Editing Termite (section)

==Life cycle==
[[File:Termite-by-RalfR.jpg|thumbnail|alt=A termite nymph looks like a smaller version of an adult but lacks the specialisations that would enable identification of its caste.|A young termite nymph featuring visible wing buds. Nymphs mainly develop into [[alate]]s.]]
[[File:20200403 Termite and shed wings on interior window sill.png|thumb|A termite alate with shed wings from other alates on an interior window sill. Shedding of wings is associated with reproductive swarming.<ref name=NewYorker_20180918>{{cite magazine |last1=Srinivasan |first1=Amia |title=What Termites Can Teach Us |url=https://www.newyorker.com/magazine/2018/09/17/what-termites-can-teach-us |magazine=The New Yorker |date=September 10, 2018 |archive-url=https://web.archive.org/web/20200307022415/https://www.newyorker.com/magazine/2018/09/17/what-termites-can-teach-us |archive-date=March 7, 2020 |url-status=live }}</ref>]]
Termites are often compared with the [[eusociality|social]] Hymenoptera (ants and various species of bees and wasps), but their differing evolutionary origins result in major differences in life cycle. In the eusocial Hymenoptera, the workers are exclusively female. Males (drones) are haploid and develop from unfertilised eggs, while females (both workers and the queen) are diploid and develop from fertilised eggs. In contrast, worker termites, which constitute the majority in a colony, are [[diploid]] individuals of both sexes and develop from fertilised eggs. Depending on species, male and female workers may have different roles in a termite colony.<ref>{{cite journal |last1=Korb |first1=J. |title=Termites, hemimetabolous diploid white ants? |journal=Frontiers in Zoology |date=2008 |volume=5 |issue=1 |pages=15 |doi=10.1186/1742-9994-5-15 |pmc=2564920 |pmid=18822181 |doi-access=free }}</ref>

The life cycle of a termite begins with an [[egg (biology)|egg]], but is different from that of a bee or ant in that it goes through a developmental process called [[incomplete metamorphosis]], going through multiple gradual pre-adult molts that are highly [https://www.sciencedirect.com/topics/medicine-and-dentistry/developmental-plasticity#:~:text=Developmental%20plasticity%20refers%20to%20the,generated%20from%20a%20single%20genotype. developmentally plastic] before becoming an adult.<ref name=":4" /><ref>{{cite web |last1=Davis |first1=P. |title=Termite Identification |url=http://agspsrv34.agric.wa.gov.au/ento/termites.htm |publisher=Entomology at Western Australian Department of Agriculture |url-status=dead |archive-url=http://webarchive.loc.gov/all/20090612011547/http://agspsrv34.agric.wa.gov.au/ento/termites.htm |archive-date=2009-06-12 }}</ref> Unlike in other [[Hemimetabolism|hemimetabolous]] insects, ''nymphs'' are more strictly defined in termites as immature young with visible wing buds, which often invariably go through a series of [[Moulting|moults]] to become winged [[Imago|adults]].<ref>{{cite journal|last1=Neoh |first1=K.B. |last2=Lee|first2=C.Y.|title=Developmental stages and caste composition of a mature and incipient colony of the drywood termite, ''Cryptotermes dudleyi'' (Isoptera: Kalotermitidae) |journal=Journal of Economic Entomology |date=2011 |volume=104 |issue=2 |pages=622–8 |pmid=21510214 |doi=10.1603/ec10346|s2cid=23356632 |doi-access=free }}</ref><ref name=":4" /> ''Larvae'', which are defined as early nymph instars with absent wing buds, exhibit the highest developmental potentiality and are able to molt into ''Alates'', ''Soldiers'', ''Neotenics'', or ''Workers''. Workers are believed to have evolved from larvae, sharing many similarities to the extent that workers can be regarded as "larval", in that both lack wings, eyes, and functional reproductive organs while maintaining varying levels of developmental flexibility, although usually to a much lesser extent in workers. The main distinction being that while larvae are wholly dependent on other nestmates to survive, workers are independent and are able to feed themselves and contribute to the colony. Workers remain wingless and across many [[Taxon|taxa]] become developmentally arrested, appearing to not change into any other caste until death.<ref name=":4" /> In some basal taxa, there is no distinction, with the "workers" (pseudergates) essentially being late instar larvae that retain the ability to change into all other castes.<ref name=":5" />

The development of larvae into adults can take months; the time period depends on food availability and nutrition, temperature, and the size of the colony. Since larvae and nymphs are unable to feed themselves, workers must feed them, but workers also take part in the social life of the colony and have certain other tasks to accomplish such as foraging, building or maintaining the nest or tending to the queen.<ref name=aus>{{cite web|title=Termites|url=https://australian.museum/learn/animals/insects/termites/|publisher=Australian Museum|access-date=8 January 2015}}</ref><ref name=Michael>{{cite web|last1=Schneider|first1=M.F.|title=Termite Life Cycle and Caste System|url=http://www.fzi.uni-freiburg.de/InsectPestKey-long%20version/termit2.htm|publisher=University of Freiburg|access-date=8 January 2015|date=1999}}</ref> Pheromones regulate the caste system in termite colonies, preventing all but a very few of the termites from becoming fertile queens.<ref>{{cite journal|last1=Simpson|first1=S.J.|last2=Sword|first2=G.A.|last3=Lo|first3=N.|title=Polyphenism in Insects|journal=Current Biology|date=2011|volume=21|issue=18|pages=738–749|doi=10.1016/j.cub.2011.06.006|pmid=21959164|s2cid=656039|doi-access=free|bibcode=2011CBio...21.R738S }}</ref>

Queens of the [[eusociality|eusocial]] termite ''[[Reticulitermes]] speratus'' are capable of a long lifespan without sacrificing [[fecundity]]. These long-lived queens have a significantly lower level of oxidative damage, including [[DNA oxidation|oxidative DNA damage]], than workers, soldiers and nymphs.<ref name="pmid28076409">{{cite journal |vauthors=Tasaki E, Kobayashi K, Matsuura K, Iuchi Y |title=An Efficient Antioxidant System in a Long-Lived Termite Queen |journal=PLOS ONE |volume=12 |issue=1 |pages=e0167412 |date=2017 |pmid=28076409 |pmc=5226355 |doi=10.1371/journal.pone.0167412 |bibcode=2017PLoSO..1267412T |doi-access=free }}</ref> The lower levels of damage appear to be due to increased [[catalase]], an enzyme that protects against [[oxidative stress]].<ref name="pmid28076409" />

===Reproduction===
[[File:Flying Termites after rain.jpg|thumbnail|alt=Hundreds of winged termite reproductives swarming after a summer rain, filling the field of the photograph.|Alates swarming during nuptial flight after rain|left]]
Termite alates (winged virgin queens and kings) only leave the colony when a [[nuptial flight]] takes place. Alate males and females pair up together and then land in search of a suitable place for a colony.<ref name=v2010>{{cite web|last1=Miller|first1=D.M.|title=Subterranean Termite Biology and Behavior|publisher=Virginia Tech (Virginia State University)|access-date=8 January 2015|url=https://pubs.ext.vt.edu/444/444-502/444-502.html|date=5 March 2010}}</ref> A termite king and queen do not mate until they find such a spot. When they do, they excavate a chamber big enough for both, close up the entrance and proceed to mate.<ref name=v2010/> After mating, the pair may never surface again, spending the rest of their lives in the nest. Nuptial flight time varies in each species. For example, alates in certain species emerge during the day in summer while others emerge during the winter.<ref name=Gouge>{{cite web|last1=Gouge|first1=D.H.|last2=Smith|first2=K.A.|last3=Olson|first3=C.|last4=Baker|first4=P.|title=Drywood Termites|url=http://ag.arizona.edu/pubs/insects/az1232/|publisher=University of Arizona|work=Cooperative Extension, College of Agriculture & Life Sciences|access-date=16 September 2015|date=2001|archive-date=10 November 2016|archive-url=https://web.archive.org/web/20161110102210/http://ag.arizona.edu/pubs/insects/az1232/|url-status=dead}}</ref> The nuptial flight may also begin at dusk, when the alates swarm around areas with many lights. The time when nuptial flight begins depends on the environmental conditions, the time of day, moisture, wind speed and precipitation.<ref name=Gouge/> The number of termites in a colony also varies, with the larger species typically having 100–1,000 individuals. However, some termite colonies, including those with many individuals, can number in the millions.<ref name=styria/>

The queen only lays 10–20 eggs in the very early stages of the colony, but lays as many as 1,000 a day when the colony is several years old.<ref name=aus/> At maturity, a primary queen has a great capacity to lay eggs. In some species, the mature queen has a greatly distended abdomen and may produce 40,000 eggs a day.<ref>{{cite journal |last1=Kaib |first1=M. |last2=Hacker |first2=M. |last3=Brandl |first3=R. |title=Egg-laying in monogynous and polygynous colonies of the termite ''Macrotermes michaelseni'' (Isoptera, Macrotermitidae)|journal=Insectes Sociaux |date=2001 |volume=48 |issue=3 |pages=231–237 |doi=10.1007/PL00001771|s2cid=35656795 }}</ref> The two mature ovaries may have some 2,000 [[ovariole]]s each.<ref>{{cite book|last1=Gilbert|first1=executive editors, G.A. Kerkut, L.I.|title=Comprehensive insect physiology, biochemistry, and pharmacology |date=1985 |publisher=Pergamon Press |location=Oxford |isbn=978-0-08-026850-7 |page=167 |edition=1st |url=https://books.google.com/books?id=bkTgBAAAQBAJ&q=termite+queen+ovarioles&pg=PA167}}</ref> The abdomen increases the queen's body length to several times more than before mating and reduces her ability to move freely; attendant workers provide assistance.

[[File:Egg grooming behaviour of Reticulitermes speratus workers in a nursery cell - pone.0000813.s006.ogv|thumb|alt=In this movie, workers pick eggs from the egg pile, groom them and then return them into the egg pile. They coat the eggs with their saliva to protect them against drying out and to protect them against infection: the saliva contains lyzozyme which has antibacterial properties. The lysozyme also acts as an egg recognition pheromone.|Egg grooming behaviour of ''[[Reticulitermes|Reticulitermes speratus]]'' workers in a nursery cell]]The king grows only slightly larger after initial mating and continues to mate with the queen for life (a termite queen can live between 30&nbsp;and 50&nbsp;years); this is very different from ant colonies, in which a queen mates once with the males and stores the gametes for life, as the male ants die shortly after mating.<ref name=Keller1998>{{cite journal |last1=Keller|first1=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><ref name=Michael/> If a queen is absent, a termite king produces pheromones which encourage the development of replacement termite queens.<ref>{{cite book|last1=Wyatt|first1=T.D.|title=Pheromones and animal behaviour: communication by smell and taste|url=https://archive.org/details/pheromonesanimal0000wyat|url-access=registration|date=2003|publisher=Cambridge University Press|location=Cambridge|isbn=978-0-521-48526-5|page=[https://archive.org/details/pheromonesanimal0000wyat/page/119 119]|edition=Repr. with corrections 2004.}}</ref> As the queen and king are monogamous, sperm competition does not occur.<ref>{{cite journal|last1=Morrow|first1=E.H.|title=How the sperm lost its tail: the evolution of aflagellate sperm.|journal=Biological Reviews of the Cambridge Philosophical Society|date=2004|volume=79|issue=4|pages=795–814|doi=10.1017/S1464793104006451|pmid=15682871|s2cid=25878093}}</ref>

Termites going through incomplete metamorphosis on the path to becoming alates form a subcaste in certain species of termite, functioning as potential supplementary reproductives. These supplementary reproductives only mature into primary reproductives upon the death of a king or queen, or when the primary reproductives are separated from the colony.<ref name="uflorida">{{cite web |title=Native subterranean termites |url=http://entnemdept.ufl.edu/creatures/urban/termites/native_subterraneans.htm |access-date=8 January 2015 |publisher=University of Florida}}</ref><ref>{{cite web|title=Supplementary reproductive|url=http://www2.hawaii.edu/~entomol/glossary/definition_supplementary_reproductive.htm|archive-url=https://web.archive.org/web/20141030112432/http://www2.hawaii.edu/~entomol/glossary/definition_supplementary_reproductive.htm |archive-date=30 October 2014|publisher=University of Hawaii|access-date=16 September 2015}}</ref> Supplementaries have the ability to replace a dead primary reproductive, and there may also be more than a single supplementary within a colony.<ref name=aus/> Some queens have the ability to switch from sexual reproduction to [[asexual reproduction]]. Studies show that while termite queens mate with the king to produce colony workers, the queens reproduce their replacements ([[Neoteny|neotenic]] queens) [[Parthenogenesis|parthenogenetically]].<ref>{{cite journal|last1=Yashiro|first1=T.|last2=Matsuura|first2=K.|title=Termite queens close the sperm gates of eggs to switch from sexual to asexual reproduction|journal=Proceedings of the National Academy of Sciences |date=2014 |volume=111 |issue=48 |pages=17212–17217 |doi=10.1073/pnas.1412481111 |pmid=25404335 |pmc=4260566 |bibcode=2014PNAS..11117212Y|doi-access=free}}</ref><ref>{{cite journal |last1=Matsuura |first1=K.|last2=Vargo |first2=E.L.|last3=Kawatsu |first3=K. |last4=Labadie |first4=P. E. |last5=Nakano |first5=H.|last6=Yashiro |first6=T. |last7=Tsuji |first7=K. |title=Queen Succession Through Asexual Reproduction in Termites |journal=Science |date=2009 |volume=323 |issue=5922 |pages=1687 |doi=10.1126/science.1169702 |pmid=19325106 |bibcode=2009Sci...323.1687M|s2cid=21785886}}</ref>

The neotropical termite ''Embiratermes neotenicus'' and several other related species produce colonies that contain a primary king accompanied by a primary queen or by up to 200 [[neoteny|neotenic]] queens that had originated through [[thelytoky|thelytokous parthenogenesis]] of a founding primary queen.<ref name="pmid26019158">{{cite journal |vauthors=Fougeyrollas R, Dolejšová K, Sillam-Dussès D, Roy V, Poteaux C, Hanus R, Roisin Y |title=Asexual queen succession in the higher termite Embiratermes neotenicus |journal=Proc. Biol. Sci. |volume=282 |issue=1809 |pages=20150260 |date=June 2015 |pmid=26019158 |pmc=4590441 |doi=10.1098/rspb.2015.0260 }}</ref> The form of [[parthenogenesis]] likely employed maintains [[zygosity|heterozygosity]] in the passage of the [[genome]] from mother to daughter, thus avoiding [[inbreeding depression]].