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===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>
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