Editing Termite (section)

==Behaviour and ecology==

===Diet===

[[File:Termite Fecal Pellets.jpg |right |thumb |alt=A dense pile of termite faecal pellets, about 10 centimeters by 20 centimeters by several centimeters in height, which have accumulated on a wooden shelf from termite activity somewhere above the frame of this photograph. |Termite faecal pellets]]

Termites are primarily [[detritivore]]s, consuming dead plants at any level of decomposition. They also play a vital role in the ecosystem by recycling waste material such as dead wood, faeces and plants.{{sfn |Bignell |Roisin |Lo |2010 |pp=13–14}}<ref>{{cite journal |last1=Freymann |first1=B.P. |last2=Buitenwerf |first2=R. |last3=Desouza |first3=O. |last4=Olff |title=The importance of termites (Isoptera) for the recycling of herbivore dung in tropical ecosystems: a review |journal=European Journal of Entomology |date=2008 |volume=105 |issue=2 |pages=165–173 |doi=10.14411/eje.2008.025 |doi-access=free}}</ref><ref>{{cite journal |last1=de Souza |first1=O.F. |last2=Brown |first2=V.K. |title=Effects of habitat fragmentation on Amazonian termite communities |journal=Journal of Tropical Ecology |date=2009 |volume=10 |issue=2 |pages=197–206 |doi=10.1017/S0266467400007847 |s2cid=85721748 }}</ref> Many species eat [[cellulose]], having a specialised midgut that breaks down the fibre.<ref>{{cite journal |last1=Tokuda |first1=G. |last2=Watanabe |first2=H. |last3=Matsumoto |first3=T. |last4=Noda |first4=H. |title=Cellulose digestion in the wood-eating higher termite, ''Nasutitermes takasagoensis'' (Shiraki): distribution of cellulases and properties of endo-beta-1,4-glucanase. |journal=Zoological Science |date=1997 |volume=14 |issue=1 |pages=83–93 |doi=10.2108/zsj.14.83 |pmid=9200983 |s2cid=2877588 |doi-access=free}}</ref> Termites are considered to be a major source (11%) of [[atmospheric methane]], one of the prime [[greenhouse gas]]es, produced from the breakdown of cellulose.<ref>{{cite book |last=Ritter |first=Michael |title=The Physical Environment: an Introduction to Physical Geography |url=http://www4.uwsp.edu/geo/faculty/ritter/geog101/textbook/atmosphere/atmospheric_composition_p2.html |archive-url=https://web.archive.org/web/20070518014844/http://www.uwsp.edu/geo/faculty/ritter/geog101/textbook/atmosphere/atmospheric_composition_p2.html |archive-date=18 May 2007 |year=2006 |publisher=University of Wisconsin |page=450}}</ref> Termites rely primarily upon a symbiotic microbial community that includes bacteria, [[flagellate]] [[protist]]s such as [[metamonad]]s and [[hypermastigid]]s. This community provides the enzymes that digests the cellulose, allowing the insects to absorb the end products for their own use.<ref>{{cite journal |last1=Ikeda-Ohtsubo |first1=W. |last2=Brune |first2=A. |title=Cospeciation of termite gut flagellates and their bacterial endosymbionts: ''Trichonympha'' species and ''Candidatus'' Endomicrobium trichonymphae |journal=Molecular Ecology |date=2009 |volume=18 |issue=2 |pages=332–342 |doi=10.1111/j.1365-294X.2008.04029.x |pmid=19192183 |bibcode=2009MolEc..18..332I |s2cid=28048145}}</ref><ref>{{cite journal |last1=Slaytor |first1=M. |title=Cellulose digestion in termites and cockroaches: What role do symbionts play? |journal=Comparative Biochemistry and Physiology B |date=1992 |volume=103 |issue=4 |pages=775–784 |doi=10.1016/0305-0491(92)90194-V}}</ref> 
[[File:Prot flag trichonymphid 2 reticulotermes.jpg |thumb |right |alt=Trichonymphid flagellate from Reticulotermes. Light microscope image of living cell. |Trichonymphid flagellate from Reticulitermes. Light microscope image of living cell.]]
The microbial ecosystem present in the termite gut contains many species found nowhere else on Earth. Termites hatch without these symbionts present in their guts, and develop them after fed a culture from other termites.<ref name="ibiology.org">{{Cite web |title=The Termite Gut and its Symbiotic Microbes |url=https://www.ibiology.org/ecology/termite-gut/ |website=iBiology |language=en-US |access-date=2020-05-16}}</ref> Gut protozoa, such as ''Trichonympha'', in turn, rely on symbiotic [[bacteria]] embedded on their surfaces to produce some of the necessary [[digestive enzyme]]s. Most higher termites, especially in the family Termitidae, can produce their own [[cellulase]] enzymes, but they rely primarily upon the bacteria. The flagellates have been lost in Termitidae.<ref>{{cite journal |last1=Watanabe |first1=H.. |last2=Noda |first2=H. |last3=Tokuda |first3=G. |last4=Lo |first4=N. |title=A cellulase gene of termite origin |journal=Nature |date=1998 |volume=394 |issue=6691 |pages=330–331 |doi=10.1038/28527 |pmid=9690469 |bibcode = 1998Natur.394..330W |s2cid=4384555}}</ref><ref>{{Cite journal |title=Hidden cellulases in termites: revision of an old hypothesis |journal=Biology Letters |volume=3 |pages=336–339 |doi=10.1098/rsbl.2007.0073 |date= 2007 |issue=3 |pmid=17374589 |last1=Tokuda |first1=G. |last2=Watanabe |first2=H. |pmc=2464699}}</ref><ref>{{cite journal |last1=Li |first1=Z.-Q. |last2=Liu |first2=B.-R. |last3=Zeng |first3=W.-H. |last4=Xiao |first4=W.-L. |last5=Li |first5=Q.-J. |last6=Zhong |first6=J.-H. |title=Character of Cellulase Activity in the Guts of Flagellate-Free Termites with Different Feeding Habits |journal=Journal of Insect Science |date=2013 |volume=13 |issue=37 |pages=37 |doi=10.1673/031.013.3701 |pmid=23895662 |pmc=3738099}}</ref> Researchers have found species of [[Spirochaete |spirochetes]] living in termite guts capable of fixing atmospheric nitrogen to a form usable by the insect.<ref name="ibiology.org"/> Scientists' understanding of the relationship between the termite digestive tract and the microbial endosymbionts is still rudimentary; what is true in all termite species, however, is that the workers feed the other members of the colony with substances derived from the digestion of plant material, either from the [[Insect mouthparts |mouth]] or anus.<ref name=trophal/><ref name= GeethaIyer >[https://scroll.in/magazine/830107/why-indians-worship-the-mound-of-the-much-hated-termite Geetha Iyer ''Scroll.in'' (Mar 09, 2017) Why Indians worship the mound of the much-hated termite] "[The soldier termites] and the reproductive castes obtain their nutrients from the workers through oral or anal trophallaxis."</ref> Judging from closely related bacterial species, it is strongly presumed that the termites' and cockroach's [[gut flora |gut microbiota]] derives from their [[dictyoptera]]n ancestors.<ref>{{cite journal |last1=Dietrich |first1=C. |last2=Kohler |first2=T. |last3=Brune |first3=A. |title=The Cockroach origin of the termite gut microbiota: patterns in bacterial community structure reflect major evolutionary events |journal=Applied and Environmental Microbiology |date=2014 |volume=80 |issue=7 |pages=2261–2269 |doi=10.1128/AEM.04206-13 |pmid=24487532 |pmc=3993134 |bibcode=2014ApEnM..80.2261D}}</ref> Despite primarily consuming decaying plant material as a group, many termite species have been observed to opportunistically feed on dead animals to supplement their dietary needs. Termites are also known to harbor bacteriophages in their gut.<ref>{{Cite journal |last1=Tikhe |first1=Chinmay V. |last2=Husseneder |first2=Claudia |date=2018 |title=Metavirome Sequencing of the Termite Gut Reveals the Presence of an Unexplored Bacteriophage Community |journal=Frontiers in Microbiology |volume=8 |page=2548 |doi=10.3389/fmicb.2017.02548 |doi-access=free |pmid=29354098 |issn=1664-302X |pmc=5759034 }}</ref><ref>{{Cite journal |last1=Tikhe |first1=Chinmay Vijay |last2=Gissendanner |first2=Chris R. |last3=Husseneder |first3=Claudia |date=2018-01-04 |title=Whole-Genome Sequence of the Novel Temperate Enterobacter Bacteriophage Tyrion, Isolated from the Gut of the Formosan Subterranean Termite |journal=Genome Announcements |language=en |volume=6 |issue=1 |doi=10.1128/genomeA.00839-17 |issn=2169-8287 |pmc=5754475 |pmid=29301895}}</ref><ref>{{Cite journal |last1=Tikhe |first1=Chinmay Vijay |last2=Gissendanner |first2=Chris R. |last3=Husseneder |first3=Claudia |date=2018-01-04 |title=Whole-Genome Sequence of the Novel Enterobacter Bacteriophage Arya with an Integrase Pseudogene, Isolated from the Gut of the Formosan Subterranean Termite |journal=Genome Announcements |language=en |volume=6 |issue=1 |doi=10.1128/genomeA.00838-17 |issn=2169-8287 |pmc=5754474 |pmid=29301894}}</ref><ref>{{Cite journal |last1=Pramono |first1=Ajeng K. |last2=Kuwahara |first2=Hirokazu |last3=Itoh |first3=Takehiko |last4=Toyoda |first4=Atsushi |last5=Yamada |first5=Akinori |last6=Hongoh |first6=Yuichi |date=2017 |title=Discovery and Complete Genome Sequence of a Bacteriophage from an Obligate Intracellular Symbiont of a Cellulolytic Protist in the Termite Gut |url=https://www.jstage.jst.go.jp/article/jsme2/32/2/32_ME16175/_article |journal=Microbes and Environments |volume=32 |issue=2 |pages=112–117 |doi=10.1264/jsme2.ME16175 |pmid=28321010 |pmc=5478533 }}</ref><ref>{{Cite journal |last1=Tikhe |first1=Chinmay Vijay |last2=Martin |first2=Thomas M. |last3=Gissendanner |first3=Chris R. |last4=Husseneder |first4=Claudia |date=2015-08-27 |title=Complete Genome Sequence of Citrobacter Phage CVT22 Isolated from the Gut of the Formosan Subterranean Termite, Coptotermes formosanus Shiraki |journal=Genome Announcements |language=en |volume=3 |issue=4 |doi=10.1128/genomeA.00408-15 |issn=2169-8287 |pmc=4505115 |pmid=26184927}}</ref> Some of these bacteriophages likely infect the symbiotic bacteria which play a key role in termite biology. The exact role and function of bacteriophages in the termite gut microbiome is not clearly understood. Termite gut bacteriophages also show similarity to bacteriophages ([[CrAssphage]]) found in the human gut. 

Certain species such as ''[[Gnathamitermes tubiformans]]'' have seasonal food habits. For example, they may preferentially consume Red three-awn (''[[Aristida longiseta]]'') during the summer, Buffalograss (''[[Buchloe dactyloides]]'') from May to August, and blue grama ''[[Bouteloua gracilis]]'' during spring, summer and autumn. Colonies of ''G.&nbsp;tubiformans'' consume less food in spring than they do during autumn when their feeding activity is high.<ref>{{cite journal |last1=Allen |first1=C.T. |last2=Foster |first2=D.E. |last3=Ueckert |first3=D.N. |title=Seasonal Food Habits of a Desert Termite, ''Gnathamitermes tubiformans'', in West Texas |journal=Environmental Entomology |date=1980 |volume=9 |issue=4 |pages=461–466 |doi=10.1093/ee/9.4.461}}</ref>

Various woods differ in their susceptibility to termite attack; the differences are attributed to such factors as moisture content, hardness, and resin and lignin content. In one study, the drywood termite ''Cryptotermes brevis'' strongly preferred [[Populus |poplar]] and [[maple]] woods to other woods that were generally rejected by the termite colony. These preferences may in part have represented conditioned or learned behaviour.<ref>{{cite journal |last1=McMahan |first1=E.A. |title=Studies of Termite Wood-feeding Preferences |journal=Hawaiian Entomological Society |date=1966 |volume=19 |issue=2 |pages=239–250 |url=http://scholarspace.manoa.hawaii.edu/bitstream/handle/10125/10922/19_239-250.pdf?sequence=1 |issn=0073-134X}}</ref>

Some species of termite practice [[fungiculture]]. They maintain a "garden" of specialised fungi of genus ''[[Termitomyces]]'', which are nourished by the excrement of the insects. When the fungi are eaten, their spores pass undamaged through the intestines of the termites to complete the cycle by germinating in the fresh faecal pellets.<ref>{{cite journal |last1=Aanen |first1=D.K. |last2=Eggleton |first2=P. |last3=Rouland-Lefevre |first3=C. |last4=Guldberg-Froslev |first4=T. |last5=Rosendahl |first5=S. |last6=Boomsma |first6=J.J. |title=The evolution of fungus-growing termites and their mutualistic fungal symbionts |journal=Proceedings of the National Academy of Sciences |date=2002 |volume=99 |issue=23 |pages=14887–14892 |doi=10.1073/pnas.222313099 |pmid=12386341 |jstor=3073687 |bibcode=2002PNAS...9914887A |pmc=137514 |doi-access=free}}</ref><ref>{{cite journal |last1=Mueller |first1=U.G. |last2=Gerardo |first2=N. |title=Fungus-farming insects: Multiple origins and diverse evolutionary histories |journal=Proceedings of the National Academy of Sciences |date=2002 |volume=99 |issue=24 |pages=15247–15249 |bibcode=2002PNAS...9915247M |doi=10.1073/pnas.242594799 |pmid=12438688 |pmc=137700 |doi-access=free}}</ref> Molecular evidence suggests that the family [[Macrotermitinae]] developed agriculture about 31 million years ago. It is assumed that more than 90 per cent of dry wood in the semiarid savannah ecosystems of Africa and Asia are reprocessed by these termites. Originally living in the rainforest, fungus farming allowed them to colonise the African savannah and other new environments, eventually expanding into Asia.<ref name="Roberts_et_al_2016">{{cite journal |last1=Roberts |first1=E.M. |last2=Todd |first2=C.N. |last3=Aanen |first3=D.K. |last4=Nobre |first4=T. |last5=Hilbert-Wolf |first5=H.L. |last6=O'Connor |first6=P.M. |last7=Tapanila |first7=L. |last8=Mtelela |first8=C. |last9=Stevens |first9=N.J. |title=Oligocene termite nests with in situ fungus gardens from the Rukwa Rift Basin, Tanzania, support a paleogene African origin for insect agriculture |journal=PLOS ONE |date=2016 |volume=11 |issue=6 |pages=e0156847 |doi=10.1371/journal.pone.0156847 |pmid=27333288 |pmc=4917219 |bibcode=2016PLoSO..1156847R |doi-access=free}}</ref>

Depending on their feeding habits, termites are placed into two groups: the lower termites and higher termites. The lower termites predominately feed on wood. As wood is difficult to digest, termites prefer to consume fungus-infected wood because it is easier to digest and the fungi are high in protein. Meanwhile, the higher termites consume a wide variety of materials, including faeces, [[humus]], grass, leaves and roots.<ref>{{cite journal |last1=Radek |first1=R. |title=Flagellates, bacteria, and fungi associated with termites: diversity and function in nutrition – a review |journal=Ecotropica |date=1999 |volume=5 |pages=183–196 |url=http://large.stanford.edu/publications/coal/references/docs/radek.pdf}}</ref> The gut of the lower termites contains many species of bacteria along with [[protozoa]] and ''[[Holomastigotoides]]'', while the higher termites only have a few species of bacteria with no protozoa.<ref>{{cite journal |last1=Breznak |first1=J.A. |last2=Brune |first2=A. |title=Role of microorganisms in the digestion of lignocellulose by termites |journal=Annual Review of Entomology |date=1993 |volume=39 |issue=1 |pages=453–487 |doi=10.1146/annurev.en.39.010194.002321}}</ref>

===Predators===
[[File:Crab Spider (Thomisidae) and Winged Termite prey (12640038823).jpg |thumbnail |[[Crab spider]] with a captured alate]]
Termites are consumed by a wide variety of [[Predation |predators]]. One termite species alone, ''[[Hodotermes |Hodotermes mossambicus]]'', was reported (1990) in the stomach contents of 65 [[bird]]s and 19 [[mammal]]s.<ref>{{cite journal |last1=Kok |first1=O.B. |last2=Hewitt |first2=P.H. |title=Bird and mammal predators of the harvester termite ''Hodotermes mossambicus'' (Hagen) in semi-arid regions of South Africa |journal=South African Journal of Science |date=1990 |volume=86 |issue=1 |pages=34–37 |issn=0038-2353}}</ref> [[Arthropod]]s such as [[ants]],<ref name=HW1990>{{cite book |last1=Hölldobler |first1=B. |last2=Wilson |first2=E.O. |title=The Ants |date=1990 |publisher=Belknap Press of Harvard University Press |pages=[https://archive.org/details/ants0000hlld/page/559 559–566] |location=Cambridge, Massachusetts |isbn=978-0-674-04075-5 |title-link=The Ants}}</ref><ref name='ant_fight_termite'>{{cite journal |last1=Culliney |first1=T.W. |last2=Grace |first2=J.K. |title=Prospects for the biological control of subterranean termites (Isoptera: Rhinotermitidae), with special reference to ''Coptotermes formosanus'' |journal=Bulletin of Entomological Research |date=2000 |volume=90 |issue=1 |pages=9–21 |doi=10.1017/S0007485300000663 |doi-broken-date=4 April 2025 |pmid=10948359}}</ref> [[centipede]]s, [[cockroach]]es, [[Cricket (insect) |crickets]], [[Dragonfly |dragonflies]], [[scorpion]]s and [[spider]]s,<ref>{{cite journal |last1=Dean |first1=W.R.J. |last2=Milton |first2=S.J. |title=Plant and invertebrate assemblages on old fields in the arid southern Karoo, South Africa |journal=African Journal of Ecology |date=1995 |volume=33 |issue=1 |pages=1–13 |doi=10.1111/j.1365-2028.1995.tb00777.x |bibcode=1995AfJEc..33....1D }}</ref> [[reptile]]s such as [[lizard]]s,<ref>{{cite book |last1=Wade |first1=W.W. |title=Ecology of Desert Systems |date=2002 |publisher=Elsevier |location=Burlington |isbn=978-0-08-050499-5 |page=216}}</ref> and [[amphibian]]s such as [[frog]]s<ref>{{cite book |last1=Reagan |first1=D.P. |last2=Waide |first2=R.B. |title=The food web of a tropical rain forest |date=1996 |publisher=University of Chicago Press |location=Chicago |isbn=978-0-226-70599-6 |page=[https://archive.org/details/foodweboftropica0000unse/page/294 294] |url=https://archive.org/details/foodweboftropica0000unse/page/294}}</ref> and [[toad]]s consume termites, with two [[spider]]s in the family [[Ammoxenidae]] being specialist termite predators.<ref name=Services2013>{{cite book |last1=Bardgett |first1=R.D. |last2=Herrick |first2=J.E. |last3=Six |first3=J. |last4=Jones |first4=T.H. |last5=Strong |first5=D.R. |last6=van der Putten |first6=W.H. |title=Soil ecology and ecosystem services |date=2013 |publisher=Oxford University Press |location=Oxford |isbn=978-0-19-968816-6 |page=178 |edition=1st}}</ref>{{sfn |Bignell |Roisin |Lo |2010 |p=509}}<ref>{{cite book |last1=Choe |first1=J.C. |last2=Crespi |first2=B.J. |title=The evolution of social behavior in insects and arachnids |url=https://archive.org/details/evolutionsocialb00choe |url-access=limited |date=1997 |publisher=Cambridge university press |location=Cambridge |isbn=978-0-521-58977-2 |page=[https://archive.org/details/evolutionsocialb00choe/page/n84 76] |edition=1st}}</ref> Other predators include [[aardvark]]s, [[Aardwolf |aardwolves]], [[anteater]]s, [[bat]]s, [[bear]]s, [[Macrotis |bilbies]], many [[bird]]s, [[echidnas]], [[fox]]es, [[galago]]s, [[numbat]]s, [[Mouse |mice]] and [[pangolin]]s.<ref name=Services2013/><ref name=wilson2014>{{cite book |last1=Abe |first1=Y. |last2=Bignell |first2=D.E. |last3=Higashi |first3=T. |title=Termites: Evolution, Sociality, Symbioses, Ecology |date=2014 |publisher=Springer |isbn=978-94-017-3223-9 |doi=10.1007/978-94-017-3223-9 |pages=124–149 |s2cid=30804981}}</ref><ref>{{cite journal |last1=Wilson |first1=D.S. |last2=Clark |first2=A.B. |title=Above ground defence in the harvester termite, ''Hodotermes mossambicus'' |journal=Journal of the Entomological Society of South Africa |date=1977 |volume=40 |pages=271–282}}</ref><ref>{{cite book |last1=Lavelle |first1=P. |last2=Spain |first2=A.V. |title=Soil ecology |url=https://archive.org/details/soilecology00lave |url-access=limited |date=2001 |publisher=Kluwer Academic |location=Dordrecht |isbn=978-0-306-48162-8 |edition=2nd |page=[https://archive.org/details/soilecology00lave/page/n340 316]}}</ref> The [[aardwolf]] is an [[insectivore |insectivorous]] [[mammal]] that primarily feeds on termites; it locates its food by sound and also by detecting the scent secreted by the soldiers; a single aardwolf is capable of consuming thousands of termites in a single night by using its long, sticky tongue.<ref>{{cite encyclopedia |last1=Richardson |first1=P.K.R. |last2=Bearder |first2=S.K. |editor1-last=MacDonald |editor1-first=D. |encyclopedia=The Encyclopedia of Mammals |title=The Hyena Family |isbn=978-0-87196-871-5 |publisher=Facts on File Publication |location=New York, NY |year=1984 |pages=[https://archive.org/details/encyclopediaofma00mals_0/page/158 158–159] |url-access=registration |url=https://archive.org/details/encyclopediaofma00mals_0/page/158}}</ref><ref>{{cite book |last1=Mills |first1=G. |last2=Harvey |first2=M. |title=African Predators |publisher=Smithsonian Institution Press |location=Washington, D.C. |year=2001 |isbn=978-1-56098-096-4 |page=71}}</ref> [[Sloth bears]] break open mounds to consume the nestmates, while [[Common chimpanzee |chimpanzees]] have [[Tool use by animals#Primates |developed tools]] to "fish" termites from their nest. Wear pattern analysis of bone tools used by the early [[hominin]] ''[[Paranthropus robustus]]'' suggests that they used these tools to dig into termite mounds.<ref>{{cite journal |last1=d'Errico |first1=F. |last2=Backwell |first2=L. |title=Assessing the function of early hominin bone tools |journal=Journal of Archaeological Science |date=2009 |volume=36 |issue=8 |pages=1764–1773 |doi=10.1016/j.jas.2009.04.005 |bibcode=2009JArSc..36.1764D |url=https://www.researchgate.net/publication/229364355}}</ref>

[[File:Megaponera analis major killing macrotermes soldier.jpg |thumbnail |left |A Matabele ant (''Megaponera analis'') kills a ''Macrotermes bellicosus'' termite soldier during a raid.]]
Among all predators, ants are the greatest enemy to termites.<ref name=HW1990/><ref name='ant_fight_termite'/> Some ant genera are specialist predators of termites. For example, ''[[Megaponera]]'' is a strictly termite-eating (termitophagous) genus that perform raiding activities, some lasting several hours.<ref name="lepage">{{cite journal |last1=Lepage |first1=M.G. |title=Étude de la prédation de ''Megaponera foetens'' (F.) sur les populations récoltantes de Macrotermitinae dans un ecosystème semi-aride (Kajiado-Kenya) |journal=Insectes Sociaux |date=1981 |volume=28 |issue=3 |pages=247–262 |doi=10.1007/BF02223627 |s2cid=28763771 |language=es}}</ref><ref name=Levieux1966>{{cite journal |last1=Levieux |first1=J. |title=Note préliminaire sur les colonnes de chasse de ''Megaponera fœtens'' F. (Hyménoptère Formicidæ) |journal=Insectes Sociaux |date=1966 |volume=13 |issue=2 |pages=117–126 |doi=10.1007/BF02223567 |s2cid=2031222 |language=fr}}</ref> ''[[Paltothyreus |Paltothyreus tarsatus]]'' is another termite-raiding species, with each individual stacking as many termites as possible in its [[Mandible (insect mouthpart)|mandibles]] before returning home, all the while recruiting additional nestmates to the raiding site through chemical trails.<ref name=HW1990/> The Malaysian basicerotine ants ''[[Eurhopalothrix |Eurhopalothrix heliscata]]'' uses a different strategy of termite hunting by pressing themselves into tight spaces, as they hunt through rotting wood housing termite colonies. Once inside, the ants seize their prey by using their short but sharp mandibles.<ref name=HW1990/> ''[[Tetramorium |Tetramorium uelense]]'' is a specialised predator species that feeds on small termites. A scout recruits 10–30 workers to an area where termites are present, killing them by immobilising them with their stinger.<ref>{{cite journal |last1=Longhurst |first1=C. |last2=Baker |first2=R. |last3=Howse |first3=P.E. |title=Chemical crypsis in predatory ants |journal=Experientia |date=1979 |volume=35 |issue=7 |pages=870–872 |doi=10.1007/BF01955119 |s2cid=39854106}}</ref> ''[[Centromyrmex]]'' and ''[[Iridomyrmex]]'' colonies sometimes nest in [[termite mound]]s, and so the termites are preyed on by these ants. No evidence for any kind of relationship (other than a predatory one) is known.<ref name=wheeler1936>{{cite journal |last1=Wheeler |first1=W.M. |title=Ecological relations of Ponerine and other ants to termites |journal=Proceedings of the American Academy of Arts and Sciences |date=1936 |volume=71 |issue=3 |pages=159–171 |doi=10.2307/20023221 |jstor=20023221 |url=https://zenodo.org/record/25265}}</ref><ref name=Shattuck>{{cite journal |last1=Shattuck |first1=S.O. |last2=Heterick |first2=B.E. |title=Revision of the ant genus ''Iridomyrmex'' (Hymenoptera : Formicidae) |date=2011 |journal=Zootaxa |volume=2845 |pages=1–74 |doi=10.11646/zootaxa.2743.1.1 |isbn=978-1-86977-676-3 |url=http://www.antwiki.org/wiki/images/a/ab/Heterick_%26_Shattuck.pdf |issn=1175-5334}}</ref> Other ants, including ''[[Acanthostichus]]'', ''[[Camponotus]]'', ''[[Crematogaster]]'', ''[[Cylindromyrmex]]'', ''[[Leptogenys]]'', ''[[Odontomachus]]'', ''[[Ophthalmopone]]'', ''[[Pachycondyla]]'', ''[[Rhytidoponera]]'', ''[[Fire ant |Solenopsis]]'' and ''[[Wasmannia]]'', also prey on termites.<ref name=wilson2014/><ref name=HW1990/><ref>{{cite journal |last1=Traniello |first1=J.F.A. |title=Enemy deterrence in the recruitment strategy of a termite: Soldier-organized foraging in ''Nasutitermes costalis'' |journal=Proceedings of the National Academy of Sciences |date=1981 |volume=78 |issue=3 |pages=1976–1979 |doi=10.1073/pnas.78.3.1976 |pmid=16592995 |pmc=319259 |bibcode=1981PNAS...78.1976T |doi-access=free}}</ref> Specialized subterranean species of army ants such as ones in the genus ''[[Dorylus]]'' are known to commonly predate on young ''[[Macrotermes]]'' colonies.<ref>{{cite journal |last1=Schöning |first1=C. |last2=Moffett |first2=M.W. |title=Driver Ants Invading a Termite Nest: why do the most catholic predators of all seldom take this abundant prey? |journal=Biotropica |date=2007 |volume=39 |issue=5 |pages=663–667 |doi=10.1111/j.1744-7429.2007.00296.x |bibcode=2007Biotr..39..663S |s2cid=13689479 |url=http://www.doctorbugs.com/dorylus.pdf |access-date=2015-09-20 |archive-date=2015-11-12 |archive-url=https://web.archive.org/web/20151112035412/http://www.doctorbugs.com/Dorylus.pdf |url-status=dead}}</ref>

Ants are not the only invertebrates that perform raids. Many [[Spheciformes |sphecoid wasp]]s and several species including ''[[Polybia]]'' and ''[[Angiopolybia pallens |Angiopolybia]]'' are known to raid termite mounds during the termites' nuptial flight.<ref>{{cite journal |last1=Mill |first1=A.E. |title=Observations on Brazilian termite alate swarms and some structures used in the dispersal of reproductives (Isoptera: Termitidae) |journal=Journal of Natural History |date=1983 |volume=17 |issue=3 |pages=309–320 |doi=10.1080/00222938300770231 |bibcode=1983JNatH..17..309M }}</ref>

===Parasites, pathogens, and viruses===
Termites are less likely to be attacked by parasites than bees, wasps and ants, as they are usually well protected in their mounds.{{sfn |Schmid-Hempel |1998 |p=61}}{{sfn |Schmid-Hempel |1998 |p=75}} Nevertheless, termites are infected by a variety of parasites. Some of these include dipteran flies,<ref>{{cite book |last1=Wilson |first1=E.O. |title=The Insect Societies |date=1971 |publisher=Belknap Press of Harvard University Press |location=Cambridge, Massachusetts |isbn=978-0-674-45495-8 |page=398 |edition=5th |volume=76}}</ref> ''[[Pyemotes]]'' mites, and a large number of [[nematode]] parasites. Most nematode parasites are in the order [[Rhabditida]];{{sfn |Schmid-Hempel |1998 |p=59}} others are in the genus ''[[Mermis]]'', ''[[Diplogaster aerivora]]'' and ''[[Heterakis gallinarum |Harteria gallinarum]]''.{{sfn |Schmid-Hempel |1998 |pp=301–302}} Under imminent threat of an attack by parasites, a colony may migrate to a new location.{{sfn |Schmid-Hempel |1998 |p=19}} Certain fungal pathogens such as ''[[Aspergillus nomius]]'' and ''[[Metarhizium anisopliae]]'' are, however, major threats to a termite colony as they are not host-specific and may infect large portions of the colony;<ref>{{cite journal |last1=Weiser |first1=J. |last2=Hrdy |first2=I. |title=Pyemotes – mites as parasites of termites |journal=Zeitschrift für Angewandte Entomologie |date=2009 |volume=51 |issue=1–4 |pages=94–97 |doi=10.1111/j.1439-0418.1962.tb04062.x}}</ref><ref name=resource>{{cite journal |last1=Chouvenc |first1=T. |last2=Efstathion |first2=C.A. |last3=Elliott |first3=M.L. |last4=Su |first4=N.Y. |title=Resource competition between two fungal parasites in subterranean termites. |journal=Die Naturwissenschaften |date=2012 |volume=99 |issue=11 |pages=949–58 |doi=10.1007/s00114-012-0977-2 |pmid=23086391 |bibcode = 2012NW.....99..949C |s2cid=16393629}}</ref> transmission usually occurs via direct physical contact.{{sfn |Schmid-Hempel |1998 |pp=38, 102}} ''M.&nbsp;anisopliae'' is known to weaken the termite immune system. Infection with ''A.&nbsp;nomius'' only occurs when a colony is under great stress. Over 34 fungal species are known to live as parasites on the exoskeleton of termites, with many being host-specific and only causing indirect harm to their host.<ref>{{cite journal |last1=Wilson |first1=Megan |last2=Barden |first2=Phillip |last3=Ware |first3=Jessica |title=A Review of Ectoparasitic Fungi Associated With Termites |journal=Annals of the Entomological Society of America |date=2021-04-30 |volume=114 |issue=4 |pages=373–396 |doi=10.1093/aesa/saab001 |doi-access=free }}</ref>

Termites are infected by viruses including [[Entomopoxvirinae]] and the [[Nuclear Polyhedrosis Virus]].<ref>{{cite journal |last1=Chouvenc |first1=T. |last2=Mullins |first2=A.J. |last3=Efstathion |first3=C.A. |last4=Su |first4=N.-Y. |title=Virus-like symptoms in a termite (Isoptera: Kalotermitidae) field colony |journal=Florida Entomologist |date=2013 |volume=96 |issue=4 |pages=1612–1614 |doi=10.1653/024.096.0450 |s2cid=73570814 |doi-access=free}}</ref><ref>{{cite journal |last1=Al Fazairy |first1=A.A. |last2=Hassan |first2=F.A. |title=Infection of Termites by ''Spodoptera littoralis'' Nuclear Polyhedrosis Virus |journal=International Journal of Tropical Insect Science |date=2011 |volume=9 |issue=1 |pages=37–39 |doi=10.1017/S1742758400009991 |s2cid=84743428}}</ref>

===Locomotion and foraging===
Because the worker and soldier castes lack wings and thus never fly, and the reproductives use their wings for just a brief amount of time, termites predominantly rely upon their legs to move about.{{sfn |Bignell |Roisin |Lo |2010 |p=11}}

Foraging behaviour depends on the type of termite. For example, certain species feed on the wood structures they inhabit, and others harvest food that is near the nest.<ref>{{cite book |last1=Traniello |first1=J.F.A. |last2=Leuthold |first2=R.H. |title=Behavior and Ecology of Foraging in Termites |date=2000 |pages=141–168 |doi=10.1007/978-94-017-3223-9_7 |isbn=978-94-017-3223-9 |publisher=Springer Netherlands}}</ref> Most workers are rarely found out in the open, and do not forage unprotected; they rely on sheeting and runways to protect them from predators.{{sfn |Bignell |Roisin |Lo |2010 |p=13}} Subterranean termites construct tunnels and galleries to look for food, and workers who manage to find food sources recruit additional nestmates by depositing a phagostimulant pheromone that attracts workers.<ref>{{cite journal |last1=Reinhard |first1=J. |last2=Kaib |first2=M. |title=Trail communication during foraging and recruitment in the subterranean termite ''Reticulitermes santonensis'' De Feytaud (Isoptera, Rhinotermitidae) |journal=Journal of Insect Behavior |date=2001 |volume=14 |issue=2 |pages=157–171 |doi=10.1023/A:1007881510237 |bibcode=2001JIBeh..14..157R |s2cid=40887791}}</ref> Foraging workers use semiochemicals to communicate with each other,<ref name=commu/> and workers who begin to forage outside of their nest release trail pheromones from their sternal glands.<ref>{{cite journal |last1=Costa-Leonardo |first1=A.M. |title=Morphology of the sternal gland in workers of ''Coptotermes gestroi'' (Isoptera, Rhinotermitidae). |journal=Micron |date=2006 |volume=37 |issue=6 |pages=551–556 |doi=10.1016/j.micron.2005.12.006 |pmid=16458523}}</ref> In one species, ''[[Nasutitermes costalis]]'', there are three phases in a foraging expedition: first, soldiers scout an area. When they find a food source, they communicate to other soldiers and a small force of workers starts to emerge. In the second phase, workers appear in large numbers at the site. The third phase is marked by a decrease in the number of soldiers present and an increase in the number of workers.<ref>{{cite journal |last1=Traniello |first1=J.F. |last2=Busher |first2=C. |title=Chemical regulation of polyethism during foraging in the neotropical termite ''Nasutitermes costalis'' |journal=Journal of Chemical Ecology |date=1985 |volume=11 |issue=3 |pages=319–32 |doi=10.1007/BF01411418 |pmid=24309963 |bibcode=1985JCEco..11..319T |s2cid=27799126}}</ref> Isolated termite workers may engage in [[Lévy flight]] behaviour as an optimised strategy for finding their nestmates or foraging for food.<ref>{{cite journal |last1=Miramontes |first1=O. |last2=DeSouza |first2=O. |last3=Paiva |first3=L.R. |last4=Marins |first4=A. |last5=Orozco |first5=S. |last6=Aegerter |first6=C.M. |title=Lévy flights and self-similar exploratory behaviour of termite workers: beyond model fitting |journal=PLOS ONE |date=2014 |volume=9 |issue=10 |pages=e111183 |doi=10.1371/journal.pone.0111183 |pmid=25353958 |pmc=4213025 |bibcode=2014PLoSO...9k1183M |arxiv = 1410.0930 |doi-access=free}}</ref>

===Competition===
Competition between two colonies always results in [[agonistic behaviour]] towards each other, resulting in fights. These fights can cause mortality on both sides and, in some cases, the gain or loss of territory.<ref>{{cite journal |last1=Jost |first1=C. |last2=Haifig |first2=I. |last3=de Camargo-Dietrich |first3=C.R.R. |last4=Costa-Leonardo |first4=A.M. |title=A comparative tunnelling network approach to assess interspecific competition effects in termites |journal=Insectes Sociaux |date=2012 |volume=59 |issue=3 |pages=369–379 |doi=10.1007/s00040-012-0229-7 |s2cid=14885485}}</ref><ref>{{cite journal |last1=Polizzi |first1=J.M. |last2=Forschler |first2=B.T. |title=Intra- and interspecific agonism in ''Reticulitermes flavipes'' (Kollar) and ''R. virginicus'' (Banks) and effects of arena and group size in laboratory assays |journal=Insectes Sociaux |date=1998 |volume=45 |issue=1 |pages=43–49 |doi=10.1007/s000400050067 |s2cid=36235510}}</ref> "Cemetery pits" may be present, where the bodies of dead termites are buried.<ref>{{cite journal |last1=Darlington |first1=J.P.E.C. |title=The underground passages and storage pits used in foraging by a nest of the termite ''Macrotermes michaelseni'' in Kajiado, Kenya |journal=Journal of Zoology |date=1982 |volume=198 |issue=2 |pages=237–247 |doi=10.1111/j.1469-7998.1982.tb02073.x}}</ref>

Studies show that when termites encounter each other in foraging areas, some of the termites deliberately block passages to prevent other termites from entering.<ref name=commu/><ref>{{cite journal |last1=Cornelius |first1=M.L. |last2=Osbrink |first2=W.L. |title=Effect of soil type and moisture availability on the foraging behavior of the Formosan subterranean termite (Isoptera: Rhinotermitidae). |journal=Journal of Economic Entomology |date=2010 |volume=103 |issue=3 |pages=799–807 |doi=10.1603/EC09250 |pmid=20568626 |s2cid=23173060 |doi-access=free}}</ref> Dead termites from other colonies found in exploratory tunnels leads to the isolation of the area and thus the need to construct new tunnels.<ref>{{cite journal |last1=Toledo Lima |first1=J. |last2=Costa-Leonardo |first2=A.M. |title=Subterranean termites (Isoptera: Rhinotermitidae): Exploitation of equivalent food resources with different forms of placement |journal=Insect Science |date=2012 |volume=19 |issue=3 |pages=412–418 |doi=10.1111/j.1744-7917.2011.01453.x |bibcode=2012InsSc..19..412T |s2cid=82046133}}</ref> Conflict between two competitors does not always occur. For example, though they might block each other's passages, colonies of ''Macrotermes bellicosus'' and ''Macrotermes subhyalinus'' are not always aggressive towards each other.<ref>{{cite journal |last1=Jmhasly |first1=P. |last2=Leuthold |first2=R.H. |title=Intraspecific colony recognition in the termites ''Macrotermes subhyalinus'' and ''Macrotermes bellicosus'' (Isoptera, Termitidae) |journal=Insectes Sociaux |date=1999 |volume=46 |issue=2 |pages=164–170 |doi=10.1007/s000400050128 |s2cid=23037986}}</ref> Suicide cramming is known in ''[[Coptotermes formosanus]]''. Since ''C.&nbsp;formosanus'' colonies may get into physical conflict, some termites squeeze tightly into foraging tunnels and die, successfully blocking the tunnel and ending all agonistic activities.<ref>{{cite journal |last1=Messenger |first1=M.T. |last2=Su |first2=N.Y. |title=Agonistic behavior between colonies of the Formosan subterranean termite (Isoptera: Rhinotermitidae) from Louis Armstrong Park, New Orleans, Louisiana |journal=Sociobiology |date=2005 |volume=45 |issue=2 |pages=331–345}}</ref>

Among the reproductive caste, neotenic queens may compete with each other to become the dominant queen when there are no primary reproductives. This struggle among the queens leads to the elimination of all but a single queen, which, with the king, takes over the colony.<ref>{{cite journal |last1=Korb |first1=J. |last2=Weil |first2=T. |last3=Hoffmann |first3=K. |last4=Foster |first4=K.R. |last5=Rehli |first5=M. |title=A gene necessary for reproductive suppression in termites |journal=Science |date=2009 |volume=324 |issue=5928 |pages=758 |doi=10.1126/science.1170660 |pmid=19423819 |bibcode=2009Sci...324..758K |s2cid=31608071}}</ref>

Ants and termites may compete with each other for nesting space. In particular, ants that prey on termites usually have a negative impact on arboreal nesting species.<ref name=arb>{{cite journal |last1=Mathew |first1=T.T.G. |last2=Reis |first2=R. |last3=DeSouza |first3=O. |last4=Ribeiro |first4=S.P. |title=Predation and interference competition between ants (Hymenoptera: Formicidae) and arboreal termites (Isoptera: Termitidae) |journal=Sociobiology |date=2005 |volume=46 |issue=2 |pages=409–419 |url=http://www.repositorio.ufop.br/bitstream/123456789/4835/1/ARTIGO_PredationInterferenceCompetition.pdf}}</ref>

===Communication===
[[File:Termites (Nasutitermes sp.) (8439859723).jpg |thumbnail |Hordes of ''Nasutitermes'' on a march for food, following and leaving trail pheromones]]
Most termites are blind, so communication primarily occurs through chemical, mechanical and pheromonal cues.<ref name=Leonardo>{{cite journal |last1=Costa-Leonardo |first1=A.M. |last2=Haifig |first2=I. |title=Pheromones and exocrine glands in Isoptera |journal=Vitamins and Hormones |date=2010 |volume=83 |pages=521–549 |doi=10.1016/S0083-6729(10)83021-3 |pmid=20831960 |isbn=9780123815163}}</ref><ref name=commu>{{cite book |last1=Costa-Leonardo |first1=A.M. |last2=Haifig |first2=I. |title=''Termite communication during different behavioral activities'' in Biocommunication of Animals |date=2013 |pages=161–190 |doi=10.1007/978-94-007-7414-8_10 |publisher=Springer Netherlands |isbn=978-94-007-7413-1}}</ref> These methods of communication are used in a variety of activities, including foraging, locating reproductives, construction of nests, recognition of nestmates, nuptial flight, locating and fighting enemies, and defending the nests.<ref name=Leonardo/><ref name=commu/> The most common way of communicating is through antennation.<ref name=commu/> A number of pheromones are known, including contact pheromones (which are transmitted when workers are engaged in trophallaxis or grooming) and [[Pheromone#Alarm |alarm]], [[Trail pheromone |trail]] and [[sex pheromone]]s. The alarm pheromone and other defensive chemicals are secreted from the frontal gland. Trail pheromones are secreted from the sternal gland, and sex pheromones derive from two glandular sources: the sternal and tergal glands.<ref name=Leonardo/> When termites go out to look for food, they forage in columns along the ground through vegetation. A trail can be identified by the faecal deposits or runways that are covered by objects. Workers leave pheromones on these trails, which are detected by other nestmates through olfactory receptors.<ref name=Britannica/> Termites can also communicate through mechanical cues, vibrations, and physical contact.<ref name=Britannica>{{cite encyclopedia |last1=Krishna |first1=K. |title=Termite |url=http://www.britannica.com/animal/termite |encyclopedia=Encyclopædia Britannica |access-date=11 September 2015}}</ref><ref name=commu/> These signals are frequently used for alarm communication or for evaluating a food source.<ref name=commu/><ref>{{cite journal |last1=Evans |first1=T.A. |last2=Inta |first2=R. |last3=Lai |first3=J.C.S. |last4=Lenz |first4=M. |title=Foraging vibration signals attract foragers and identify food size in the drywood termite, ''Cryptotermes secundus'' |journal=Insectes Sociaux |date=2007 |volume=54 |issue=4 |pages=374–382 |doi=10.1007/s00040-007-0958-1 |s2cid=40214049}}</ref>

When termites construct their nests, they use predominantly indirect communication. No single termite would be in charge of any particular construction project. Individual termites react rather than think, but at a group level, they exhibit a sort of collective cognition. Specific structures or other objects such as pellets of soil or pillars cause termites to start building. The termite adds these objects onto existing structures, and such behaviour encourages building behaviour in other workers. The result is a self-organised process whereby the information that directs termite activity results from changes in the environment rather than from direct contact among individuals.<ref name=commu/>

Termites can distinguish nestmates and non-nestmates through chemical communication and gut symbionts: chemicals consisting of hydrocarbons released from the cuticle allow the recognition of alien termite species.<ref>{{cite journal |last1=Costa-Leonardo |first1=A.M. |last2=Casarin |first2=F.E. |last3=Lima |first3=J.T. |title=Chemical communication in isoptera |journal=Neotropical Entomology |date=2009 |volume=38 |issue=1 |pages=747–52 |doi=10.1590/S1519-566X2009000100001 |pmid=19347093 |doi-access=free |hdl=11449/19749 |hdl-access=free}}</ref><ref>{{cite journal |last1=Richard |first1=F.-J. |last2=Hunt |first2=J.H. |title=Intracolony chemical communication in social insects |journal=Insectes Sociaux |date=2013 |volume=60 |issue=3 |pages=275–291 |doi=10.1007/s00040-013-0306-6 |s2cid=8108234 |url=http://www4.ncsu.edu/~jhhunt/Richard%20and%20Hunt.pdf |access-date=2015-10-08 |archive-date=2016-03-04 |archive-url=https://web.archive.org/web/20160304114501/http://www4.ncsu.edu/~jhhunt/Richard%20and%20Hunt.pdf |url-status=dead}}</ref> Each colony has its own distinct odour. This odour is a result of genetic and environmental factors such as the termites' diet and the composition of the bacteria within the termites' intestines.<ref>{{cite journal |last1=Dronnet |first1=S. |last2=Lohou |first2=C. |last3=Christides |first3=J.P. |last4=Bagnères |first4=A.G. |title=Cuticular hydrocarbon composition reflects genetic relationship among colonies of the introduced termite ''Reticulitermes santonensis'' Feytaud |journal=Journal of Chemical Ecology |date=2006 |volume=32 |issue=5 |pages=1027–1042 |doi=10.1007/s10886-006-9043-x |pmid=16739021 |bibcode=2006JCEco..32.1027D |s2cid=23956394}}</ref>

===Defence===<!-- Note this spelling is correct in UK English, if you came here to change this to Defense, please note that articles on Wikipedia choose between US and UK spelling based on priority. See WP:ENGVAR.-->
{{see also |Defense in insects |l1=Insect defences}}
[[File:Termites rush to damaged portion of mound.jpg |thumbnail |180px |alt=To demonstrate termite repair behaviour, a hole was bored into a termite nest. Over a dozen worker termites with pale heads are visible in this close-up photo, most facing the camera as they engage in repair activities from the inside of the hole. About a dozen soldier termites with orange heads are also visible, some facing outwards from the hole, others patrolling the surrounding area. |Termites rush to a damaged area of the nest.]]
Termites rely on alarm communication to defend a colony.<ref name=commu/> Alarm pheromones can be released when the nest has been breached or is being attacked by enemies or potential pathogens. Termites always avoid nestmates infected with ''[[Metarhizium anisopliae]]'' spores, through vibrational signals released by infected nestmates.<ref name=rosengaus1999>{{cite journal |last1=Rosengaus |first1=R.B. |last2=Traniello |first2=J.F.A. |last3=Chen |first3=T. |last4=Brown |first4=J.J. |last5=Karp |first5=R.D. |title=Immunity in a social insect |journal=Naturwissenschaften |date=1999 |volume=86 |issue=12 |pages=588–591 |doi=10.1007/s001140050679 |bibcode=1999NW.....86..588R |s2cid=10769345 }}</ref> Other methods of defence include headbanging and secretion of fluids from the frontal gland and defecating faeces containing alarm pheromones.<ref name=commu/><ref>{{cite journal |last1=Wilson |first1=D.S. |title=Above ground predator defense in the harvester termite, ''Hodotermes mossambicus'' (Hagen) |journal=Journal of the Entomological Society of Southern Africa |date=1977 |volume=40 |pages=271–282}}</ref>

In some species, some soldiers block tunnels to prevent their enemies from entering the nest, and they may deliberately rupture themselves as an act of defence.<ref>{{cite book |last1=Belbin |first1=R.M. |title=The Coming Shape of Organization |date=2013 |publisher=Routledge | location=New York |isbn=978-1-136-01553-3 |page=27}}</ref> In cases where the intrusion is coming from a breach that is larger than the soldier's head, soldiers form a [[Phalanx formation |phalanx]]-like formation around the breach and bite at intruders.<ref name=wilson/> If an invasion carried out by ''[[Megaponera analis]]'' is successful, an entire colony may be destroyed, although this scenario is rare.<ref name=wilson>{{cite book |last1=Wilson |first1=E.O. |author-link=E. O. Wilson |title=A window on eternity: a biologist's walk through Gorongosa National Park |date=2014 |publisher=Simon & Schuster |isbn=978-1-4767-4741-5 |pages=85, 90 |edition=First}}</ref>

To termites, any breach of their tunnels or nests is a cause for alarm. When termites detect a potential breach, the soldiers usually bang their heads, apparently to attract other soldiers for defence and to recruit additional workers to repair any breach.<ref name=Britannica/> Additionally, an alarmed termite bumps into other termites which causes them to be alarmed and to leave pheromone trails to the disturbed area, which is also a way to recruit extra workers.<ref name=Britannica/>

[[File:Nasute termite soldiers.png |thumbnail |left |Nasute termite soldiers on rotten wood]]
The pantropical subfamily [[Nasutitermitinae]] has a specialised caste of soldiers, known as nasutes, that have the ability to exude noxious liquids through a [[fontanellar gun |horn-like frontal projection]] that they use for defence.<ref>{{cite journal |last1=Miura |first1=T. |last2=Matsumoto |first2=T. |title=Soldier morphogenesis in a nasute termite: discovery of a disc-like structure forming a soldier nasus |journal=Proceedings of the Royal Society B: Biological Sciences |date=2000 |volume=267 |issue=1449 |pages=1185–1189 |doi=10.1098/rspb.2000.1127 |pmc=1690655 |pmid=10902684}}</ref> Nasutes have lost their mandibles through the course of evolution and must be fed by workers.<ref name=biosyn/> A wide variety of [[monoterpene]] hydrocarbon [[solvent]]s have been identified in the liquids that nasutes secrete.<ref>{{cite journal |last1=Prestwich |first1=G.D. |last2=Chen |first2=D. |title=Soldier defense secretions of ''Trinervitermes bettonianus'' (Isoptera, Nasutitermitinae): Chemical variation in allopatric populations |journal=Journal of Chemical Ecology |date=1981 |volume=7 |issue=1 |pages=147–157 |doi=10.1007/BF00988642 |pmid=24420434 |bibcode=1981JCEco...7..147P |s2cid=27654745 }}</ref> Similarly, [[Formosan subterranean termite]]s have been known to secrete [[naphthalene]] to protect their nests.<ref>{{Cite journal |last1=Chen |first1=J. |last2=Henderson |first2=G. |last3=Grimm |first3=C. C. |last4=Lloyd |first4=S. W. |last5=Laine |first5=R. A. |date=1998-04-09 |title=Termites fumigate their nests with naphthalene |journal=Nature |volume=392 |issue=6676 |pages=558–559 |doi=10.1038/33305 |bibcode=1998Natur.392..558C |s2cid=4419882 }}</ref>

Soldiers of the species ''[[Globitermes sulphureus]]'' commit suicide by [[autothysis]]&nbsp;– rupturing a large gland just beneath the surface of their cuticles. The thick, yellow fluid in the gland becomes very sticky on contact with the air, entangling ants or other insects that are trying to invade the nest.<ref name="Extraordinary_Animals" >{{citation |title=Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals |first=Ross |last=Piper |year=2007 |page=26 |isbn=978-0-313-33922-6 |url=https://books.google.com/books?id=eqegRf2UstIC&q=termite |publisher=Greenwood Press }}</ref><ref>{{cite journal |last1=Bordereau |first1=C. |last2=Robert |first2=A. |last3=Van Tuyen |first3=V. |last4=Peppuy |first4=A. |title=Suicidal defensive behaviour by frontal gland dehiscence in ''Globitermes sulphureus'' Haviland soldiers (Isoptera) |journal=Insectes Sociaux |date=1997 |volume=44 |issue=3 |pages=289–297 |doi=10.1007/s000400050049 |s2cid=19770804}}</ref> Another termite, ''[[Neocapriterme taracua]]'', also engages in suicidal defence. Workers physically unable to use their mandibles while in a fight form a pouch full of chemicals, then deliberately rupture themselves, releasing toxic chemicals that paralyse and kill their enemies.<ref>{{cite journal |last1=Sobotnik |first1=J. |last2=Bourguignon |first2=T. |last3=Hanus |first3=R. |last4=Demianova |first4=Z. |last5=Pytelkova |first5=J. |last6=Mares |first6=M. |last7=Foltynova |first7=P. |last8=Preisler |first8=J. |last9=Cvacka |first9=J. |last10=Krasulova |first10=J. |last11=Roisin |first11=Y. |title=Explosive backpacks in old termite workers |journal=Science |date=2012 |volume=337 |issue=6093 |pages=436 |doi=10.1126/science.1219129 |pmid=22837520 |bibcode=2012Sci...337..436S |s2cid=206540025 }}</ref> The soldiers of the [[Neotropical realm |neotropical]] termite family [[Serritermitidae]] have a defence strategy which involves front gland autothysis, with the body rupturing between the head and abdomen. When soldiers guarding nest entrances are attacked by intruders, they engage in autothysis, creating a block that denies entry to any attacker.<ref>{{Cite journal | journal = Biological Journal of the Linnean Society | title = Structure and function of defensive glands in soldiers of ''Glossotermes oculatus'' (Isoptera: Serritermitidae) | first5 = Y. | volume = 99 | pages = 839–848 | doi = 10.1111/j.1095-8312.2010.01392.x | year = 2010 | last1 = ŠobotnÍk | last5 = Roisin | last2 = Bourguignon | first1 = J. | first2 = T. | last3 = Hanus | last4 = Weyda | first3 = R. | first4 = F. | issue = 4 | doi-access = free}}</ref>

Workers use several different strategies to deal with their dead, including burying, cannibalism, and avoiding a corpse altogether.<ref>{{cite journal |last1=Ulyshen |first1=M.D. |last2=Shelton |first2=T.G. |title=Evidence of cue synergism in termite corpse response behavior |journal=Naturwissenschaften |date=2011 |volume=99 |issue=2 |pages=89–93 |doi=10.1007/s00114-011-0871-3 |pmid=22167071 |bibcode=2012NW.....99...89U |s2cid=2616753}}</ref><ref>{{cite journal |last1=Su |first1=N.Y. |title=Response of the Formosan subterranean termites (Isoptera: Rhinotermitidae) to baits or nonrepellent termiticides in extended foraging arenas. |journal=Journal of Economic Entomology |date=2005 |volume=98 |issue=6 |pages=2143–2152 |doi=10.1603/0022-0493-98.6.2143 |pmid=16539144 |s2cid=196618597}}</ref><ref>{{cite journal |last1=Sun |first1=Q. |last2=Haynes |first2=K.F. |last3=Zhou |first3=X. |title=Differential undertaking response of a lower termite to congeneric and conspecific corpses |journal=Scientific Reports |date=2013 |volume=3 |pages=1650 |doi=10.1038/srep01650 |pmid=23598990 |pmc=3629736 |bibcode=2013NatSR...3.1650S}}</ref> To avoid [[pathogens]], termites occasionally engage in [[necrophoresis]], in which a nestmate carries away a corpse from the colony to dispose of it elsewhere.<ref name=termnecro>{{cite journal |last1=Neoh |first1=K.-B. |last2=Yeap |first2=B.-K. |last3=Tsunoda |first3=K. |last4=Yoshimura |first4=T. |last5=Lee |first5=C.Y. |last6=Korb |first6=J. |title=Do termites avoid carcasses? behavioral responses depend on the nature of the carcasses |journal=PLOS ONE |date=2012 |volume=7 |issue=4 |pages=e36375 |doi=10.1371/journal.pone.0036375 |pmid=22558452 |pmc=3338677 |bibcode = 2012PLoSO...736375N |doi-access=free}}</ref> Which strategy is used depends on the nature of the corpse a worker is dealing with (i.e. the age of the carcass).<ref name=termnecro/>

===Relationship with other organisms===
[[File:Rhizanthella gardneri — Fred Hort.jpg |thumbnail |180px |alt=The Western Underground Orchid lives completely underground. It is unable to photosynthesize, and it is dependent on underground insects such as termites for pollination. The flower head shown is only about 1.5 centimetres across. Dozens of tiny rose-coloured florets are arranged in a tight cluster, surrounded by petals that give the flower the appearance of a pale miniature tulip. |''[[Rhizanthella gardneri]]'' is the only orchid known to be pollinated by termites.]]
A species of [[fungus]] is known to mimic termite eggs, successfully avoiding its natural predators. These small brown balls, known as "termite balls", rarely kill the eggs, and in some cases the workers tend to them.<ref>{{cite journal |last1=Matsuura |first1=K. |title=Termite-egg mimicry by a sclerotium-forming fungus |journal=Proceedings of the Royal Society B: Biological Sciences |date=2006 |volume=273 |issue=1591 |pages=1203–1209 |doi=10.1098/rspb.2005.3434 |pmid=16720392 |pmc=1560272}}</ref> This fungus mimics these eggs by producing cellulose-digesting enzymes known as [[glucosidases]].<ref>{{cite journal |last1=Matsuura |first1=K. |last2=Yashiro |first2=T. |last3=Shimizu |first3=K. |last4=Tatsumi |first4=S. |last5=Tamura |first5=T. |title=Cuckoo fungus mimics termite eggs by producing the cellulose-digesting enzyme β-glucosidase |journal=Current Biology |date=2009 |volume=19 |issue=1 |pages=30–36 |doi=10.1016/j.cub.2008.11.030 |pmid=19110429 |s2cid=18604426 |doi-access=free |bibcode=2009CBio...19...30M }}</ref> A unique mimicking behaviour exists between various species of ''[[Trichopsenius]]'' beetles and certain termite species within ''[[Reticulitermes]]''. The beetles share the same [[cuticular |cuticle]] [[hydrocarbon]]s as the termites and even biosynthesize them. This chemical mimicry allows the beetles to integrate themselves within the termite colonies.<ref>{{cite journal |last1=Howard |first1=R.W. |last2=McDaniel |first2=C.A. |last3=Blomquist |first3=G.J. |title=Chemical mimicry as an integrating mechanism: cuticular hydrocarbons of a termitophile and its host |journal=Science |date=1980 |volume=210 |issue=4468 |pages=431–433 |doi=10.1126/science.210.4468.431 |pmid=17837424 |bibcode=1980Sci...210..431H |s2cid=33221252}}</ref> The developed [[appendages]] on the physogastric abdomen of ''[[Austrospirachtha mimetes]]'' allows the beetle to mimic a termite worker.<ref>{{cite journal |last1=Watson |first1=J.A.L. |title=''Austrospirachtha mimetes'' a new termitophilous corotocine from Northern Australia (Coleoptera: Staphylinidae) |journal=Australian Journal of Entomology |date=1973 |volume=12 |issue=4 |pages=307–310 |doi=10.1111/j.1440-6055.1973.tb01678.x |doi-access=free}}</ref>

Some species of ant are known to capture termites to use as a fresh food source later on, rather than killing them. For example, ''[[Formica nigra]]'' captures termites, and those that try to escape are immediately seized and driven underground.<ref>{{cite journal |last1=Forbes |first1=H.O. |title=Termites Kept in Captivity by Ants |journal=Nature |date=1878 |volume=19 |issue=471 |pages=4–5 |doi=10.1038/019004b0 |bibcode = 1878Natur..19....4F |s2cid=4125839 |url=https://zenodo.org/record/1429239}} {{subscription required}}</ref> Certain species of ants in the subfamily [[Ponerinae]] conduct these raids although other ant species go in alone to steal the eggs or nymphs.<ref name=arb/> Ants such as ''Megaponera analis'' attack the outside of mounds and [[Dorylinae]] ants attack underground.<ref name=arb/><ref>{{cite journal |last1=Darlington |first1=J. |title=Attacks by doryline ants and termite nest defences (Hymenoptera; Formicidae; Isoptera; Termitidae) |journal=Sociobiology |date=1985 |volume=11 |pages=189–200}}</ref> Despite this, some termites and ants can coexist peacefully. Some species of termite, including ''[[Nasutitermes corniger]]'', form associations with certain ant species to keep away predatory ant species.<ref>{{cite journal |journal= Journal of Insect Behavior |title= Behavioural Interactions Between ''Crematogaster brevispinosa rochai'' Forel (Hymenoptera: Formicidae) and Two Nasutitermes Species (Isoptera: Termitidae) |volume=18 |issue= 1 |pages=1–17 |year= 2005 |author=Quinet Y, Tekule N & de Biseau JC |doi=10.1007/s10905-005-9343-y |bibcode= 2005JIBeh..18....1Q |s2cid= 33487814}}</ref> The earliest known association between ''[[Azteca (ant) |Azteca]]'' ants and ''Nasutitermes'' termites date back to the Oligocene to Miocene period.<ref>{{cite journal |last1=Coty |first1=D. |last2=Aria |first2=C. |last3=Garrouste |first3=R. |last4=Wils |first4=P. |last5=Legendre |first5=F. |last6=Nel |first6=A. |last7=Korb |first7=J. |title=The First Ant-Termite Syninclusion in Amber with CT-Scan Analysis of Taphonomy |journal=PLOS ONE |date=2014 |volume=9 |issue=8 |pages=e104410 |doi=10.1371/journal.pone.0104410 |pmid=25140873 |pmc=4139309 |bibcode=2014PLoSO...9j4410C |doi-access=free}}</ref>

[[File:Megaponera analis raid collecting termites.jpg |thumbnail |left |An ant raiding party collecting ''Pseudocanthotermes militaris'' termites after a successful raid]]
54 species of ants are known to inhabit ''Nasutitermes'' mounds, both occupied and abandoned ones.<ref name=santos2010>{{cite journal |last1=Santos |first1=P.P. |last2=Vasconcellos |first2=A. |last3=Jahyny |first3=B. |last4=Delabie |first4=J.H.C. |title=Ant fauna (Hymenoptera, Formicidae) associated to arboreal nests of Nasutitermes spp: (Isoptera, Termitidae) in a cacao plantation in southeastern Bahia, Brazil |journal=Revista Brasileira de Entomologia |date=2010 |volume=54 |issue=3 |pages=450–454 |doi=10.1590/S0085-56262010000300016 |doi-access=free}}</ref> One reason many ants live in ''Nasutitermes'' mounds is due to the termites' frequent occurrence in their geographical range; another is to protect themselves from floods.<ref name=santos2010/><ref>{{cite journal |last1=Jaffe |first1=K. |last2=Ramos |first2=C. |last3=Issa |first3=S. |title=Trophic Interactions Between Ants and Termites that Share Common Nests |journal=Annals of the Entomological Society of America |date=1995 |volume=88 |issue=3 |pages=328–333 |doi=10.1093/aesa/88.3.328}}</ref> ''Iridomyrmex'' also inhabits termite mounds although no evidence for any kind of relationship (other than a predatory one) is known.<ref name=wheeler1936/> In rare cases, certain species of termites live inside active ant colonies.<ref>{{cite journal |last1=Trager |first1=J.C. |title=A Revision of the fire ants, ''Solenopsis geminata'' group (Hymenoptera: Formicidae: Myrmicinae) |journal=Journal of the New York Entomological Society |date=1991 |volume=99 |issue=2 |pages=141–198 |doi=10.5281/zenodo.24912 |jstor=25009890}}</ref> Some invertebrate organisms such as beetles, caterpillars, flies and millipedes are termitophiles and dwell inside termite colonies (they are unable to survive independently).<ref name=Britannica/> As a result, certain beetles and flies have evolved with their hosts. They have developed a gland that secrete a substance that attracts the workers by licking them. Mounds may also provide shelter and warmth to birds, lizards, snakes and scorpions.<ref name=Britannica/>

Termites are known to carry pollen and regularly visit flowers,<ref name=Cingel>{{cite book |last1=Cingel |first1=N.A. van der |title=An atlas of orchid pollination: America, Africa, Asia and Australia |date=2001 |publisher=Balkema |location=Rotterdam |isbn=978-90-5410-486-5 |page=224}}</ref> so are regarded as potential pollinators for a number of flowering plants.<ref>{{cite web |last1=McHatton |first1=R. |title=Orchid Pollination: exploring a fascinating world |url=http://staugorchidsociety.org/PDF/OrchidPollinationbyRonMcHatton.pdf |publisher=The American Orchid Society |access-date=5 September 2015 |page=344 |date=2011}}</ref> One flower in particular, ''[[Rhizanthella gardneri]]'', is regularly pollinated by foraging workers, and it is perhaps the only [[Orchidaceae]] flower in the world to be pollinated by termites.<ref name=Cingel/>

Many plants have developed effective defences against termites. However, seedlings are vulnerable to termite attacks and need additional protection, as their defence mechanisms only develop when they have passed the seedling stage.<ref>{{cite book |last1=Cowie |first1=R. |title=Journey to a Waterfall a biologist in Africa |date=2014 |publisher=Lulu Press |location=Raleigh, North Carolina |isbn=978-1-304-66939-1 |page=169}}</ref> Defence is typically achieved by secreting antifeedant chemicals into the woody cell walls.<ref name=envirostud/> This reduces the ability of termites to efficiently digest the [[cellulose]]. A commercial product, "Blockaid", has been developed in Australia that uses a range of plant extracts to create a paint-on nontoxic [[Termite barriers |termite barrier]] for buildings.<ref name=envirostud>{{cite book |last1=Tan |first1=K.H. |title=Environmental Soil Science |date=2009 |publisher=CRC Press |location=Boca Raton, Florida |isbn=978-1-4398-9501-6 |pages=105–106 |edition=3rd}}</ref> An extract of a species of Australian figwort, ''[[Eremophila (plant) |Eremophila]]'', has been shown to repel termites;<ref name=abc2005>{{cite news |url=http://www.abc.net.au/news/newsitems/200511/s1507502.htm |title=Plant extract stops termites dead |archive-url=https://web.archive.org/web/20090615163604/http://www.abc.net.au/news/newsitems/200511/s1507502.htm |first=Sarah |last=Clark |newspaper=ABC |archive-date=15 June 2009 |date=15 November 2005 |access-date=8 February 2014}}</ref> tests have shown that termites are strongly repelled by the toxic material to the extent that they will starve rather than consume the food. When kept close to the extract, they become disoriented and eventually die.<ref name=abc2005/>

===Relationship with the environment===
Termite populations can be substantially impacted by environmental changes including those caused by human intervention. A Brazilian study investigated the termite assemblages of three sites of [[Caatinga]] under different levels of anthropogenic disturbance in the semi-arid region of northeastern [[Brazil]] were sampled using 65 x 2 m transects.<ref name=termiteassemblages2010>{{cite journal |last1=Vasconcellos |first1=Alexandre |last2=Bandeira |first2=Adelmar G. |last3=Moura |first3=Flávia Maria S. |last4=Araújo |first4=Virgínia Farias P. |last5=Gusmão |first5=Maria Avany B. |last6=Reginaldo |first6=Constantino |title=Termite assemblages in three habitats under different disturbance regimes in the semi-arid Caatinga of NE Brazil |journal=Journal of Arid Environments |publisher=Elsevier |date=February 2010 |volume=74 |issue=2 |pages=298–302 |doi=10.1016/j.jaridenv.2009.07.007 |issn=0140-1963 |bibcode=2010JArEn..74..298V }}</ref> A total of 26 species of termites were present in the three sites, and 196 encounters were recorded in the transects. The termite assemblages were considerably different among sites, with a conspicuous reduction in both diversity and abundance with increased disturbance, related to the reduction of tree density and soil cover, and with the intensity of trampling by cattle and goats. The wood-feeders were the most severely affected feeding group.