Editing Termite (section)

===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>