Editing Ant (section)

==Life cycle==
[[Image:Meat eater ant nest swarming02.jpg|thumb|[[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 [[pupa]]l 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 [[Regurgitation (digestion)|regurgitates]] liquid food held in its [[Crop (anatomy)|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 egg]]s, pieces of prey, and seeds brought by workers.<ref>{{cite book| vauthors = Hölldobler B, Wilson EO |title=The Ants|url=https://archive.org/details/ants0000hlld |url-access=registration |year=1990 |publisher=Harvard University Press|isbn=978-0-674-04075-5 |page=[https://archive.org/details/ants0000hlld/page/n316 291]}}</ref>

The larvae grow through a series of four or five [[moult]]s and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a [[Chrysalis|butterfly pupa]].<ref>{{cite book|author=Gillott, Cedric |year=1995|title=Entomology|publisher=Springer|isbn=978-0-306-44967-3|page=325}}</ref> The differentiation into queens and workers (which are both female), and different [[caste (biology)|castes]] of workers, is influenced in some species by the nutrition the larvae obtain. Genetic influences and the [[polyphenism|control of gene expression]] by the developmental environment are complex and the determination of caste continues to be a subject of research.<ref>{{cite journal|title=The causes and consequences of genetic caste determination in ants (Hymenoptera: Formicidae) |first1=Kirk E. |last1=Anderson |first2=Timothy A. |last2=Linksvayer |first3=Chris R. |last3=Smith  | name-list-style=vanc |journal=Myrmecol. News |volume=11 |pages=119–132 |year=2008 |url=https://myrmecologicalnews.org/cms/index.php?option=com_content&view=category&id=250&Itemid=73}}</ref> Winged male ants, called drones (termed "aner" in old literature<ref name=":1">{{Cite journal| vauthors = Gaul AT |date=1951|title=A Glossary of Terms and Phrases Used in the Study of Social Insects|url=https://academic.oup.com/aesa/article-lookup/doi/10.1093/aesa/44.3.473|journal=Annals of the Entomological Society of America|volume=44|issue=3|pages=473–484|doi=10.1093/aesa/44.3.473|issn=1938-2901}}</ref>), emerge from pupae along with the usually winged breeding females. Some species, such as [[army ant]]s, 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.<ref>Hölldobler & Wilson (1990), pp. 351, 372</ref>

A new ergate spends the first few days of its adult life caring for the queen and young. She then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. Such age-based task-specialization or [[polyethism]] has been suggested as having evolved due to the high casualties involved in foraging and defence, making it an acceptable risk only for ants who are older and likely to die sooner from natural causes.<ref>{{cite journal|journal=Annual Review of Entomology|year=1989|volume=34|pages=191–210|title=Foraging strategies of ants|author=Traniello JFA|doi=10.1146/annurev.en.34.010189.001203}}</ref><ref>{{cite journal|vauthors=Sorensen A, Busch TM, Vinson SB |title=Behavioral flexibility of temporal sub-castes in the fire ant, ''Solenopsis invicta'', in response to food|journal=Psyche: A Journal of Entomology|volume=91|pages=319–332|year=1984|doi=10.1155/1984/39236|issue=3–4|doi-access=free}}</ref> In the Brazilian ant ''[[Forelius pusillus]],'' the nest entrance is closed from the outside to protect the colony from predatory ant species at sunset each day. About one to eight workers seal the nest entrance from the outside and they have no chance of returning to the nest and are in effect sacrificed.<ref>{{Cite journal |last1=Tofilski |first1=Adam |last2=Couvillon |first2=Margaret J. |last3=Evison |first3=Sophie E. F. |last4=Helanterä |first4=Heikki |last5=Robinson |first5=Elva J. H. |last6=Ratnieks |first6=Francis L. W. |date=2008 |title=Preemptive Defensive Self-Sacrifice by Ant Workers |journal=The American Naturalist|volume=172 |issue=5 |pages=E239–E243 |doi=10.1086/591688 |pmid=18928332 |bibcode=2008ANat..172E.239T |s2cid=7052340 |issn=0003-0147}}</ref> Whether these seemingly suicidal workers are older workers has not been determined.<ref>{{Cite journal |last1=Shorter |first1=J. R. |last2=Rueppell |first2=O. |date=2012 |title=A review on self-destructive defense behaviors in social insects |url=http://link.springer.com/10.1007/s00040-011-0210-x |journal=Insectes Sociaux|volume=59 |issue=1 |pages=1–10 |doi=10.1007/s00040-011-0210-x |s2cid=253634662 |issn=0020-1812}}</ref>

Ant colonies can be long-lived. The queens can live for up to 30&nbsp;years, and workers live from 1 to 3&nbsp;years. Males, however, are more transitory, being quite short-lived and surviving for only a few weeks.<ref>{{cite journal |author=Keller L|year=1998 |title=Queen lifespan and colony characteristics in ants and termites |journal=[[Insectes Sociaux]] |volume=45 |pages=235–246 |doi=10.1007/s000400050084 |issue=3|s2cid=24541087 }}</ref> Ant queens are estimated to live 100 times as long as solitary insects of a similar size.<ref name=insencyc>{{cite book|editor-link1=Nigel R. Franks|veditors=Franks NR, Resh VH, Cardé RT|year=2003|title=Encyclopedia of Insects|pages=[https://archive.org/details/encyclopediaofin00bada/page/29 29–32]|isbn=978-0-12-586990-4|publisher=Academic Press|location=San Diego|url=https://archive.org/details/encyclopediaofin00bada/page/29}}</ref>

Ants are active all year long in the tropics; however, in cooler regions, they survive the winter in a state of dormancy known as [[hibernation]]. The forms of inactivity are varied and some temperate species have larvae going into the inactive state ([[diapause]]), while in others, the adults alone pass the winter in a state of reduced activity.<ref>{{cite journal|author=Kipyatkov VE|year=2001|title=Seasonal life cycles and the forms of dormancy in ants (Hymenoptera, Formicoidea)|journal=Acta Societatis Zoologicae Bohemicae|volume=65|issue=2|pages=198–217}}</ref>

===Reproduction===
[[Image:FlyingAnts.jpg|thumb|Honey ant (''[[Prenolepis imparis]]'') mating, the drone is much smaller than the queen]]
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 [[thelytoky|thelytokous parthenogenesis]].<ref>{{cite journal| vauthors = Heinze J, Tsuji K |year=1995|title=Ant reproductive strategies|journal=Res. Popul. Ecol.|volume=37|issue=2|pages=135–149|url=http://meme.biology.tohoku.ac.jp/POPECOL/RP%20PDF/37(2)/pp.135.pdf|doi=10.1007/BF02515814|bibcode=1995PopEc..37..135H |s2cid=21948488|access-date=2009-04-16|archive-url=https://web.archive.org/web/20110527051529/http://meme.biology.tohoku.ac.jp/POPECOL/RP%20PDF/37(2)/pp.135.pdf|archive-date=2011-05-27|url-status=dead}}</ref> Secretions from the male accessory glands in some species can plug the female genital opening and prevent females from re-mating.<ref name="Mikheyev, a. S. 2003. pp.401">{{Cite journal | doi = 10.1007/s00040-003-0697-x | title = Evidence for mating plugs in the fire ant ''Solenopsis invicta'' | journal = Insectes Sociaux | volume = 50 | issue = 4 | pages = 401–402 | year = 2003 | vauthors = Mikheyev AS | s2cid = 43492133 }}</ref> 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 "[[gamergate (ant)|gamergates]]" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right.<ref>{{cite journal | vauthors = Peeters C, Hölldobler B | title = Reproductive cooperation between queens and their mated workers: the complex life history of an ant with a valuable nest | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 92 | issue = 24 | pages = 10977–10979 | date = November 1995 | pmid = 11607589 | pmc = 40553 | doi = 10.1073/pnas.92.24.10977 | bibcode = 1995PNAS...9210977P | doi-access = free }}</ref>

Drones can also mate with existing queens by entering a foreign colony, such as in [[army ant]]s. 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.<ref name="Franks N. 2008">{{Cite journal | vauthors = Franks NR, Hölldobler B | doi = 10.1111/j.1095-8312.1987.tb00298.x | title = Sexual competition during colony reproduction in army ants | journal = Biological Journal of the Linnean Society | volume = 30 | issue = 3 | pages = 229–243 | year = 1987 }}</ref> 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.<ref>{{Cite journal | doi = 10.1007/BF01175395 | title = Pheromonal manipulation of workers by a fighting male to kill his rival males in the ant ''Cardiocondyla wroughtonii'' | journal = Naturwissenschaften | volume = 79 | issue = 6 | pages = 274–276 | year = 1992 | vauthors = Yamauchi K, Kawase N | bibcode = 1992NW.....79..274Y | s2cid = 31191187 }}</ref>

Most ants are [[univoltine]], producing a new generation each year.<ref name="bloodywasp">{{cite book| vauthors = Taylor RW |year=2007 |chapter=Bloody funny wasps! Speculations on the evolution of eusociality in ants|pages=580–609|veditors=Snelling RR, Fisher BL, Ward PS|title=Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson&nbsp;– 50 years of contributions. Memoirs of the American Entomological Institute, 80|publisher=American Entomological Institute | chapter-url = http://antbase.org/ants/publications/21292/21292.pdf |access-date=2015-12-13}}</ref> During the species-specific breeding period, winged females and winged males, known to [[entomologists]] as [[alate]]s, 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.<ref name="nuptial flight 2">{{cite journal| vauthors = Wilson EO |year=1957|title=The organization of a nuptial flight of the ant ''Pheidole sitarches'' Wheeler | journal = Psyche: A Journal of Entomology | url = http://www.antwiki.org/wiki/images/7/7f/Wilson_1958d.pdf |volume=64|issue= 2|pages=46–50|doi=10.1155/1957/68319|doi-access=free}}</ref> 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 [[Spermatozoon|sperm]] in their [[spermatheca]]e.<ref name="HolldoblerWilsonAnts2">Hölldobler & Wilson (1990), pp. 143–179</ref> The genus ''[[Cardiocondyla]]'' have species with both winged and wingless males, where the latter will only mate with females living in the same nest. Some species in the genus have lost winged males completely, and only produce wingless males.<ref>{{Cite journal |last=Heinze |first=Jürgen |date=2017 |title=Life-history evolution in ants: the case of Cardiocondyla |journal=  Proceedings of the Royal Society B: Biological Sciences|language=en |volume=284 |issue=1850 |pages=20161406 |doi=10.1098/rspb.2016.1406 |issn=0962-8452 |pmc=5360909 |pmid=28298341}}</ref> In ''[[Cardiocondyla elegans|C. elegans]],'' workers may transport newly emerged queens to other conspecific nests where the wingless males from unrelated colonies can mate with them, a behavioural adaptation that may reduce the chances of inbreeding.<ref>{{cite journal | vauthors = Vidal M, Königseder F, Giehr J, Schrempf A, Lucas C, Heinze J | title = Worker ants promote outbreeding by transporting young queens to alien nests | journal = Communications Biology | volume = 4 | issue = 1 | pages = 515 | date = May 2021 | pmid = 33941829 | pmc = 8093424 | doi = 10.1038/s42003-021-02016-1 }}</ref>

[[Image:Meat eater ant qeen excavating hole.jpg|thumb|left|Fertilised meat-eater ant queen beginning to dig a new colony]]

Mated females then seek a suitable place to begin a colony. There, they break off their wings using their tibial spurs and begin to lay and care for eggs. The females can selectively fertilise future eggs with the sperm stored to produce diploid workers or lay unfertilized haploid eggs to produce drones. The first workers to hatch, known as nanitics,<ref name="SuddFranks2013">{{cite book | vauthors = Sudd JH, Franks NR |title=The Behavioural Ecology of Ants|year=2013|publisher=Springer Science & Business Media|isbn=978-9400931237|page=41}}</ref> 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,<ref name="HolldoblerWilsonAnts2"/> a process akin to [[Swarming (honey bee)|swarming]] in [[honeybee]]s.


=== Nests, colonies, and supercolonies ===
The typical ant species has a colony occupying a single nest, housing one or more queens, where the brood is raised. There are however more than 150 species of ants in 49 genera that are known to have colonies consisting of multiple spatially separated nests. These polydomous (as opposed to monodomous) colonies have food and workers moving between the nests.<ref>{{Cite journal |last1=Cook |first1=Zoe |last2=Franks |first2=Daniel W. |last3=Robinson |first3=Elva J.H. |date=2013 |title=Exploration versus exploitation in polydomous ant colonies |url=https://linkinghub.elsevier.com/retrieve/pii/S0022519313000477 |journal=Journal of Theoretical Biology|volume=323 |pages=49–56 |doi=10.1016/j.jtbi.2013.01.022|pmid=23380232 |bibcode=2013JThBi.323...49C }}</ref> Membership to a colony is identified by the response of worker ants which identify whether another individual belongs to their own colony or not. A signature cocktail of body surface chemicals (also known as cuticular hydrocarbons or CHCs) forms the so-called colony odor which other members can recognize.<ref>{{Cite journal |last1=Bos |first1=Nick |last2=d’Ettorre |first2=Patrizia |date=2012 |title=Recognition of Social Identity in Ants |journal=Frontiers in Psychology |volume=3 |page=83 |doi=10.3389/fpsyg.2012.00083 |issn=1664-1078 |pmc=3309994 |pmid=22461777 |doi-access=free }}</ref> Some ant species appear to be less discriminating and in the Argentine ant ''Linepithema humile,'' workers carried from a colony anywhere in the southern US and Mexico are acceptable within other colonies in the same region. Similarly workers from colonies that have established in Europe are accepted by any other colonies within Europe but not by the colonies in the Americas. The interpretation of these observations has been debated and some have been termed these large populations as supercolonies<ref>{{Cite journal |last=Moffett |first=Mark W. |date=2012 |title=Supercolonies of billions in an invasive ant: What is a society? |url=https://academic.oup.com/beheco/article-lookup/doi/10.1093/beheco/ars043 |journal=Behavioral Ecology|volume=23 |issue=5 |pages=925–933 |doi=10.1093/beheco/ars043 |issn=1465-7279}}</ref><ref>{{Cite journal |last1=Van Wilgenburg |first1=Ellen |last2=Torres |first2=Candice W. |last3=Tsutsui |first3=Neil D. |date=2010 |title=The global expansion of a single ant supercolony |journal=Evolutionary Applications|volume=3 |issue=2 |pages=136–143 |doi=10.1111/j.1752-4571.2009.00114.x |issn=1752-4571 |pmc=3352483 |pmid=25567914}}</ref><ref>{{Cite journal |last=Heikki Helanterä |date=2022 |title=Supercolonies of ants (Hymenoptera: Formicidae): ecological patterns, behavioural processes and their implications for social evolution |url=https://myrmecologicalnews.org/cms/index.php?option=com_content&view=category&id=1593&Itemid=441 |journal=Myrmecological News |volume=32 |pages=1–22 |doi=10.25849/MYRMECOL.NEWS_032:001}}</ref> while others have termed the populations as unicolonial.<ref>{{Cite journal |last1=Helanterä |first1=Heikki |last2=Strassmann |first2=Joan E. |last3=Carrillo |first3=Juli |last4=Queller |first4=David C. |date=2009 |title=Unicolonial ants: where do they come from, what are they and where are they going? |url=https://linkinghub.elsevier.com/retrieve/pii/S0169534709000895 |journal=Trends in Ecology & Evolution|volume=24 |issue=6 |pages=341–349 |doi=10.1016/j.tree.2009.01.013|pmid=19328589 |bibcode=2009TEcoE..24..341H }}</ref>