–263,weblink 10.1046/j.1365-3113.1996.d01-12.x, 3, JOURNAL, Taylor RW, 1968, The Australian workerless inquiline ant, Strumigenys xenos Brown (Hymenoptera-Formicidae) recorded from New Zealand, New Zealand Entomologist, 4, 1, 47–49,weblink 10.1080/00779962.1968.9722888, This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen may enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest.Hölldobler & Wilson (1990), pp. 436—448A conflict between the sexes of a species is seen in some species of ants with these reproducers apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process whereby a diploid egg loses its maternal contribution to produce haploid males who are clones of the father.JOURNAL, Fournier D, Estoup A, Orivel J, Foucaud J, Jourdan H, Le Breton J, Keller L, Clonal reproduction by males and females in the little fire ant, Nature, 435, 7046, 1230–4, June 2005, 15988525, 10.1038/nature03705, 2005Natur.435.1230F,

Relationships with other organisms

File:Myrmarachne plataleoides female thailand.jpg|thumb|The spider Myrmarachne plataleoides (female shown) mimics weaver antweaver antAnts form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They also are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae, and eggs, consuming the food stores of the ants, or avoiding predators. These inquilines may bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.JOURNAL, Ant-mimicry in Panamanian clubionid and salticid spiders (Araneae: Clubionidae, Salticidae), Reiskind J, Biotropica, 9, 1, 1977, 1–8, 10.2307/2387854, 2387854, JOURNAL, Myrmecomorphy and myrmecophily in spiders: A Review, Cushing PE, The Florida Entomologist, 80, 2, 1997, 165–193, 10.2307/3495552,weblink PDF, 3495552,
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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.JOURNAL, Styrsky JD, Eubanks MD, Ecological consequences of interactions between ants and honeydew-producing insects, Proceedings. Biological Sciences, 274, 1607, 151–64, January 2007, 17148245, 1685857, 10.1098/rspb.2006.3701,weblink In some cases, the aphids secrete the honeydew in response to ants tapping them with their antennae. The ants in turn keep predators away from the aphids and will move them from one feeding location to another. When migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs may become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.JOURNAL, 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, Myrmecophilous (ant-loving) caterpillars of the butterfly family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants.JOURNAL, DeVries PJ, 1992, Singing caterpillars, ants and symbiosis, Scientific American, 267, 76–82, 10.1038/scientificamerican1092-76, 4, 1992SciAm.267d..76D, A similar adaptation can be seen in Grizzled skipper butterflies that emit vibrations by expanding their wings in order to communicate with ants, which are natural predators of these butterflies.JOURNAL, Elfferich, Nico W., vanc, 1998, Is the larval and imaginal signalling of Lycaenidae and other Lepidoptera related to communication with ants,weblink Deinsea, 4, 1, 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.JOURNAL, Pierce NE, Braby MF, Heath A, Lohman DJ, Mathew J, Rand DB, Travassos MA, The ecology and evolution of ant association in the Lycaenidae (Lepidoptera), Annual Review of Entomology, 47, 733–71, 2002, 11729090, 10.1146/annurev.ento.47.091201.145257, A number of specialized bacterial have been found as endosymbionts in ant guts. Some of the dominant bacteria belong to the order Rhizobiales whose members are known for being nitrogen-fixing symbionts in legumes but the species found in ant lack the ability to fix nitrogen.JOURNAL, Kautz S, Rubin BE, Russell JA, Moreau CS, Surveying the microbiome of ants: comparing 454 pyrosequencing with traditional methods to uncover bacterial diversity, Applied and Environmental Microbiology, 79, 2, 525–34, January 2013, 23124239, 3553759, 10.1128/AEM.03107-12, JOURNAL, Neuvonen MM, Tamarit D, Näslund K, Liebig J, Feldhaar H, Moran NA, Guy L, Andersson SG, The genome of Rhizobiales bacteria in predatory ants reveals urease gene functions but no genes for nitrogen fixation, En, Scientific Reports, 6, 1, 39197, December 2016, 27976703, 5156944, 10.1038/srep39197, 2016NatSR...639197N, Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the genera Leucoagaricus or Leucocoprinus of the family Agaricaceae. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant, Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.JOURNAL, Dejean A, Solano PJ, Ayroles J, Corbara B, Orivel J, Insect behaviour: arboreal ants build traps to capture prey, Nature, 434, 7036, 973, April 2005, 15846335, 10.1038/434973a, 2005Natur.434..973D, File:Ants on a dandelion.jpg|thumb|Ants may obtain nectar from flowers such as the dandeliondandelionLemon 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.JOURNAL, Frederickson ME, Gordon DM, The devil to pay: a cost of mutualism with Myrmelachista schumanni ants in 'devil's gardens' is increased herbivory on Duroia hirsuta trees, Proceedings. Biological Sciences, 274, 1613, 1117–23, April 2007, 17301016, 2124481, 10.1098/rspb.2006.0415, Although some ants obtain nectar from flowers, pollination by ants is somewhat rare.JOURNAL, Ant Inhibition of Pollen Function: A Possible Reason Why Ant Pollination is Rare, Andrew J., Beattie, Turnbull, Christine, Knox, R. B., Williams, E. G., vanc, American Journal of Botany, 71, 3, 1984, 421–426, 10.2307/2443499, 2443499, 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.JOURNAL, Katayama N, Suzuki N, Role of extrafloral nectaries of Vicia faba in attraction of ants and herbivore exclusion by ants, 2004, Entomological Science, 119–124, 7, 2, 10.1111/j.1479-8298.2004.00057.x, Species such as the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) who defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants.JOURNAL, Do ants feed plants? A 15N labelling study of nitrogen fluxes from ants to plants in the mutualism of Pheidole and Piper, Fischer RC, Wanek W, Richter A, Mayer V, 2003, Journal of Ecology, 91, 126–134, 10.1046/j.1365-2745.2003.00747.x, 2009JEcol..98...74B, In return, the ants obtain food from protein- and lipid-rich Beltian bodies. In Fiji Philidris nagasau (Dolichoderinae) are known to selectively grow species of epiphytic Squamellaria (Rubiaceae) which produce large domatia inside which the ant colonies nest. The ants plant the seeds and the domatia of young seedling are immediately occupied and the ant faeces in them contribute to rapid growth.JOURNAL, Chomicki G, Renner SS, Obligate plant farming by a specialized ant, Nature Plants, 2, 12, 16181, November 2016, 27869787, 10.1038/nplants.2016.181, Similar dispersal associations are found with other dolichoderines in the region as well.JOURNAL, Chomicki G, Janda M, Renner SS, The assembly of ant-farmed gardens: mutualism specialization following host broadening, Proceedings. Biological Sciences, 284, 1850, 20161759, March 2017, 28298344, 5360912, 10.1098/rspb.2016.1759, Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.JOURNAL, Fiala B, Maschwitz U, Pong TY, Helbig AJ, Studies of a South East Asian ant-plant association: protection of Macaranga trees by Crematogaster borneensis, Oecologia, 79, 4, 463–470, June 1989, 28313479, 10.1007/bf00378662, 1989Oecol..79..463F,weblink Submitted manuscript, Many tropical tree species have seeds that are dispersed by ants.JOURNAL, Hanzawa FM, Beattie AJ, Culver DC, 1988, Directed dispersal: demographic analysis of an ant-seed mutualism, American Naturalist, 131, 1, 1–13, 10.1086/284769, Seed dispersal by ants or myrmecochory is widespread and new estimates suggest that nearly 9% of all plant species may have such ant associations.JOURNAL, Giladi I, Choosing benefits or partners: a review of the evidence for the evolution of myrmecochory, Oikos, 112, 3, 2006, 481–492, 10.1111/j.0030-1299.2006.14258.x, 10.1.1.530.1306, JOURNAL, Convergent evolution of seed dispersal by ants, and phylogeny and biogeography in flowering plants: A global survey, Szabolcs, Lengyel, Aaron D., Gove, Andrew M., Latimer, Jonathan D., Majer, Robert R., Dunn, vanc, 2010, Perspectives in Plant Ecology, Evolution and Systematics, 12, 43–55, 10.1016/j.ppees.2009.08.001, 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.JOURNAL, Fischer RC, Ölzant SM, Wanek W, Mayer V, The fate of Corydalis cava elaiosomes within an ant colony of Myrmica rubra: elaiosomes are preferentially fed to larvae, Insectes Sociaux, 2005, 52, 1, 55–62, 10.1007/s00040-004-0773-x, 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.JOURNAL, Hughes L, Westoby M, 1992, Capitula on stick insect eggs and elaiosomes on seeds: convergent adaptations for burial by ants, Functional Ecology, 6, 642–648, 10.2307/2389958, 6, 2389958, File:Common jassid nymph and ant02.jpg|thumb|left|A meat ant tending a common leafhopperleafhopperMost 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. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away predatory ant species.JOURNAL, Journal of Insect Behavior, Behavioural Interactions Between Crematogaster brevispinosa rochai Forel (Hymenoptera: Formicidae) and Two Nasutitermes Species (Isoptera: Termitidae), 18, 1, 1–17, 2005, Quinet Y, Tekule N, de Biseau JC, 10.1007/s10905-005-9343-y, The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellent chemical.JOURNAL, Jeanne RL, 1972, Social biology of the neotropical wasp Mischocyttarus drewseni, Bull. Mus. Comp. Zool., 144, 63–150,weblink It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Other wasps, such as A. multipicta, defend against ants by blasting them off the nest with bursts of wing buzzing.JOURNAL,weblink Foraging in Social Wasps: Agelaia lacks recruitment to food (Hymenoptera: Vespidae), Jeanne, Robert, vanc, July 1995, Journal of the Kansas Entomological Society, Stingless bees (Trigona and Melipona) use chemical defences against ants.Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants.JOURNAL, Sivinski J, Marshall S, Petersson E, 1999, Kleptoparasitism and phoresy in the Diptera, Florida Entomologist, 82, 2, 179–197,weblink PDF, 10.2307/3496570, 3496570, 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.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 JOURNAL, Schaechter E, 2000, Some weird and wonderful fungi, Microbiology Today, 27, 3, 116–117, in a microhabitat that best suits the fungus.JOURNAL, Andersen SB, Gerritsma S, Yusah KM, Mayntz D, Hywel-Jones NL, Billen J, Boomsma JJ, Hughes DP, The life of a dead ant: the expression of an adaptive extended phenotype, The American Naturalist, 174, 3, 424–33, September 2009, 19627240, 10.1086/603640, Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates.JOURNAL, Wojcik DP, 1989, Behavioral interactions between ants and their parasites, The Florida Entomologist, 72, 1, 43–51, 10.2307/3494966, 3494966,weblink PDF, 2017-10-25,weblink 2017-09-22, yes, A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black-coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, causing them to carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits, such as Hyeronima alchorneoides, and eaten. The droppings of the bird are collected by other ants and fed to their young, leading to further spread of the nematode.JOURNAL, Poinar G, Yanoviak SP, Myrmeconema neotropicum n. g., n. sp., a new tetradonematid nematode parasitising South American populations of Cephalotes atratus (Hymenoptera: Formicidae), with the discovery of an apparent parasite-induced host morph, Systematic Parasitology, 69, 2, 145–53, 2008, 18038201, 10.1007/s11230-007-9125-3, (File:Jumping spider with prey.jpg|thumb|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.JOURNAL, Caldwell JP, 1996, The evolution of myrmecophagy and its correlates in poison frogs (Family Dendrobatidae), Journal of Zoology, 240, 1, 75–101, 10.1111/j.1469-7998.1996.tb05487.x, 2010JZoo..281..263G, Army ants forage in a wide roving column, attacking any animals in that path that are unable to escape. In Central and South America, Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers.JOURNAL, 10.1146/annurev.es.09.110178.001331, Willis E, Oniki Y, 1978, Birds and Army Ants, Annual Review of Ecology and Systematics, 9, 243–263, JOURNAL, Vellely AC, Foraging at army ant swarms by fifty bird species in the highlands of Costa Rica, Ornitologia Neotropical, 12, 2001, 271–275,weblink PDF, 8 June 2008,
This behaviour was once considered mutualistic, but later studies found the birds to be parasitic. Direct kleptoparasitism (birds stealing food from the ants' grasp) is rare and has been noted in Inca doves which pick seeds at nest entrances as they are being transported by species of Pogonomyrmex.JOURNAL, Inzunza, Ernesto Ruelas, Martínez-Leyva, J. Eduardo, Valenzuela-González, Jorge E., vanc, Doves kleptoparasitize ants, The Southwestern Naturalist, 60, 1, 103–106, 10.1894/msh-03.1, 2015, Birds that follow ants eat many prey insects and thus decrease the foraging success of ants.JOURNAL, Wrege PH, Antbirds parasitize foraging army ants, Ecology, 86, 2005, 555–559, 10.1890/04-1133, Wikelski, Martin, Mandel, James T., Rassweiler, Thomas, Couzin, Iain D., vanc, 3, Birds indulge in a peculiar behaviour called anting that, as yet, is not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may be a means to remove ectoparasites from the birds.
Anteaters, aardvarks, pangolins, echidnas and numbats have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants. About 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.JOURNAL, 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, 10.1139/cjz-77-4-551,

Relationship with humans

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- AntsStitchingLeave.jpg -
upWeaver 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.Hölldobler & Wilson (1990), pp. 619–629 On the other hand, ants may become nuisances when they invade buildings, or cause economic losses.In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.JOURNAL, Gottrup F, Leaper D, 2004, Wound healing: Historical aspects, EWMA Journal, 4, 2,weblinkweblink" title="web.archive.org/web/20070616090223weblink">weblink 2007-06-16, PDF, 5, JOURNAL, Gudger EW, 1925, Stitching wounds with the mandibles of ants and beetles, Journal of the American Medical Association, 84, 1861–1864, 10.1001/jama.1925.02660500069048, 24, BOOK, Robert M., Sapolsky, vanc, Robert Sapolsky, 2001, A Primate's Memoir: A Neuroscientist's Unconventional Life Among the Baboons, 156, Simon and Schuster, 978-0-7432-0241-1, The large heads of the dinergates (soldiers) of the leafcutting ant Atta cephalotes are also used by native surgeons in closing wounds.BOOK, Wheeler, William M., vanc, William Morton Wheeler, Ants: Their Structure, Development and Behavior, 1910, Columbia University Biological Series, 9, Columbia University Press, 978-0-231-00121-2, 10, 10.5962/bhl.title.1937, 10008253//r88, 560205, s:Index:Ants, Wheeler (1910).djvu, Some ants have toxic venom and are of medical importance. The species include Paraponera clavata (tocandira) and Dinoponera spp. (false tocandiras) of South America JOURNAL, Haddad V, Cardoso JL, Moraes RH, 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), Revista Do Instituto de Medicina Tropical de São Paulo, 47, 4, 235–238, 10.1590/S0036-46652005000400012, and the Myrmecia ants of Australia.JOURNAL, McGain F, Winkel KD, Ant sting mortality in Australia, Toxicon, 40, 8, 1095–100, August 2002, 12165310, 10.1016/S0041-0101(02)00097-1, In South Africa, ants are used to help harvest the seeds of rooibos (Aspalathus linearis), a plant used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds may be collected from one ant-heap.WEB,weblink PDF, Innovative mechanisms for sharing benefits of biodiversity and related knowledge, Downes D, Laird SA, 1999, The Center for International Environmental Law, 8 June 2008,weblink" title="web.archive.org/web/20080423211123weblink">weblink 23 April 2008, no, JOURNAL, Economic Botany, Rooibos tea, a South African contribution to world beverages, 17, 3, 186–194, 1963, Cheney RH, Scholtz E,weblink 10.1007/BF02859435, Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These tend to be island species that have evolved specialized traits and risk being displaced by introduced ant species. Examples include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.JOURNAL, The influence of sociality on the conservation biology of social insects, Chapman RE, Bourke AF, Ecology Letters, 4, 6, 2001, 650–662, 10.1046/j.1461-0248.2001.00253.x, 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) (i.e. between 1015 and 1016). 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.BOOK, Holldobler B, Wilson EO, The Superorganism: The Beauty, Elegance, and Strangeness of Insect Societies, 2009, W. W. Norton, New York, 5, 978-0-393-06704-0, Also, according to this estimate, there are approximately 1 million ants for every human on Earth.WEB, How many ants are there for every one person on earth?,weblink info.com, 27 July 2013, yes,weblink" title="web.archive.org/web/20130813234547weblink">weblink 13 August 2013,

As food

{{See also|Entomophagy}}(File:Ants For Food SG.jpg|thumb|left|Roasted ants in Colombia)
missing image!
- Ants Eggs Market Thailand.jpg -
Ant larvae for sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are used in Mexican escamoles. They are considered a form of insect caviar and can sell for as much as US$40 per pound ($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.JOURNAL, DeFoliart GR, Insects as food: why the western attitude is important, Annual Review of Entomology, 44, 21–50, 1999, 9990715, 10.1146/annurev.ento.44.1.21, 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.BOOK, Bingham CT, 1903, Fauna of British India. Hymenoptera Volume 3, 311, Weaver ant eggs and larvae, as well as the ants, may be used in a Thai salad, yam (), in a dish called yam khai mot daeng () 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".JOURNAL, 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,weblink In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickling, acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).

As pests

{{See also|Ants of medical importance}}File:Monomorium pharaonis.jpg|thumb|The tiny pharaoh antpharaoh antSome ant species are considered as pests, primarily those that occur in human habitations, where their presence is often problematic. For example, the presence of ants would be undesirable in sterile places such as hospitals or kitchens. Some species or genera commonly categorized as pests include the Argentine ant, pavement ant, yellow crazy ant, banded sugar ant, Pharaoh ant, carpenter ants, odorous house ant, red imported fire ant, and European fire ant. Some ants will raid stored food, others may damage indoor structures, some can damage agricultural crops directly (or by aiding sucking pests), and some will sting or bite.WEB,weblink Pest Notes: Ants (Publication 7411), University of California Agriculture and Natural Resources, 2007, 5 June 2008, The adaptive nature of ant colonies make it nearly impossible to eliminate entire colonies and most pest management practices aim to control local populations and tend to be temporary solutions. Ant populations are managed by a combination of approaches that make use of chemical, biological and physical methods. Chemical methods include the use of insecticidal bait which is gathered by ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Management is based on the species and techniques can vary according to the location and circumstance.

In science and technology

File:Formicariums.jpg|thumb|right{{See also|Myrmecology|Biomimetics|Ant colony optimization algorithms}}Observed by humans since the dawn of history, the behaviour of ants has been documented and the subject of early writings and fables passed from one century to another. Those using scientific methods, myrmecologists, study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbock in 1881.JOURNAL, 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, 110–112, 10.1111/j.1096-3642.1882.tb02275.x, 90, 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.BOOK, Stadler B, Dixon AF, 2008, Mutualism: Ants and their insect partners, Cambridge University Press, 978-0-521-86035-2, Ant colonies may be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures.JOURNAL, Myrmecological technique. IV. Collecting ants by rearing pupae, Kennedy CH, The Ohio Journal of Science, 51, 1, 1951, 17–20, 1811/3802, Individuals may be tracked for study by marking them with dots of colours.JOURNAL, Wojcik DP, Burges RJ, Blanton CM, Focks DA, 2000, An improved and quantified technique for marking individual fire ants (Hymenoptera: Formicidae), The Florida Entomologist, 83, 1, 74–78, 10.2307/3496231,weblink PDF, 3496231, 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.JOURNAL, Dicke E, Byde A, Cliff D, Layzell P, 2004, An ant-inspired technique for storage area network design, Proceedings of Biologically Inspired Approaches to Advanced Information Technology: First International Workshop, BioADIT 2004 LNCS 3141, 364–379, A. J. Ispeert, M. Murata, N. Wakamiya,

As pets

From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Some later commercial versions use transparent gel instead of soil, allowing greater visibility at the cost of stressing the ants with unnatural light.PATENT, US, 5803014, granted, Habitat media for ants and other invertebrates, 8 September 1998, Guri A, Plant Cell Technology Inc,

In culture

missing image!
- The Ant and the Grasshopper - Project Gutenberg etext 19994.jpg -
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 also are mentioned in religious texts.BOOK, Quran, The Ant, The Ants, Surah 27, 18–19,weblink yes,weblink" title="web.archive.org/web/20070101181115weblink">weblink 2007-01-01, BOOK, Sahih, Bukhari, vanc, Sunnah, 4 Book 54, 536, Beginning of Creation,weblinkweblink" title="web.archive.org/web/20000818092520weblink">weblink yes, 2000-08-18, 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 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, BOOK, Mawil Y. Izzi, Deen, vanc, Islamic Environmental Ethics, Law, and Society, Ethics of Environment and Development, Engel JR, JG Engel, 1990, Bellhaven Press, London,weblinkweblink" title="web.archive.org/web/20110714055330weblink">weblink yes, 2011-07-14, PDF, In parts of Africa, ants are considered to be the messengers of the deities. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief.JOURNAL, Balee WL, Antiquity of traditional ethnobiological knowledge in Amazonia: The Tupi-Guarani family and time, Ethnohistory (journal), Ethnohistory, 47, 2, 2000, 399–422, 10.1215/00141801-47-2-399, Ant bites are used in the initiation ceremonies of some Amazon Indian cultures as a test of endurance.BOOK, Cesard N, Deturche J, Erikson P, 2003, Les Insectes dans les pratiques médicinales et rituelles d'Amazonie indigène, Motte-Florac, E., J. M. C. Thomas, Les insectes dans la tradition orale, Peeters-Selaf, Paris, 395–406, fr, JOURNAL, Schmidt RJ, The super-nettles. A dermatologist's guide to ants in the plants, International Journal of Dermatology, 24, 4, 204–10, May 1985, 3891647, 10.1111/j.1365-4362.1985.tb05760.x,weblink 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.BOOK, 22 The Black Forest and Its Treasures, A Tramp Abroad, Twain, Mark, vanc, 1880,weblink 978-0-19-510137-9, Oxford University Press, New York, 2015-12-13, 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. H.G. Wells wrote about intelligent ants destroying human settlements in Brazil and threatening human civilization in his 1905 science-fiction short story, The Empire of the Ants. In more recent times, animated cartoons and 3-D animated films featuring ants have been produced including Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Ferdy the Ant and Atom Ant. Renowned myrmecologist E. O. Wilson wrote a short story, "Trailhead" in 2010 for The New Yorker magazine, which describes the life and death of an ant-queen and the rise and fall of her colony, from an ants' point of view.WEB, Wilson, EO,weblink Trailhead, The New Yorker, 25 January 2010, 56–62, The French neuroanatomist, psychiatrist and eugenicist Auguste Forel believed that ant societies were models for human society. He published a five volume work from 1921 to 1923 that examined ant biology and society.JOURNAL, Parent, André, vanc, 2003, Auguste Forel on Ants and Neurology,weblink The Canadian Journal of Neurological Sciences, 30, 3, 284–291, 10.1017/s0317167100002754, In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".WEB,weblinkweblink" title="web.archive.org/web/20090611002722weblink">weblink 2009-06-11, 1992 Excellence in Software Awards Winners, 3 April 2008, Software & Information Industry Association, Ants also are quite popular inspiration for many science-fiction insectoids, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the films Them! and Empire of the Ants, Marvel Comics' super hero Ant-Man, and ants mutated into super-intelligence in Phase IV. In computer strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.JOURNAL, Robots, insects and swarm intelligence, Artificial Intelligence Review, 26, 4, 2006, Sharkey AJC, 10.1007/s10462-007-9057-y, 255–268,

See also

{{Wikipedia books|Ants}}

References

{{Reflist|32em}}

Cited texts

  • BOOK, Borror DJ, Triplehorn CA, Delong DM, Introduction to the Study of Insects, 6th Edition, Saunders College Publishing, 1989, 978-0-03-025397-3,
  • BOOK, Hölldobler B, Wilson EO, The Ants, Harvard University Press, 1990, 978-0-674-04075-5, The Ants,

Further reading

  • BOOK, Bolton, Barry, Harvard University Press, 1995, 978-0-674-61514-4, A New General Catalogue of the Ants of the World,
  • BOOK, Hölldobler B, Wilson EO, 1998, Journey to the Ants: A Story of Scientific Exploration, Belknap Press, 978-0-674-48526-6,
  • BOOK, Hölldobler B, Wilson EO, 2009, The Superorganism: The Beauty, Elegance and Strangeness of Insect Societies, Norton & Co., 978-0-393-06704-0,

External links

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