Editing Termite (section)

===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/>