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===Relationship with other organisms=== [[File:Rhizanthella gardneri β 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=1203β1209 |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 Ξ²-glucosidase |journal=Current Biology |date=2009 |volume=19 |issue=1 |pages=30β36 |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=431β433 |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=307β310 |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=4β5 |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=189β200}}</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=1β17 |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=450β454 |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=328β333 |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=141β198 |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=105β106 |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/>
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