View source for Termite

{{short description|Social insects related to cockroaches}}
{{About|the social insects}}
{{Distinguish|Thermite|Turmite}}
{{Good article}}
{{Use British English|date=December 2010}}
{{Automatic taxobox
| fossil_range = {{fossilrange|Early Cretaceous | Recent}}
| image = Coptotermes formosanus shiraki USGov k8204-7.jpg
| image_caption = [[Formosan subterranean termite]] (''Coptotermes formosanus'')<br>Soldiers (red-coloured heads)<br>Workers (pale-coloured heads)
| taxon = Isoptera
| authority = [[Gaspard Auguste Brullé|Brullé]], 1832
| display_parents = 3
| subdivision_ranks = Families
| subdivision = {{Plain list|
* † [[Cratomastotermitidae]]<ref name="fossilworks" />
* [[Mastotermitidae]]
* † [[Melqartitermitidae]]
* † [[Mylacrotermitidae]]
* † [[Krishnatermitidae]]
* † [[Termopsidae]]<ref name=Engel2009/>
* † [[Arceotermitidae]]
* [[Stolotermitidae]]
* [[Hodotermopsidae]]
* [[Hodotermitidae]]
* [[Archotermopsidae]]
* [[Kalotermitidae]]
* † [[Tanytermitidae]]
* † [[Archeorhinotermitidae]]
* [[Stylotermitidae]]
* [[Serritermitidae]]
* [[Rhinotermitidae]]
* [[Termitidae]]
}}
}}
'''Termites''' are a group of [[detritivore|detritophagous]] [[Eusociality|eusocial]] [[insect]]s which consume a variety of [[Detritus|decaying plant material]], generally in the form of [[wood]], [[Plant litter|leaf litter]], and [[Humus|soil humus]]. They are distinguished by their moniliform antennae and the soft-bodied and often unpigmented worker caste for which they have been commonly termed "'''white ants'''"; however, they are not [[ant]]s, to which they are only distantly related.<ref name=":1">{{cite web |title=Termite |url=https://www.merriam-webster.com/dictionary/termite?show=0&t=1420442739 |work=Merriam-Webster.com| date=23 May 2023 }}</ref> About 2,972 extant [[species]] are currently described, 2,105 of which are members of the family [[Termitidae]].
Termites comprise the [[infraorder]] '''Isoptera''', or alternatively the [[Taxonomic rank#All ranks|epifamily]] '''Termitoidae''', within the order [[Blattodea]] (along with [[cockroach]]es). Termites were once classified in a separate [[Order (biology)|order]] from cockroaches, but recent [[phylogenetic]] studies indicate that they evolved from cockroaches, as they are deeply nested within the group, and the [[sister group]] to wood eating cockroaches of the genus ''[[Cryptocercus]]''. Previous estimates suggested the divergence took place during the [[Jurassic]] or [[Triassic]]. More recent estimates suggest that they have an origin during the [[Late Jurassic]],<ref name=":0">{{Cite journal|last1=Evangelista|first1=Dominic A.|last2=Wipfler|first2=Benjamin|last3=Béthoux|first3=Olivier|last4=Donath|first4=Alexander|last5=Fujita|first5=Mari|last6=Kohli|first6=Manpreet K.|last7=Legendre|first7=Frédéric|last8=Liu|first8=Shanlin|last9=Machida|first9=Ryuichiro|last10=Misof|first10=Bernhard|last11=Peters|first11=Ralph S.|date=2019-01-30|title=An integrative phylogenomic approach illuminates the evolutionary history of cockroaches and termites (Blattodea)|journal=Proceedings of the Royal Society B: Biological Sciences|language=en|volume=286|issue=1895|pages=20182076|doi=10.1098/rspb.2018.2076|issn=0962-8452|pmc=6364590|pmid=30963947}}</ref> with the first fossil records in the [[Early Cretaceous]].
Similarly to ants and some [[bee]]s and [[wasp]]s from the separate order [[Hymenoptera]], most termites have an analogous "worker" and "soldier" caste system consisting of mostly sterile individuals which are physically and behaviorally distinct. Unlike ants, most colonies begin from sexually mature individuals known as the "king" and "queen" that together form a lifelong [[Monogamy in animals|monogamous]] pair.<ref>{{Cite journal |last1=Nalepa |first1=Christine A. |last2=Jones |first2=Susan C. |title=Evolution of Monogamy in Termites |date=February 1991 |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1469-185X.1991.tb01136.x |journal=Biological Reviews |language=en |volume=66 |issue=1 |pages=83–97 |doi=10.1111/j.1469-185X.1991.tb01136.x |s2cid=84398573 |issn=1464-7931}}</ref> Also unlike ants, which undergo a [[complete metamorphosis]], termites undergo an [[incomplete metamorphosis]] that proceeds through egg, [[nymph (biology)|nymph]], and [[Imago|adult]] stages. Termite colonies are commonly described as [[superorganism]]s due to the collective behaviors of the individuals which form a self-governing entity: the colony itself.{{sfn|Bignell|Roisin|Lo|2010|p=2}} Their colonies range in size from a few hundred individuals to enormous societies with several million individuals. Most species are rarely seen, having a cryptic life-history where they remain hidden within the galleries and tunnels of their nests for most of their lives.
Termites' success as a group has led to them colonizing almost every global landmass, with the highest diversity occurring in the tropics where they are estimated to constitute 10% of the animal [[Biomass (ecology)|biomass]], particularly in [[Africa]] which has the richest diversity with more than 1000 described species.<ref>{{Cite journal |last=van Huis |first=Arnold |date=2017-01-26 |title=Cultural significance of termites in sub-Saharan Africa |journal=Journal of Ethnobiology and Ethnomedicine |volume=13 |issue=1 |pages=8 |doi=10.1186/s13002-017-0137-z |issn=1746-4269 |pmc=5270236 |pmid=28126033 |doi-access=free }}</ref> They are important decomposers of decaying plant matter in the [[Subtropics|subtropical]] and [[Tropics|tropical]] regions of the world, and their recycling of wood and plant matter is of considerable ecological importance. Many species are [[ecosystem engineer]]s capable of altering [[soil]] characteristics such as [[hydrology]], decomposition, [[nutrient cycling]], vegetative growth, and consequently surrounding [[biodiversity]] through the large [[Termite mound|mounds]] constructed by certain species.<ref>{{Cite journal |last1=Jouquet |first1=Pascal |last2=Traoré |first2=Saran |last3=Choosai |first3=Chutinan |last4=Hartmann |first4=Christian |last5=Bignell |first5=David |date=2011-07-01 |title=Influence of termites on ecosystem functioning. Ecosystem services provided by termites |url=https://www.sciencedirect.com/science/article/pii/S1164556311000422 |journal=European Journal of Soil Biology |language=en |volume=47 |issue=4 |pages=215–222 |doi=10.1016/j.ejsobi.2011.05.005 |bibcode=2011EJSB...47..215J |issn=1164-5563}}</ref>
Termites have several impacts on humans. They are a delicacy in the diet of some human cultures such as the [[Makiritare]] in the [[Alto Orinoco Municipality|Alto Orinoco]] province of [[Venezuela]], where they are commonly used as a spice.<ref>{{Cite journal |last1=Paoletti |first1=M. G. |last2=Buscardo |first2=E. |last3=Vanderjagt |first3=D. J. |last4=Pastuszyn |first4=A. |last5=Pizzoferrato |first5=L. |last6=Huang |first6=Y.-S. |last7=Chuang |first7=L.-T. |last8=Glew |first8=R. H. |last9=Millson |first9=M. |last10=Cerda |first10=H. |date=March 2003 |title=Nutrient content of termites (syntermes soldiers) consumed bymakiritare amerindians of the altoorinoco of Venezuela |url=http://dx.doi.org/10.1080/036702403902-2255177 |journal=Ecology of Food and Nutrition |volume=42 |issue=2 |pages=177–191 |doi=10.1080/036702403902-2255177 |bibcode=2003EcoFN..42..177P |s2cid=73373107 |issn=0367-0244}}</ref> They are also used in [[Traditional medicine|traditional medicinal treatments]] of various diseases and ailments, such as influenza, asthma, bronchitis, etc.<ref>{{Cite journal |last=Alves |first=Rômulo RN |date=December 2009 |title=Fauna used in popular medicine in Northeast Brazil |journal=Journal of Ethnobiology and Ethnomedicine |language=en |volume=5 |issue=1 |pages=1 |doi=10.1186/1746-4269-5-1 |pmid=19128461 |pmc=2628872 |issn=1746-4269 |doi-access=free }}</ref><ref>{{Cite journal |last1=Alves |first1=Rômulo R. N. |last2=Dias |first2=Thelma L. P. |date=June 2010 |title=Usos de invertebrados na medicina popular no Brasil e suas implicações para conservação |journal=Tropical Conservation Science |volume=3 |issue=2 |pages=159–174 |doi=10.1177/194008291000300204 |s2cid=86904054 |issn=1940-0829|doi-access=free }}</ref> Termites are most famous for being structural pests; however, the vast majority of termite species are innocuous, with the regional numbers of economically significant species being: [[North America]], 9; [[Australia]], 16; [[Indian subcontinent]], 26; [[Afrotropical realm|tropical Africa]], 24; [[Central America]] and the [[West Indies]], 17. Of known pest species, 28 of the most invasive and structurally damaging belong to the genus ''[[Coptotermes]]''.<ref>{{Cite journal |last=Govorushko |first=Sergey |date=March 2019 |title=Economic and ecological importance of termites: A global review: Termites: a global review  |journal=Entomological Science |language=en |volume=22 |issue=1 |pages=21–35 |doi=10.1111/ens.12328|s2cid=92474272 }}</ref> The distribution of most known pest species is expected to increase over time as a consequence of [[climate change]].<ref>{{Cite journal |last1=Buczkowski |first1=Grzegorz |last2=Bertelsmeier |first2=Cleo |date=February 2017 |title=Invasive termites in a changing climate: A global perspective |journal=Ecology and Evolution |language=en |volume=7 |issue=3 |pages=974–985 |doi=10.1002/ece3.2674 |pmc=5288252 |pmid=28168033|bibcode=2017EcoEv...7..974B }}</ref> Increased urbanization and connectivity is also predicted to expand the range of some pest termites.<ref>{{Cite journal |last1=Duquesne |first1=Edouard |last2=Fournier |first2=Denis |date=2024-04-30 |title=Connectivity and climate change drive the global distribution of highly invasive termites |url=https://neobiota.pensoft.net/article/115411/ |journal=NeoBiota |language=en |volume=92 |pages=281–314 |doi=10.3897/neobiota.92.115411 |doi-access=free |issn=1314-2488}}</ref>
==Etymology==
The infraorder name Isoptera is derived from the [[Greek language|Greek words]] ''iso'' (equal) and ''ptera'' (winged), which refers to the nearly equal size of the fore and hind wings.<ref name="Bugsrule">{{cite book|last1=Cranshaw|first1=W.|title=Bugs Rule!: An Introduction to the World of Insects|date=2013|publisher=Princeton University Press|location=Princeton, New Jersey|isbn=978-0-691-12495-7|page=188|chapter-url=https://books.google.com/books?id=p_3c0qGP_t0C&q=Etymology+of+termite&pg=PA188|chapter=11}}</ref> "Termite" derives from the [[Latin]] and [[Late Latin]] word ''termes'' ("woodworm, white ant"), altered by the influence of Latin ''terere'' ("to rub, wear, erode") from the earlier word ''tarmes''. A termite nest is also known as a ''termitary'' or ''termitarium'' (plural ''termitaria'' or ''termitariums'').<ref>{{cite book|last1=Lobeck|first1=A. Kohl|title=Geomorphology; an Introduction to the Study of Landscapes|date=1939|publisher=McGraw Hill Book Company, Incorporated|location=University of California|pages=431–432|edition=1st|asin=B002P5O9SC}}</ref> The word was first used in English in 1781.<ref>{{cite web|title=Termite|url=http://www.merriam-webster.com/dictionary/termite?show=0&t=1420442739|publisher=Merriam-Webster Online Dictionary|access-date=5 January 2015}}</ref> Earlier attested designations were "wood ants" or "white ants",<ref name="termitety">{{OEtymD|Termite}}</ref> though these may never have been in wide use as termites do not exist in the [[British Isles]].
==Taxonomy and evolution==
[[File:Mastotermes darwiniensis.jpg|left|thumb|upright|alt=The giant northern termite is the most primitive living termite. Its body plan has been described as a cockroach's abdomen stuck to a termite's fore part. Its wings have the same form as roach wings, and like roaches, it lays its eggs in a case.|The external appearance of the giant northern termite ''[[Mastotermes darwiniensis]]'' is suggestive of the close relationship between termites and cockroaches.]]
Termites were formerly placed in the order Isoptera. As early as 1934 suggestions were made that they were closely related to wood-eating cockroaches (genus ''[[Cryptocercus]]'', the woodroach) based on the similarity of their symbiotic gut [[flagellate|flagellates.]]<ref>{{cite journal |last1=Cleveland |first1=L.R. |last2=Hall |first2=S.K. |last3=Sanders |first3=E.P. |last4=Collier |first4=J. |title=The Wood-Feeding Roach ''Cryptocercus'', its protozoa, and the symbiosis between protozoa and roach |journal=Memoirs of the American Academy of Arts and Sciences |date=1934 |volume=17 |issue=2 |pages=185–382 |doi=10.1093/aesa/28.2.216 |doi-access=free}}</ref> In the 1960s additional evidence supporting that hypothesis emerged when F.&nbsp;A. McKittrick noted similar morphological characteristics between some termites and ''Cryptocercus'' [[Nymph (biology)|nymphs]].<ref>{{cite journal |last1=McKittrick |first1=F.A. |date=1965 |title=A contribution to the understanding of cockroach-termite affinities |journal=Annals of the Entomological Society of America |volume=58 |issue=1 |pages=18–22 |doi=10.1093/aesa/58.1.18 |pmid=5834489 |doi-access=free}}</ref> In 2008 [[DNA analysis]] from [[MT-RNR2|16S rRNA]] sequences<ref>{{cite journal|last1=Ware|first1=J.L.|last2=Litman|first2=J.|last3=Klass|first3=K.-D.|last4=Spearman|first4=L.A. |title=Relationships among the major lineages of Dictyoptera: the effect of outgroup selection on dictyopteran tree topology|journal=Systematic Entomology|date=2008|volume=33|issue=3 |pages=429–450 |doi=10.1111/j.1365-3113.2008.00424.x |bibcode=2008SysEn..33..429W |s2cid=86777253}}</ref> supported the position of termites being nested within the evolutionary tree containing the order [[Blattodea]], which included the cockroaches.<ref name="inward">{{cite journal|last1=Inward|first1=D.|last2=Beccaloni|first2=G.|last3=Eggleton|first3=P.|title=Death of an order: a comprehensive molecular phylogenetic study confirms that termites are eusocial cockroaches.|journal=Biology Letters|date=2007|volume=3|issue=3|pages=331–5|doi=10.1098/rsbl.2007.0102|pmid=17412673|pmc=2464702}}</ref><ref>{{cite journal|last1=Eggleton|first1=P.|last2=Beccaloni|first2=G.|last3=Inward|first3=D.|title=Response to Lo ''et al.''|journal=Biology Letters|date=2007|volume=3|issue=5|pages=564–565|doi=10.1098/rsbl.2007.0367|pmc=2391203}}</ref>  The cockroach genus ''Cryptocercus'' shares the strongest phylogenetical similarity with termites and is considered to be a sister-group to termites.<ref>{{cite journal|last1=Ohkuma|first1=M.|last2=Noda|first2=S. |last3=Hongoh|first3=Y.|last4=Nalepa|first4=C.A.|last5=Inoue|first5=T.|date=2009|title=Inheritance and diversification of symbiotic trichonymphid flagellates from a common ancestor of termites and the cockroach ''Cryptocercus''|journal=Proceedings of the Royal Society B: Biological Sciences|volume=276|issue=1655|pages=239–245 |doi=10.1098/rspb.2008.1094|pmc=2674353|pmid=18812290}}</ref><ref>{{cite journal |last1=Lo|first1=N. |last2=Tokuda |first2=G. |last3=Watanabe |first3=H. |last4=Rose |first4=H. |last5=Slaytor |first5=M. |last6=Maekawa |first6=K.|last7=Bandi |first7=C. |last8=Noda |first8=H.|date=June 2000|title=Evidence from multiple gene sequences indicates that termites evolved from wood-feeding cockroaches|journal=Current Biology|volume=10 |issue=13 |pages=801–814 |doi=10.1016/S0960-9822(00)00561-3|pmid=10898984|s2cid=14059547|doi-access=free |bibcode=2000CBio...10..801L }}</ref> Termites and ''Cryptocercus'' share similar morphological and social features: for example, most cockroaches do not exhibit social characteristics, but ''Cryptocercus'' takes care of its young and exhibits other [[social behavior|social behaviour]] such as [[trophallaxis]] and [[Social grooming|allogrooming]].<ref>{{cite book|title=Evolution of the insects|url=https://archive.org/details/evolutioninsects00grim_110|url-access=limited|last1=Grimaldi|first1=D.|last2=Engel|first2=M.S. |date=2005|publisher=Cambridge University Press|isbn=978-0-521-82149-0|edition=1st|location=Cambridge|page=[https://archive.org/details/evolutioninsects00grim_110/page/n251 237]}}</ref> Termites are thought to be the descendants of the genus ''Cryptocercus''.<ref name="inward" /><ref>{{cite journal|last1=Klass|first1=K.D. |last2=Nalepa|first2=C.|last3=Lo|first3=N.|title=Wood-feeding cockroaches as models for termite evolution (Insecta: Dictyoptera): ''Cryptocercus'' vs. ''Parasphaeria boleiriana''|journal=Molecular Phylogenetics & Evolution|date=2008|volume=46|issue=3|pages=809–817|doi=10.1016/j.ympev.2007.11.028|pmid=18226554|bibcode=2008MolPE..46..809K }}</ref> Some researchers have suggested a more conservative measure of retaining the termites as the Termitoidae, an [[epifamily]] within the cockroach order, which preserves the classification of termites at family level and below.<ref>{{cite journal|last1=Lo|first1=N.|last2=Engel|first2=M.S.|last3=Cameron|first3=S.|last4=Nalepa|first4=C.A.|last5=Tokuda|first5=G.|last6=Grimaldi|first6=D.|last7=Kitade|first7=O..|last8=Krishna|first8=K.|last9=Klass|first9=K.-D.|last10=Maekawa|first10=K.|last11=Miura|first11=T.|last12=Thompson|first12=G.J.|title=Comment. Save Isoptera: a comment on Inward ''et al.''|journal=Biology Letters|date=2007|volume=3|issue=5|pages=562–563 |doi=10.1098/rsbl.2007.0264 |pmid=17698448|pmc=2391185}}</ref> Termites have long been accepted to be closely related to cockroaches and [[Mantis|mantids]], and they are classified in the same superorder ([[Dictyoptera]]).<ref name="Costa">{{cite book|title=The other insect societies|last=Costa|first=James|publisher=Harvard University Press|year=2006|isbn=978-0-674-02163-1|pages=135–136}}</ref><ref name="cap2008">{{cite book|title=Encyclopedia of Entomology|url=https://archive.org/details/encyclopediaento00capi|url-access=limited|last1=Capinera|first1=J.L. |date=2008|publisher=Springer|isbn=978-1-4020-6242-1|edition=2nd|location=Dordrecht|pages=[https://archive.org/details/encyclopediaento00capi/page/n3096 3033]–3037, 3754}}</ref>
The oldest unambiguous termite [[fossils]] date to the early [[Cretaceous]], but given the diversity of Cretaceous termites and early fossil records showing mutualism between microorganisms and these insects, they possibly originated earlier in the Jurassic or Triassic.<ref>{{cite journal|last1=Vrsanky|first1=P.|last2=Aristov|first2=D. |title=Termites (Isoptera) from the Jurassic/Cretaceous boundary: Evidence for the longevity of their earliest genera|journal=European Journal of Entomology|date=2014|volume=111|issue=1|pages=137–141 |doi=10.14411/eje.2014.014 |doi-access=free}}</ref><ref>{{cite journal|last1=Poinar|first1=G.O.|title=Description of an early Cretaceous termite (Isoptera: Kalotermitidae) and its associated intestinal protozoa, with comments on their co-evolution|journal=Parasites & Vectors|date=2009|volume=2|issue=1–17 |pages=12|doi=10.1186/1756-3305-2-12 |pmid=19226475|pmc=2669471 |doi-access=free }}</ref><ref>{{cite journal|last1=Legendre|first1=F.|last2=Nel |first2=A.|last3=Svenson|first3=G.J.|last4=Robillard|first4=T.|last5=Pellens|first5=R.|last6=Grandcolas|first6=P.|last7=Escriva|first7=H.|title=Phylogeny of Dictyoptera: Dating the Origin of Cockroaches, Praying Mantises and Termites with Molecular Data and Controlled Fossil Evidence|journal=PLOS ONE|date=2015|volume=10|issue=7|pages=e0130127 |doi=10.1371/journal.pone.0130127 |pmid=26200914 |pmc=4511787|bibcode = 2015PLoSO..1030127L|doi-access=free}}</ref> Possible evidence of a Jurassic origin is the assumption that the extinct [[Mammaliaformes|mammaliaform]] ''[[Fruitafossor]]'' from [[Morrison Formation]] consumed termites, judging from its morphological similarity to modern termite-eating mammals.<ref>{{cite journal|last1=Luo|first1=Z.X.|last2=Wible|first2=J.R.|title=A Late Jurassic digging mammal and early mammalian diversification.|journal=Science|date=2005|volume=308|issue=5718 |pages=103–107 |doi=10.1126/science.1108875 |pmid=15802602|bibcode=2005Sci...308..103L|s2cid=7031381}}</ref> Morrison Formation also yields social insect nest fossils close to that of termites.<ref>{{Cite journal |last1=Smith |first1=Elliott Armour |last2=Loewen |first2=Mark A. |last3=Kirkland |first3=James I. |date=2020-08-29 |title=New social insect nests from the Upper Jurassic Morrison Formation of Utah |url=https://www.giw.utahgeology.org/giw/index.php/GIW/article/view/84 |journal=Geology of the Intermountain West |language=en |volume=7 |pages=281–299 |doi=10.31711/giw.v7.pp281-299 |s2cid=225189490 |issn=2380-7601|doi-access=free }}</ref> The oldest termite nest discovered is believed to be from the [[Upper Cretaceous]] in [[West Texas]], where the oldest known faecal pellets were also discovered.<ref>{{cite journal|last1=Rohr|first1=D.M.|last2=Boucot|first2=A. J.|last3=Miller |first3=J. |last4=Abbott|first4=M.|title=Oldest termite nest from the Upper Cretaceous of west Texas|journal=Geology |volume=14|issue=1|pages=87|doi=10.1130/0091-7613(1986)14<87:OTNFTU>2.0.CO;2|bibcode=1986Geo....14...87R|year=1986}}</ref> Claims that termites emerged earlier have faced controversy. For example, F. M. Weesner indicated that the [[Mastotermitidae]] termites may go back to the [[Late Permian]], 251 million years ago,<ref>{{cite journal|last1=Weesner|first1=F.M.|title=Evolution and Biology of the Termites|journal=Annual Review of Entomology|date=1960|volume=5|issue=1|pages=153–170|doi=10.1146/annurev.en.05.010160.001101}}</ref> and fossil wings that have a close resemblance to the wings of ''Mastotermes'' of the Mastotermitidae, the most primitive living termite, have been discovered in the [[Permian]] layers in Kansas.<ref name="Tilyard">{{cite journal|last1=Tilyard|first1=R.J.|title=Kansas Permian insects. Part XX the cockroaches, or order Blattaria|journal=American Journal of Science|date=1937|volume=34|issue=201|pages=169–202, 249–276 |bibcode=1937AmJS...34..169T|doi=10.2475/ajs.s5-34.201.169}}</ref> It is even possible that the first termites emerged during the [[Carboniferous]].<ref>{{cite book|last1=Henry|first1=M.S.|title=Symbiosis: Associations of Invertebrates, Birds, Ruminants, and Other Biota|date=2013|publisher=Elsevier|location=New York, New York|isbn=978-1-4832-7592-5|page=59}}</ref> The folded wings of the fossil wood roach ''[[Pycnoblattina]]'', arranged in a convex pattern between segments 1a&nbsp;and 2a, resemble those seen in ''Mastotermes'', the only living insect with the same pattern.<ref name="Tilyard" /> [[Kumar Krishna]] ''et al.'', though, consider that all of the Paleozoic and Triassic insects tentatively classified as termites are in fact unrelated to termites and should be excluded from the Isoptera.<ref name="Krishna, K. 2013">{{cite journal|last1=Krishna|first1=K.|last2=Grimaldi|first2=D.A.|last3=Krishna|first3=V.|last4=Engel|first4=M.S.|year=2013|title=Treatise on the Isoptera of the world|url=http://digitallibrary.amnh.org/bitstream/handle/2246/6430/B377%20vol.%201.pdf?sequence=1&isAllowed=y|journal=Bulletin of the American Museum of Natural History|volume=377|issue=7|series=1|pages=1–200|doi=10.1206/377.1|s2cid=87276148}}</ref> Other studies suggest that the origin of termites is more recent, having diverged from ''Cryptocercus'' sometime during the [[Early Cretaceous]].<ref name=":0" /> [[File:Isoptera.jpg|thumb|Macro image of a worker.]]The primitive [[giant northern termite]] (''Mastotermes darwiniensis'') exhibits numerous cockroach-like characteristics that are not shared with other termites, such as laying its eggs in rafts and having anal lobes on the wings.<ref>{{cite book|last1=Bell|first1=W.J.|last2=Roth|first2=L.M.|last3=Nalepa|first3=C.A.|title=Cockroaches: ecology, behavior, and natural history|url=https://archive.org/details/cockroachesecolo00bell|url-access=limited|date=2007|publisher=Johns Hopkins University Press|location=Baltimore, Md.|isbn=978-0-8018-8616-4|page=[https://archive.org/details/cockroachesecolo00bell/page/n177 161]}}</ref> It has been proposed that the Isoptera and Cryptocercidae be grouped in the clade "[[Xylophagodea]]".<ref>{{cite journal|last1=Engel|first1=M.|title=Family-group names for termites (Isoptera), redux|journal=ZooKeys|date=2011|issue=148|pages=171–184|doi=10.3897/zookeys.148.1682|pmid=22287896|pmc=3264418|doi-access=free|bibcode=2011ZooK..148..171E }}</ref> Termites are sometimes called "white ants", but the only resemblance to the ants is due to their sociality which is due to [[convergent evolution]]<ref>{{cite journal|last=Thorne|first=Barbara L|year=1997|title=Evolution of eusociality in termites|journal=Annual Review of Ecology and Systematics|volume=28|issue=5|pages=27–53|url=http://www.thornelab.umd.edu/Termite_PDFS/EvolutionEusocialityTermites.pdf|doi=10.1146/annurev.ecolsys.28.1.27|pmc=349550|url-status=dead|archive-url=https://web.archive.org/web/20100530162505/http://www.thornelab.umd.edu/Termite_PDFS/EvolutionEusocialityTermites.pdf |archive-date=2010-05-30}}</ref><ref name="Harrison2018">{{cite journal|last1=Harrison|first1=Mark C.|last2=Jongepier|first2=Evelien|last3=Robertson|first3=Hugh M.|last4=Arning |first4=Nicolas|last5=Bitard-Feildel|first5=Tristan|last6=Chao|first6=Hsu |last7=Childers|first7=Christopher P.|last8=Dinh|first8=Huyen |last9=Doddapaneni|first9=Harshavardhan|last10=Dugan|first10=Shannon|last11=Gowin|first11=Johannes|last12=Greiner|first12=Carolin|last13=Han|first13=Yi|last14=Hu|first14=Haofu|last15=Hughes|first15=Daniel S. T.|last16=Huylmans|first16=Ann-Kathrin|last17=Kemena|first17=Carsten|last18=Kremer|first18=Lukas P. M.|last19=Lee|first19=Sandra L.|last20=Lopez-Ezquerra|first20=Alberto|last21=Mallet|first21=Ludovic|last22=Monroy-Kuhn|first22=Jose M.|last23=Moser|first23=Annabell|last24=Murali|first24=Shwetha C.|last25=Muzny|first25=Donna M.|last26=Otani|first26=Saria|last27=Piulachs|first27=Maria-Dolors|last28=Poelchau|first28=Monica |last29=Qu |first29=Jiaxin|last30=Schaub|first30=Florentine|last31=Wada-Katsumata|first31=Ayako|last32=Worley |first32=Kim C. |last33=Xie |first33=Qiaolin|last34=Ylla|first34=Guillem |last35=Poulsen|first35=Michael|last36=Gibbs |first36=Richard A. |last37=Schal|first37=Coby|author37-link=Coby Schal|last38=Richards |first38=Stephen |last39=Belles |first39=Xavier|last40=Korb |first40=Judith|last41=Bornberg-Bauer|first41=Erich |display-authors=6 |title=Hemimetabolous genomes reveal molecular basis of termite eusociality|journal=Nature Ecology & Evolution |date=2018|volume=2|issue=3|pages=557–566|doi=10.1038/s41559-017-0459-1|pmid=29403074|pmc=6482461|bibcode=2018NatEE...2..557H }}</ref> with termites being the first social insects to evolve a caste system more than 100 million years ago.<ref name="Nature_2016">{{Cite journal|date=2016|title=Termites had first castes|journal=Nature |volume=530 |issue=7590 |pages=256 |bibcode=2016Natur.530Q.256.|doi=10.1038/530256a|s2cid=49905391|doi-access=free}}</ref> Termite genomes are generally relatively large compared to those of other insects; the first fully sequenced termite genome, of ''[[Zootermopsis nevadensis]]'', which was published in the journal ''[[Nature Communications]]'', consists of roughly 500Mb,<ref>{{cite journal|last1=Terrapon|first1=Nicolas|last2=Li|first2=Cai |last3=Robertson|first3=Hugh M.|last4=Ji|first4=Lu|last5=Meng|first5=Xuehong|last6=Booth |first6=Warren|last7=Chen |first7=Zhensheng |last8=Childers |first8=Christopher P.|last9=Glastad|first9=Karl M.|last10=Gokhale|first10=Kaustubh |display-authors=etal |title=Molecular traces of alternative social organization in a termite genome |journal=Nature Communications|date=2014|volume=5|pages=3636 |doi=10.1038/ncomms4636 |pmid=24845553 |bibcode=2014NatCo...5.3636T |doi-access=free|hdl=2286/R.I.44873|hdl-access=free}}</ref> while two subsequently published genomes, ''[[Macrotermes natalensis]]'' and ''[[Cryptotermes secundus]]'', are considerably larger at around 1.3Gb.<ref>{{cite journal|last1=Poulsen|first1=Michael|last2=Hu|first2=Haofu |last3=Li|first3=Cai |last4=Chen |first4=Zhensheng |last5=Xu|first5=Luohao|last6=Otani|first6=Saria |last7=Nygaard|first7=Sanne |last8=Nobre |first8=Tania|last9=Klaubauf|first9=Sylvia|last10=Schindler|first10=Philipp M .|display-authors=etal |title=Complementary symbiont contributions to plant decomposition in a fungus-farming termite|journal=Proceedings of the National Academy of Sciences|date=2014|volume=111 |issue=40|pages=14500–14505|doi=10.1073/pnas.1319718111 |pmid=25246537|pmc=4209977|bibcode=2014PNAS..11114500P|doi-access=free}}</ref><ref name="Harrison2018" />
External phylogeny showing relationship of termites with other insect groups:<ref>{{cite journal |doi=10.1098/rspb.2018.2076 |title=An integrative phylogenomic approach illuminates the evolutionary history of cockroaches and termites (Blattodea) |year=2019 |last1=Evangelista |first1=Dominic A. |last2=Wipfler |first2=Benjamin |last3=Béthoux |first3=Olivier |last4=Donath |first4=Alexander |last5=Fujita |first5=Mari |last6=Kohli |first6=Manpreet K. |last7=Legendre |first7=Frédéric |last8=Liu |first8=Shanlin |last9=Machida |first9=Ryuichiro |last10=Misof |first10=Bernhard |last11=Peters |first11=Ralph S. |last12=Podsiadlowski |first12=Lars |last13=Rust |first13=Jes |last14=Schuette |first14=Kai |last15=Tollenaar |first15=Ward |last16=Ware |first16=Jessica L. |last17=Wappler |first17=Torsten |last18=Zhou |first18=Xin |last19=Meusemann |first19=Karen |last20=Simon |first20=Sabrina |display-authors=6 |journal=Proceedings of the Royal Society B: Biological Sciences |volume=286 |issue=1895 |pmid=30963947 |pmc=6364590 }}</ref>
{{clade
|label1='''Dictyoptera'''
|1={{clade
|label1=[[Mantodea]]
|1=&nbsp;(Mantises)
|label2=[[Blattodea]]
|2={{clade
|1=[[Blaberoidea]]
|label2=[[Solumblattodea]]
|2={{clade
|1=[[Corydiodea]]
|label2=[[Blattoidea]]
|2={{clade
|1=[[Blattoidae]]
|label2=[[Kittrickea]]
|2={{clade
|1=[[Lamproblattidae]]
|label2=[[Xylophagodea]] |sublabel2=(=[[Tutricablattae]])
|2={{clade
|1=[[Cryptocercidae]] (brown-hooded cockroaches)
|2='''Isoptera (Termites)'''
}}
}}
}}
}}
}}
}}
}}
Internal phylogeny showing relationship of extant termite families:<ref name=":7">{{Cite journal |last1=Hellemans |first1=Simon |last2=Wang |first2=Menglin |last3=Hasegawa |first3=Nonno |last4=Šobotník |first4=Jan |last5=Scheffrahn |first5=Rudolf H. |last6=Bourguignon |first6=Thomas |date=2022-03-02 |title=Using ultraconserved elements to reconstruct the termite tree of life |journal=Molecular Phylogenetics and Evolution |volume=173 |language=en |pages=2021.12.09.472027 |doi=10.1016/j.ympev.2022.107520|biorxiv=10.1101/2021.12.09.472027 |pmid=35577300 |s2cid=245133526 |doi-access=free |bibcode=2022MolPE.17307520H }}</ref><ref>{{Cite journal |last1=Wang |first1=Menglin |last2=Hellemans |first2=Simon |last3=Šobotník |first3=Jan |last4=Arora |first4=Jigyasa |last5=Buček |first5=Aleš |last6=Sillam-Dussès |first6=David |last7=Clitheroe |first7=Crystal |last8=Lu |first8=Tomer |last9=Lo |first9=Nathan |last10=Engel |first10=Michael S. |last11=Roisin |first11=Yves |last12=Evans |first12=Theodore A. |last13=Bourguignon |first13=Thomas |date=2022-04-29 |title=Phylogeny, biogeography and classification of Teletisoptera (Blattaria: Isoptera) |url=http://dx.doi.org/10.1111/syen.12548 |journal=Systematic Entomology |volume=47 |issue=4 |pages=581–590 |doi=10.1111/syen.12548 |bibcode=2022SysEn..47..581W |s2cid=248457693 |issn=0307-6970}}</ref>
{{clade
|label1='''Isoptera'''
|1={{Clade
|1=[[Mastotermitidae]]
|2={{clade
|label1=[[Euisoptera]]
|1={{clade
|label1=[[Teletisoptera]]
|1={SUBCLADE_A}
|label2=[[Icoisoptera]]
|2={SUBCLADE_B}
}}
}}
|targetA={SUBCLADE_A}
|subcladeA={{Clade
|1=[[Stolotermitidae]]
|2={{clade
|1=[[Hodotermopsidae]]
|2={{clade
|1=[[Hodotermitidae]]
|2=[[Archotermopsidae]]
}}
}}
}}
|targetB={SUBCLADE_B}
|subcladeB={{clade
|1=[[Kalotermitidae]]
|2={{clade
|label1=[[Neoisoptera]]
|1={SUBCLADE_C}
}}
}}
|targetC={SUBCLADE_C}
|subcladeC={{Clade
|1=[[Stylotermitidae]]
|2={{clade
|1=[[Serritermitidae]]
|2={{clade
|1=[[Rhinotermitidae]]
|2=[[Termitidae]]
}}
}}
}}
}}
}}
There are currently 3,173 living and fossil termite [[species]] recognised, classified in 12 families; reproductive and/or soldier castes are usually required for identification. The infraorder Isoptera is divided into the following clade and family groups, showing the subfamilies in their respective classification:<ref name="Krishna, K. 2013"/><ref>Constantino, Reginaldo; Termite taxonomist authority, University of Brazil: http://164.41.140.9/catal/statistics.php?filtro=fossil
http://164.41.140.9/catal/statistics.php?filtro=extant
Total: 3,173 extant and extinct sp in Catalogue
http://www.pesquisar.unb.br/professor/reginaldo-constantino</ref>
===Early-diverging termite families===
: '''Infraorder Isoptera''' <small>[[Gaspard Auguste Brullé|Brullé]], 1832</small>
:::::: Family {{extinct}}[[Cratomastotermitidae]] <small>[[Michael S. Engel|Engel]], [[David Grimaldi (entomologist)|Grimaldi]], & [[Kumar Krishna|Krishna]], 2009</small>
:::::: Family [[Mastotermitidae]] <small>[[Jules Desneux|Desneux]], 1904</small>
:: '''Parvorder Euisoptera''' <small>Engel, Grimaldi, & Krishna, 2009</small>
:::::: Family {{extinct}}[[Melqartitermitidae]] <small>Engel, 2021</small>
:::::: Family {{extinct}}[[Mylacrotermitidae]] <small>Engel, 2021</small>
:::::: Family {{extinct}}[[Krishnatermitidae]] <small>Engel, 2021</small>
:::::: Family {{extinct}}[[Termopsidae]] <small>[[Nils Holmgren|Holmgren]], 1911</small>
:::::: Family {{extinct}}[[Carinatermitidae]] <small>Krishna & Grimaldi, 2000</small>
::: '''Minorder Teletisoptera''' <small>Barden & Engel, 2021</small>
:::::: Family [[Archotermopsidae]] <small>Engel, Grimaldi, & Krishna, 2009</small>
:::::: Family [[Hodotermitidae]] <small>Desneux, 1904</small>
:::::: Family [[Hodotermopsidae]] <small>Engel, 2021</small>
::::::: subfamily {{extinct}}[[Hodotermopsellinae]] <small>Engel & Jouault, 2024</small>
::::::: subfamily [[Hodotermopsinae]] <small>Engel, 2021</small>
:::::: Family {{extinct}}[[Arceotermitidae]] <small>Engel, 2021</small>
::::::: subfamily {{extinct}}[[Arceotermitinae]] <small>Engel, 2021</small>
::::::: subfamily {{extinct}}[[Cosmotermitinae]] <small>Engel, 2021</small>
:::::: Family [[Stolotermitidae]] <small>Holmgren, 1910</small>
::::::: subfamily [[Stolotermitinae]] <small>Holmgren, 1910</small>
::::::: subfamily [[Porotermitinae]] <small>[[Alfred E. Emerson|Emerson]], 1942</small>
::: '''Minorder Artisoptera''' <small>Engel, 2021</small>
:::::: Family {{extinct}}[[Tanytermitidae]] <small>Engel, 2021</small>
:::: '''Microrder Icoisoptera''' <small>Engel, 2013</small>
:::::: Family [[Kalotermitidae]] <small>[[Walter Wilson Froggatt|Froggatt]], 1897</small>
::::: ''' Nanorder [[Neoisoptera]]''' <small>Engel, Grimaldi, & Krishna, 2009</small>
:::::: see below for families and subfamilies
===Neoisoptera===
The [[Neoisoptera]], literally meaning "newer termites" (in an evolutionary sense), are a recently coined clade that include families such as the Heterotermitidae, [[Rhinotermitidae]] and [[Termitidae]]. ''Neoisopterans'' have a bifurcated caste development with true workers, and so notably lack pseudergates (except in some basal taxa such as ''Serritermitidae'': see [[#Caste system|below]]). All ''Neoisopterans'' have a fontanelle, which appears as a circular pore or series of pores in a depressed region within the middle of the head. The fontanelle connects to the frontal gland, a novel organ unique to Neoisopteran termites which evolved to excrete an array of defensive chemicals and secretions, and so is typically most developed in the soldier caste.<ref>{{Cite journal |last1=Šobotník |first1=Jan |last2=Bourguignon |first2=Thomas |last3=Hanus |first3=Robert |last4=Sillam-Dussès |first4=David |last5=Pflegerová |first5=Jitka |last6=Weyda |first6=František |last7=Kutalová |first7=Kateřina |last8=Vytisková |first8=Blahoslava |last9=Roisin |first9=Yves |date=2010-12-30 |title=Not Only Soldiers Have Weapons: Evolution of the Frontal Gland in Imagoes of the Termite Families Rhinotermitidae and Serritermitidae |journal=PLOS ONE |volume=5 |issue=12 |pages=e15761 |doi=10.1371/journal.pone.0015761 |pmid=21209882 |pmc=3012694 |bibcode=2010PLoSO...515761S |issn=1932-6203|doi-access=free }}</ref> Cellulose digestion in the family ''Termitidae'' has co-evolved with [[bacteria]]l gut microbiota<ref>{{cite journal | last1 = Kohler | first1 = T | last2 = Dietrich | first2 = C | last3 = Scheffrahn | first3 = RH | last4 = Brune | first4 = A | year = 2012 | title = High-resolution analysis of gut environment and bacterial microbiota reveals functional compartmentation of the gut in wood-feeding higher termites (''Nasutitermes'' spp.) | journal = Applied and Environmental Microbiology | volume = 78 | issue = 13| pages = 4691–4701 | doi=10.1128/aem.00683-12| pmid = 22544239 | pmc = 3370480 | bibcode = 2012ApEnM..78.4691K }}</ref> and many [[Taxon|taxa]] have evolved additional symbiotic relationships such as with the fungus ''[[Termitomyces]]''; in contrast, basal ''Neoisopterans'' and all other ''Euisoptera'' have [[flagellate]]s and [[prokaryote]]s in their hindguts. Extant families and subfamilies are organized as follows:<ref name=":7" /><ref>{{cite journal | last1 = Hellemans | first1 = Simon | last2 = Rocha | first2 = Mauricio M. | last3 = Wang | first3 = Menglin | last4 = Romero Arias | first4 = Johanna | year = 2024 | title = Genomic data provide insights into the classification of extant termites | journal = Nature Communications | volume = 15 | issue = 1 | page = 6724 | doi=10.1038/s41467-024-51028-y| pmid = 39112457 | pmc = 11306793 }}</ref>
: '''Early-Diverging Neoisoptera (Non-Geoisoptera)'''
:: Family {{extinct}}[[Archeorhinotermitidae]] <small>Krishna & Grimaldi, 2003</small>
:: Family [[Stylotermitidae]] <small>Holmgren & Holmgren, 1917</small>
:: Family [[Serritermitidae]] <small>Holmgren, 1910</small>
:: Family [[Rhinotermitidae]] <small>Froggatt, 1897</small>
:: Family [[Termitogetonidae]] <small>Holmgren, 1910</small>
:: Family [[Psammotermitidae]] <small>Holmgren, 1910</small>
::: Subfamily [[Prorhinotermitinae]] <small>Quennedey & Deligne, 1975</small>
::: Subfamily [[Psammotermitinae]] <small>Holmgren, 1910</small>
: '''<big>Clade Geoisoptera</big>''' <small>Engel, Hellemans, & Bourguignon, 2024</small>
:: Family [[Heterotermitinae|Heterotermitidae]] <small>Froggatt, 1897</small> (<small>=[[Coptotermitinae]] Holmgren, 1910</small>)
:: '''Family [[Termitidae]]''' <small>Latreille, 1802</small>
::: Subfamily [[Sphaerotermitinae]] <small>Engel & Krishna, 2004</small>
::: Subfamily [[Macrotermitinae]] <small>Kemner, 1934, nomen protectum [ICZN 2003]</small>
::: Subfamily [[Foraminitermitinae]] <small>Holmgren, 1912</small>
::: Subfamily [[Apicotermitinae]] <small>Grassé & Noirot, 1954 [1955]</small>
::: Subfamily [[Microcerotermitinae]] <small>Holmgren, 1910</small>
::: Subfamily [[Syntermitinae]] <small>Engel & Krishna, 2004</small>
::: Subfamily [[Forficulitermitinae]] <small>Hellemans, Engel, & Bourguignon, 2024</small>
::: Subfamily [[Engelitermitinae]] <small>Romero Arias, Roisin, & Scheffrahn, 2024</small>
::: Subfamily [[Crepititermitinae]] <small>Hellemans, Engel, & Bourguignon, 2024</small>
::: Subfamily [[Protohamitermitinae]] <small>Hellemans, Engel, & Bourguignon, 2024</small>
::: Subfamily [[Cylindrotermitinae]] <small>Hellemans, Engel, & Bourguignon, 2024</small>
::: Subfamily [[Neocapritermitinae]] <small>Hellemans, Engel, & Bourguignon, 2024</small>
::: Subfamily [[Nasutitermitinae]] <small>Hare, 1937</small>
::: Subfamily [[Promirotermitinae]] <small>Hellemans, Engel, & Bourguignon, 2024</small>
::: Subfamily [[Mirocapritermitinae]] <small>Kemner, 1934</small>
::: Subfamily [[Amitermitinae]] <small>Kemner, 1934</small>
::: Subfamily [[Cubitermitinae]] <small>Weidner, 1956</small>
::: Subfamily [[Termitinae]] <small>Latreille, 1802</small>
==Distribution and diversity==
Termites are found on all continents except [[Antarctica]]. The diversity of termite species is low in [[North America]] and [[Europe]] (10 species known in Europe and 50 in North America), but is high in [[South America]], where over 400 species are known.<ref name=tbiology>{{cite web|title=Termite Biology and Ecology|url=http://www.chem.unep.ch/pops/termites/termite_ch2.htm|work=Division of Technology, Industry and Economics Chemicals Branch|publisher=United Nations Environment Programme|access-date=12 January 2015|url-status=dead|archive-url=https://web.archive.org/web/20141110052539/http://www.chem.unep.ch/pops/termites/termite_ch2.htm|archive-date=10 November 2014}}</ref> Of the 2,972 extant termite species currently classified, 1,000 are found in [[Africa]], where mounds are extremely abundant in certain regions. Approximately 1.1&nbsp;million active termite mounds can be found in the northern [[Kruger National Park]] alone.<ref>{{cite journal|last1=Meyer|first1=V.W.|last2=Braack |first2=L.E.O.|last3=Biggs |first3=H.C.|last4=Ebersohn|first4=C.|title=Distribution and density of termite mounds in the northern Kruger National Park, with specific reference to those constructed by ''Macrotermes'' Holmgren (Isoptera: Termitidae)|journal=African Entomology|date=1999|volume=7|issue=1|pages=123–130|url=https://www.researchgate.net/publication/259487863}}</ref> In [[Asia]], there are 435 species of termites, which are mainly distributed in [[China]]. Within China, termite species are restricted to mild [[tropical]] and [[subtropical]] habitats south of the Yangtze River.<ref name=tbiology/> In [[Australia]], all ecological groups of termites (dampwood, drywood, subterranean) are [[Endemism|endemic]] to the country, with over 360 classified species.<ref name=tbiology/> Because termites are highly social and abundant, they represent a disproportionate amount of the world's insect [[biomass (ecology)|biomass]]. Termites and [[ant]]s comprise about 1% of insect species, but represent more than 50% of insect biomass.<ref>{{cite journal|doi=10.1146/annurev-environ-012420-050035|doi-access=free|title=The State of the World's Insects|year=2020|last1=Eggleton|first1=Paul|journal=Annual Review of Environment and Resources|volume=45|pages=61–82}}</ref>
Due to their soft cuticles, termites do not inhabit cool or cold habitats.<ref>{{cite journal|last1=Sanderson|first1=M.G.|title=Biomass of termites and their emissions of methane and carbon dioxide: A global database|journal=Global Biogeochemical Cycles|date=1996|volume=10|issue=4|pages=543–557|doi=10.1029/96GB01893|bibcode=1996GBioC..10..543S}}</ref> There are three ecological groups of termites: dampwood, drywood and subterranean. Dampwood termites are found only in coniferous forests, and drywood termites are found in hardwood forests; subterranean termites live in widely diverse areas.<ref name=tbiology/> One species in the drywood group is the West Indian drywood termite ''([[Cryptotermes brevis]])'', which is an invasive species in Australia.<ref name=Heather1971>{{cite journal|last1=Heather|first1=N.W.|title=The exotic drywood termite ''Cryptotermes brevis'' (Walker) (Isoptera : Kalotermitidae) and endemic Australian drywood termites in Queensland|journal=Australian Journal of Entomology|date=1971|volume=10|issue=2|pages=134–141|doi=10.1111/j.1440-6055.1971.tb00022.x|doi-access=free}}</ref>
{| class="wikitable" style="text-align: center; width:80%;"
|+ Diversity of Isoptera by continent:
|-
|
! scope="col" | [[Asia]]
! scope="col" | [[Africa]]
! scope="col" | [[North America]]
! scope="col" | [[South America]]
! scope="col" | [[Europe]]
! scope="col" | [[Australia]]
|-
! scope="row" | Estimated number of species
| 435
| 1,000
| 50
| 400
| 10
| 360
|}
==Description==
[[File:TermiteAnatomy.png|thumb|General anatomy of a worker termite with Imago (reproductive) and soldier visualized; note the reduction and fusion of [[sclerite]]s on the thorax and more membranous body compared to other [[Dictyoptera]]. Mandible descriptive terminology on the bottom right. The fontanelle is absent in basal termites, being found only in Neoisopteran termites.]]
Termites are usually small, measuring between {{convert|4|and|15|mm|in|frac=16}} in length.<ref name=tbiology/> The largest of all extant termites are the queens of the species ''[[Macrotermes bellicosus]]'', measuring up to over 10 centimetres (4&nbsp;in) in length.<ref>{{cite book|last1=Claybourne|first1=Anna|title=A colony of ants, and other insect groups|date=2013|publisher=Heinemann Library|location=Chicago, Ill.|isbn=978-1-4329-6487-0|page=38}}</ref> Another giant termite, the extinct ''Gyatermes styriensis'', flourished in [[Austria]] during the [[Miocene]] and had a wingspan of {{convert|76|mm|in|frac=4}} and a body length of {{convert|25|mm|in|0}}.<ref name=styria>{{cite journal|last1=Engel|first1=M.S.|last2=Gross|first2=M.|title=A giant termite from the Late Miocene of Styria, Austria (Isoptera)|journal=Naturwissenschaften|date=2008|volume=96|issue=2|pages=289–295|doi=10.1007/s00114-008-0480-y|pmid=19052720|bibcode=2009NW.....96..289E|s2cid=21795900}}</ref>{{refn|It is unknown whether the termite was female or male. If it was a female, the body length would be far greater than 25 millimetres when mature.|group=note}}
Most worker and soldier termites are completely blind as they do not have a pair of eyes. However, some species, such as ''[[Hodotermes|Hodotermes mossambicus]]'', have [[compound eyes]] which they use for orientation and to distinguish sunlight from moonlight.<ref>{{cite journal|last1=Heidecker|first1=J.L.|last2=Leuthold|first2=R.H.|title=The organisation of collective foraging in the harvester termite ''Hodotermes mossambicus'' (Isoptera)|journal=Behavioral Ecology and Sociobiology|date=1984|volume=14|issue=3|pages=195–202|doi=10.1007/BF00299619|s2cid=22158321}}</ref> The [[alate]]s (winged males and females) have eyes along with lateral [[Simple eye in invertebrates|ocelli]]. Lateral ocelli, however, are not found in all termites, absent in the families [[Hodotermitidae]], [[Termopsidae]], and [[Archotermopsidae]].<ref name=Leonardo/>{{sfn|Bignell|Roisin|Lo|2010|p=7}} Like other insects, termites have a small tongue-shaped [[Labrum (arthropod mouthpart)|labrum]] and a [[Clypeus (arthropod anatomy)|clypeus]]; the clypeus is divided into a postclypeus and anteclypeus. Termite antennae have a number of functions such as the sensing of touch, taste, odours (including pheromones), heat and vibration. The three basic segments of a termite antenna include a [[Antenna (biology)#Structure|scape]], a pedicel (typically shorter than the scape), and the flagellum (all segments beyond the scape and pedicel).{{sfn|Bignell|Roisin|Lo|2010|p=7}} The mouth parts contain a [[Maxilla (arthropod mouthpart)|maxillae]], a labium, and a set of [[Mandible (insect mouthpart)|mandibles]]. The maxillae and labium have [[Pedipalp|palps]] that help termites sense food and handling.{{sfn|Bignell|Roisin|Lo|2010|p=7}} The [[Arthropod cuticle|cuticle]] of most castes is soft and flexible due to a resulting lack of sclerotization, particularly of the abdomen which often appears translucent. Pigmentation and sclerotization of the cuticle correlates with [[Life history theory|life history]], with species that spend more time in the surface in the open tending to have a more sclerotized and pigmented exoskeleton.
Consistent with all insects, the anatomy of the termite [[Thorax (insect anatomy)|thorax]] consists of three segments: the [[prothorax]], the [[mesothorax]] and the [[metathorax]].{{sfn|Bignell|Roisin|Lo|2010|p=7}} Each segment contains a pair of [[Arthropod leg|leg]]s. On alates, the wings are located at the mesothorax and metathorax, which is consistent with all four-winged insects. The mesothorax and metathorax have well-developed exoskeletal plates; the prothorax has smaller plates.{{sfn|Bignell|Roisin|Lo|2010|pp=7–9}}
[[File:ZooKeys-148-105-g003 Parastylotermes krishnai.jpg|thumbnail|Diagram showing a wing, along with the [[clypeus (arthropod anatomy)|clypeus]] and leg]]
Termites have a ten-segmented abdomen with two plates, the [[Tergum|tergites]] and the [[Sternum (arthropod anatomy)|sternites]].{{sfn|Bignell|Roisin|Lo|2010|p=11}} The tenth abdominal segment has a pair of short [[Cercus|cerci]].<ref>{{cite book|last1=Robinson|first1=W.H.|title=Urban Insects and Arachnids: A Handbook of Urban Entomology|url=https://archive.org/details/urbaninsectsarac00robi|url-access=limited|date=2005|publisher=Cambridge University Press|location=Cambridge|isbn=978-1-139-44347-0|page=[https://archive.org/details/urbaninsectsarac00robi/page/n299 291]}}</ref> There are ten tergites, of which nine are wide and one is elongated.{{sfn|Bignell|Roisin|Lo|2010|p=12}} The reproductive organs are similar to those in cockroaches but are more simplified. For example, the [[intromittent organ]] is not present in male alates, and the sperm is either immotile or aflagellate. However, Mastotermitidae termites have multiflagellate sperm with limited [[motility]].<ref>{{cite journal|last1=Riparbelli|first1=M.G|last2=Dallai|first2=R|last3=Mercati|first3=D|last4=Bu|first4=Y|last5=Callaini|first5=G|title=Centriole symmetry: a big tale from small organisms|journal=Cell Motility and the Cytoskeleton|date=2009|volume=66|issue=12|pages=1100–5|doi=10.1002/cm.20417|pmid=19746415}}</ref> The genitals in females are also simplified. Unlike in other termites, Mastotermitidae females have an [[ovipositor]], a feature strikingly similar to that in female cockroaches.<ref>{{cite journal|last1=Nalepa|first1=C.A.|last2=Lenz|first2=M.|title=The ootheca of ''Mastotermes darwiniensis'' Froggatt (Isoptera: Mastotermitidae): homology with cockroach oothecae|journal=Proceedings of the Royal Society B: Biological Sciences|date=2000|volume=267|issue=1454|pages=1809–1813|doi=10.1098/rspb.2000.1214|pmid=12233781|pmc=1690738}}</ref>
The non-reproductive castes of termites are wingless and rely exclusively on their six legs for locomotion. The alates fly only for a brief amount of time, so they also rely on their legs.{{sfn|Bignell|Roisin|Lo|2010|p=11}} The appearance of the legs is similar in each caste, but the soldiers have larger and heavier legs. The structure of the legs is consistent with other insects: the parts of a leg include a [[Arthropod leg#Coxa|coxa]], [[Arthropod leg#Trochanter|trochanter]], [[Arthropod leg#Femur|femur]], [[Arthropod leg#Tibia|tibia]] and the [[Arthropod leg#Tarsus|tarsus]].{{sfn|Bignell|Roisin|Lo|2010|p=11}} The number of tibial spurs on an individual's leg varies. Some species of termite have an arolium, located between the [[Chela (organ)|claws]], which is present in species that climb on smooth surfaces but is absent in most termites.<ref>{{cite journal|last1=Crosland|first1=M.W.J.|last2=Su|first2=N.Y.|last3=Scheffrahn|first3=R.H.|title=Arolia in termites (Isoptera): functional significance and evolutionary loss|journal=Insectes Sociaux|date=2005|volume=52|issue=1|pages=63–66|doi=10.1007/s00040-004-0779-4|s2cid=26873138}}</ref>
Unlike in ants, the hind-wings and fore-wings are of equal length.<ref name=Bugsrule/> Most of the time, the alates are poor flyers; their technique is to launch themselves in the air and fly in a random direction.{{sfn|Bignell|Roisin|Lo|2010|p=9}} Studies show that in comparison to larger termites, smaller termites cannot fly long distances. When a termite is in flight, its wings remain at a right angle, and when the termite is at rest, its wings remain parallel to the body.{{sfn|Bignell|Roisin|Lo|2010|p=10}}
===Caste system===
Due to termites being [[Hemimetabolism|hemimetabolous insects]], where the young go through multiple and gradual [[wikt:adultoid|adultoid]] molts before becoming an adult, the advent of eusociality has significantly altered the developmental patterns of this group of insects of which, although similar, is not homologous to that of the eusocial [[Hymenoptera]]. Unlike ants, bees, and wasps which undergo a [[Holometabolism|complete metamorphosis]] and as a result only exhibit developmental plasticity at the immobile [[Larva|larval stage]], the mobile [[Instar|adultoid instars]] of termites remain developmentally flexible throughout all life stages up to the [[Imago|final molt]], which has uniquely allowed for the evolution of distinct yet flexible castes amongst the immatures. As a result the caste system of termites consists mostly of [[Neoteny|neotenous]] or juvenile individuals that undertake the most labor in the colony, which is in contrast to the eusocial Hymenoptera where work is strictly undertaken by the adults.
The developmental plasticity in termites can be described similarly to [[cell potency]], where each molt offers a varying level of phenotypic potency. Early instars typically exhibit the highest phenotypic potency and can be described as ''totipotent'' (''able to molt into all alternative phenotypes''), whereas following instars can be ''pluripotent'' (''able to molt into reproductives and non-reproductives but cannot molt into at least one phenotype''), to ''multipotent'' (''able to molt into either reproductive or non-reproductive phenotypes''), to ''unipotent'' (''able to molt into developmentally close phenotypes''), and then finally ''committed'' (''no longer able to change phenotype, functionally an adult.)''<ref name=":4" /> In most termites, phenotypic potency decreases with every successive molt. Notable exceptions are basal taxa such as the [[Archotermopsidae]], which are able to retain high developmental plasticity even up to the late instars. In these basal taxa, the immatures are able to go through ''progressive'' (''nymph-to-imago''), ''regressive'' (''winged-to-wingless'') and ''stationary'' (''size increase, remains wingless'') molts, which typically indicates the developmental trajectory an individual follows.<ref name=":5" /><ref name=":2" />
There is significant variation of the developmental patterns in termites even across closely related taxa, but can typically be generalized into the following two patterns: The first is the ''linear developmental pathway'', where all immatures are capable of developing into winged adults (''[[Alate]]s''), exhibit high phenotypic potency, and where there exists no true sterile caste other than the soldier. The second is the ''bifurcated developmental pathway'', where immatures diverge into two distinct developmental lineages known as the ''<u>nymphal</u>'' (winged) and ''<u>apterous</u>'' (wingless) lines. The bifurcation occurs early, either at the egg or the first two instars, and represents an irreversible and committed development to either the reproductive or non-reproductive lifestyles. As such, the apterous lineage consists mostly of wingless and truly [[Altruism (biology)|altruistic]] sterile individuals (true workers, soldiers), whereas the nymphal lineage consists mainly of fertile individuals destined to become winged reproductives. The bifurcated developmental pathway is found mainly in the derived taxa (i.e. [[Neoisoptera]]), and is believed to have evolved in tandem with the sterile worker caste as species moved to foraging for food beyond their nests, as opposed to the nest also being the food (such as in obligate wood-dwellers).<ref name=":6" /><ref name=":5" />
There are three main castes which are discussed below:
[[File:Termite vs Ant Developmental biology.png|thumb|Developmental biology of ants versus termites. As opposed to ants which have a linear and irreversible development from larval instars to adult (imago), termites exhibit a more complex and often bifurcated development which allows for more flexible caste pathways. Although in most termites, caste development is restricted to closely related pathways dependent on a variety of factors such as pheromonal cues, sex and size of an individual.]]
''Worker'' termites undertake the most labor within the colony, being responsible for foraging, food storage, and brood and nest maintenance.{{sfn|Bignell|Roisin|Lo|2010|p=13}}<ref name="aus" /> Workers are tasked with the digestion of [[cellulose]] in food and are thus the most likely caste to be found in infested wood. The process of worker termites feeding other nestmates is known as [[trophallaxis]]. Trophallaxis is an effective nutritional tactic to convert and recycle nitrogenous components.<ref name="trophal">{{cite journal|last1=Machida|first1=M.|last2=Kitade|first2=O.|last3=Miura|first3=T.|last4=Matsumoto|first4=T.|title=Nitrogen recycling through proctodeal trophallaxis in the Japanese damp-wood termite ''Hodotermopsis japonica'' (Isoptera, Termopsidae)|journal=Insectes Sociaux|date=2001|volume=48|issue=1|pages=52–56|doi=10.1007/PL00001745|s2cid=21310420|issn=1420-9098}}</ref> It frees the parents from feeding all but the first generation of offspring, allowing for the group to grow much larger and ensuring that the necessary gut symbionts are transferred from one generation to another. Workers are believed to have evolved from older wingless immatures (''Larvae'') that evolved cooperative behaviors; and indeed in some basal [[Taxon|taxa]] the late instar larvae are known to undertake the role of workers without differentiating as a true separate caste.<ref name="aus" /><ref name=":4">{{Cite journal |last1=Revely |first1=Lewis |last2=Sumner |first2=Seirian |last3=Eggleton |first3=Paul |date=2021-02-18 |title=The Plasticity and Developmental Potential of Termites |journal=Frontiers in Ecology and Evolution |volume=9 |doi=10.3389/fevo.2021.552624 |issn=2296-701X|doi-access=free }}</ref> Workers can either be male or female, although in some species with [[Polymorphism (biology)|polymorphic]] workers either sex may be restricted to a certain developmental path. Workers may also be fertile or sterile, however the term "worker" is normally reserved for the latter, having evolved in taxa that exhibit a bifurcated developmental pathway.<ref name=":6">{{Cite journal |last1=Higashi |first1=Masahiko |last2=Yamamura |first2=Norio |last3=Abe |first3=Takuya |last4=Burns |first4=Thomas P. |date=1991-10-22 |title=Why don't all termite species have a sterile worker caste? |journal=Proceedings of the Royal Society B: Biological Sciences |language=en |volume=246 |issue=1315 |pages=25–29 |doi=10.1098/rspb.1991.0120 |issn=0962-8452 |pmid=1684665 |s2cid=23067349}}</ref> As a result, sterile workers like in the family [[Termitidae]] are termed ''true workers'' and are the most derived, while those that are undifferentiated and fertile as in the wood-nesting [[Archotermopsidae]] are termed ''pseudergates'', which are the most basal.<ref name=":2" /> ''True workers'' are individuals which irreversibly develop from the ''apterous lineage and'' have completely forgo development into a winged adult. They display altruistic behaviors and either have terminal molts or exhibit a low level of phenotypical potency. True workers across different termite taxa (''Mastotermitidae'', ''Hodotermitidae'', ''Rhinotermitidae'' & ''Termitidae)'' can widely vary in the level of developmental plasticity even between closely related taxa, with many species having true workers that can molt into the other apterous castes such as ''[[ergatoid]]s'' (worker reproductive; apterous neotenics), soldiers, or the other worker castes. ''Pseudergates sensu stricto'' are individuals which arise from the linear developmental pathway that have regressively molted and lost their wing buds, and are regarded as totipotent immatures. They are capable of performing work but are overall less involved in labor and considered more cooperative than truly altruistic. ''Pseudergates sensu lato'', otherwise known as ''false workers'', are most represented in basal lineages (''Kalotermitidae'', ''Archotermopsidae'', ''Hodotermopsidae'', ''Serritermitidae'') and closely resemble true workers in which they also perform most of the work and are similarly altruistic, however differ in developing from the linear developmental pathway where they exist in a stationary molt; i.e they have halted development before the growth of wing buds, and are regarded as pluripotent immatures.<ref name=":2">{{Cite book |last=Jürgen |first=Korb, Judith Thomas-Poulsen, Michael Hu, Haofu Li, Cai Boomsma, Jacobus Jan Zhang, Guojie Liebig |url=http://worldcat.org/oclc/937913325 |title=A genomic comparison of two termites with different social complexity |date=2015 |oclc=937913325}}</ref><ref name=":5">{{Cite book |url=https://link.springer.com/book/10.1007/978-90-481-3977-4 |title=Biology of Termites: a Modern Synthesis |year=2011 |language=en |doi=10.1007/978-90-481-3977-4|isbn=978-90-481-3976-7 |editor-last1=Bignell |editor-last2=Roisin |editor-last3=Lo |editor-first1=David Edward |editor-first2=Yves |editor-first3=Nathan }}</ref>
The ''soldier'' caste is the most anatomically and behaviorally specialized, and their sole purpose is to defend the colony.{{sfn|Bignell|Roisin|Lo|2010|p=18}} Many soldiers have large heads with highly modified powerful jaws so enlarged that they cannot feed themselves. Instead, like juveniles, they are fed by workers.{{sfn|Bignell|Roisin|Lo|2010|p=18}}<ref name="Britannica" /> [[Fontanelle]]s, simple holes in the forehead that lead to a gland which exudes defensive secretions, are a feature of the clade [[Neoisoptera]] and are present in all extant taxa such as Rhinotermitidae.<ref>{{cite book|last1=Busvine|first1=J.R.|title=Insects and Hygiene The biology and control of insect pests of medical and domestic importance|date=2013|publisher=Springer US|location=Boston, MA|isbn=978-1-4899-3198-6|page=545|edition=3rd}}</ref> The majority of termite species have mandibulate soldiers which are easily identified by the disproportionately large sclerotized head and mandibles.<ref name="aus" />{{sfn|Bignell|Roisin|Lo|2010|p=18}} Among certain termites, the soldier caste has evolved globular (phragmotic) heads to block their narrow tunnels such as seen in [[Cryptotermes]].<ref>{{cite book|last1=Meek|first1=S.P.|title=Termite Control at an Ordnance Storage Depot|date=1934|publisher=American Defense Preparedness Association|page=159}}</ref> Amongst mandibulate soldiers, the mandibles have been adapted for a variety of defensive strategies: Biting/crushing (''[[Incisitermes]]''), slashing (''[[Cubitermes]]''), slashing/snapping (''[[Dentispicotermes]]''), symmetrical snapping (''[[Termes (insect)|Termes]]''), asymmetrical snapping (''[[Neocapritermes]]''), and piercing (''[[Armitermes]]'').<ref>{{Cite web|url=https://www.researchgate.net/publication/237089558|title=Worker mandible shape and feeding groups in termites}}</ref> In the more derived termite taxa, the soldier caste can be polymorphic and include minor and major forms. Other morphologically specialized soldiers includes the [[Nasutitermitinae|Nasutes]], which have a horn-like nozzle projection ([[Fontanellar gun|nasus]]) on the head.<ref name="aus" /> These unique soldiers are able to spray noxious, sticky secretions containing [[diterpene]]s at their enemies.<ref name="biosyn">{{cite journal|last1=Prestwich|first1=G.D.|title=From tetracycles to macrocycles|journal=Tetrahedron|date=1982|volume=38|issue=13|pages=1911–1919|doi=10.1016/0040-4020(82)80040-9}}</ref> [[Nitrogen fixation]] plays an important role in Nasute nutrition.<ref>{{cite journal|last1=Prestwich|first1=G. D.|last2=Bentley|first2=B.L.|last3=Carpenter|first3=E.J.|title=Nitrogen sources for neotropical nasute termites: Fixation and selective foraging|journal=Oecologia|date=1980|volume=46|issue=3|pages=397–401|doi=10.1007/BF00346270|pmid=28310050|issn=1432-1939|bibcode=1980Oecol..46..397P|s2cid=6134800}}</ref> Soldiers are normally a committed sterile caste and so do not molt into anything else, but in certain basal taxa like the Archotermopsidae they are known to rarely molt into neotenic forms that develop functional sexual organs.<ref name=":3">{{Cite journal|last1=Thorne|first1=B. L.|last2=Breisch|first2=N. L.|last3=Muscedere|first3=M. L.|date=2003-10-28|title=Evolution of eusociality and the soldier caste in termites: Influence of intraspecific competition and accelerated inheritance|journal=Proceedings of the National Academy of Sciences|language=en|volume=100|issue=22|pages=12808–12813|doi=10.1073/pnas.2133530100|issn=0027-8424|pmc=240700|pmid=14555764|bibcode=2003PNAS..10012808T|doi-access=free}}</ref> In species with the linear developmental pathway, soldiers develop from apterous immatures and constitute the only true sterile caste in these taxa.<ref name=":3" />
The primary reproductive caste of a colony consists of the fertile adult (''imago'') female and male individuals, colloquially known as the queen and king.<ref name=techreport>{{cite tech report |first1=M.A.|last1=Horwood|first2=R.H.|last2=Eldridge|title=Termites in New South Wales Part 1. Termite biology |number=96-38 |institution=Forest Resources Research |year=2005|url=http://www.dpi.nsw.gov.au/__data/assets/pdf_file/0016/252007/Termites-Part-1--Termite-biology.pdf|issn=0155-7548}}</ref> The queen of the colony is responsible for egg production of the colony. Unlike in ants, the male and female reproductives form lifelong pairs where the king will continue to mate with the queen throughout their lives.<ref name=Keller1998/> In some species, the abdomen of the queen swells up dramatically to increase [[fecundity]], a characteristic known as [[physogastrism]].{{sfn|Bignell|Roisin|Lo|2010|p=13}}<ref name=techreport/> Depending on the species, the queen starts producing reproductive alates at a certain time of the year, and huge swarms emerge from the colony when [[nuptial flight]] begins. These swarms attract a wide variety of predators.<ref name=techreport/> The queens can be particularly long-lived for insects, with some reportedly living as long as 30 or 50 years. In both the linear and bifurcated developmental pathways, the primary reproductives only develop from winged immatures (nymphs). These winged immatures are capable of regressively molting into a form known as ''brachypterous neotenics'' (''nymphoids''), which retain juvenile and adult characteristics. ''BN''<nowiki/>'s can be found in both the derived and basal termite taxa, and generally serve as supplementary reproductives.<ref name=":4" /><ref name=":5" />
==Life cycle==
[[File:Termite-by-RalfR.jpg|thumbnail|alt=A termite nymph looks like a smaller version of an adult but lacks the specialisations that would enable identification of its caste.|A young termite nymph featuring visible wing buds. Nymphs mainly develop into [[alate]]s.]]
[[File:20200403 Termite and shed wings on interior window sill.png|thumb|A termite alate with shed wings from other alates on an interior window sill. Shedding of wings is associated with reproductive swarming.<ref name=NewYorker_20180918>{{cite magazine |last1=Srinivasan |first1=Amia |title=What Termites Can Teach Us |url=https://www.newyorker.com/magazine/2018/09/17/what-termites-can-teach-us |magazine=The New Yorker |date=September 10, 2018 |archive-url=https://web.archive.org/web/20200307022415/https://www.newyorker.com/magazine/2018/09/17/what-termites-can-teach-us |archive-date=March 7, 2020 |url-status=live }}</ref>]]
Termites are often compared with the [[eusociality|social]] Hymenoptera (ants and various species of bees and wasps), but their differing evolutionary origins result in major differences in life cycle. In the eusocial Hymenoptera, the workers are exclusively female. Males (drones) are haploid and develop from unfertilised eggs, while females (both workers and the queen) are diploid and develop from fertilised eggs. In contrast, worker termites, which constitute the majority in a colony, are [[diploid]] individuals of both sexes and develop from fertilised eggs. Depending on species, male and female workers may have different roles in a termite colony.<ref>{{cite journal |last1=Korb |first1=J. |title=Termites, hemimetabolous diploid white ants? |journal=Frontiers in Zoology |date=2008 |volume=5 |issue=1 |pages=15 |doi=10.1186/1742-9994-5-15 |pmc=2564920 |pmid=18822181 |doi-access=free }}</ref>
The life cycle of a termite begins with an [[egg (biology)|egg]], but is different from that of a bee or ant in that it goes through a developmental process called [[incomplete metamorphosis]], going through multiple gradual pre-adult molts that are highly [https://www.sciencedirect.com/topics/medicine-and-dentistry/developmental-plasticity#:~:text=Developmental%20plasticity%20refers%20to%20the,generated%20from%20a%20single%20genotype. developmentally plastic] before becoming an adult.<ref name=":4" /><ref>{{cite web |last1=Davis |first1=P. |title=Termite Identification |url=http://agspsrv34.agric.wa.gov.au/ento/termites.htm |publisher=Entomology at Western Australian Department of Agriculture |url-status=dead |archive-url=http://webarchive.loc.gov/all/20090612011547/http://agspsrv34.agric.wa.gov.au/ento/termites.htm |archive-date=2009-06-12 }}</ref> Unlike in other [[Hemimetabolism|hemimetabolous]] insects, ''nymphs'' are more strictly defined in termites as immature young with visible wing buds, which often invariably go through a series of [[Moulting|moults]] to become winged [[Imago|adults]].<ref>{{cite journal|last1=Neoh |first1=K.B. |last2=Lee|first2=C.Y.|title=Developmental stages and caste composition of a mature and incipient colony of the drywood termite, ''Cryptotermes dudleyi'' (Isoptera: Kalotermitidae) |journal=Journal of Economic Entomology |date=2011 |volume=104 |issue=2 |pages=622–8 |pmid=21510214 |doi=10.1603/ec10346|s2cid=23356632 |doi-access=free }}</ref><ref name=":4" /> ''Larvae'', which are defined as early nymph instars with absent wing buds, exhibit the highest developmental potentiality and are able to molt into ''Alates'', ''Soldiers'', ''Neotenics'', or ''Workers''. Workers are believed to have evolved from larvae, sharing many similarities to the extent that workers can be regarded as "larval", in that both lack wings, eyes, and functional reproductive organs while maintaining varying levels of developmental flexibility, although usually to a much lesser extent in workers. The main distinction being that while larvae are wholly dependent on other nestmates to survive, workers are independent and are able to feed themselves and contribute to the colony. Workers remain wingless and across many [[Taxon|taxa]] become developmentally arrested, appearing to not change into any other caste until death.<ref name=":4" /> In some basal taxa, there is no distinction, with the "workers" (pseudergates) essentially being late instar larvae that retain the ability to change into all other castes.<ref name=":5" />
The development of larvae into adults can take months; the time period depends on food availability and nutrition, temperature, and the size of the colony. Since larvae and nymphs are unable to feed themselves, workers must feed them, but workers also take part in the social life of the colony and have certain other tasks to accomplish such as foraging, building or maintaining the nest or tending to the queen.<ref name=aus>{{cite web|title=Termites|url=https://australian.museum/learn/animals/insects/termites/|publisher=Australian Museum|access-date=8 January 2015}}</ref><ref name=Michael>{{cite web|last1=Schneider|first1=M.F.|title=Termite Life Cycle and Caste System|url=http://www.fzi.uni-freiburg.de/InsectPestKey-long%20version/termit2.htm|publisher=University of Freiburg|access-date=8 January 2015|date=1999}}</ref> Pheromones regulate the caste system in termite colonies, preventing all but a very few of the termites from becoming fertile queens.<ref>{{cite journal|last1=Simpson|first1=S.J.|last2=Sword|first2=G.A.|last3=Lo|first3=N.|title=Polyphenism in Insects|journal=Current Biology|date=2011|volume=21|issue=18|pages=738–749|doi=10.1016/j.cub.2011.06.006|pmid=21959164|s2cid=656039|doi-access=free|bibcode=2011CBio...21.R738S }}</ref>
Queens of the [[eusociality|eusocial]] termite ''[[Reticulitermes]] speratus'' are capable of a long lifespan without sacrificing [[fecundity]]. These long-lived queens have a significantly lower level of oxidative damage, including [[DNA oxidation|oxidative DNA damage]], than workers, soldiers and nymphs.<ref name="pmid28076409">{{cite journal |vauthors=Tasaki E, Kobayashi K, Matsuura K, Iuchi Y |title=An Efficient Antioxidant System in a Long-Lived Termite Queen |journal=PLOS ONE |volume=12 |issue=1 |pages=e0167412 |date=2017 |pmid=28076409 |pmc=5226355 |doi=10.1371/journal.pone.0167412 |bibcode=2017PLoSO..1267412T |doi-access=free }}</ref> The lower levels of damage appear to be due to increased [[catalase]], an enzyme that protects against [[oxidative stress]].<ref name="pmid28076409" />
===Reproduction===
[[File:Flying Termites after rain.jpg|thumbnail|alt=Hundreds of winged termite reproductives swarming after a summer rain, filling the field of the photograph.|Alates swarming during nuptial flight after rain|left]]
Termite alates (winged virgin queens and kings) only leave the colony when a [[nuptial flight]] takes place. Alate males and females pair up together and then land in search of a suitable place for a colony.<ref name=v2010>{{cite web|last1=Miller|first1=D.M.|title=Subterranean Termite Biology and Behavior|publisher=Virginia Tech (Virginia State University)|access-date=8 January 2015|url=https://pubs.ext.vt.edu/444/444-502/444-502.html|date=5 March 2010}}</ref> A termite king and queen do not mate until they find such a spot. When they do, they excavate a chamber big enough for both, close up the entrance and proceed to mate.<ref name=v2010/> After mating, the pair may never surface again, spending the rest of their lives in the nest. Nuptial flight time varies in each species. For example, alates in certain species emerge during the day in summer while others emerge during the winter.<ref name=Gouge>{{cite web|last1=Gouge|first1=D.H.|last2=Smith|first2=K.A.|last3=Olson|first3=C.|last4=Baker|first4=P.|title=Drywood Termites|url=http://ag.arizona.edu/pubs/insects/az1232/|publisher=University of Arizona|work=Cooperative Extension, College of Agriculture & Life Sciences|access-date=16 September 2015|date=2001|archive-date=10 November 2016|archive-url=https://web.archive.org/web/20161110102210/http://ag.arizona.edu/pubs/insects/az1232/|url-status=dead}}</ref> The nuptial flight may also begin at dusk, when the alates swarm around areas with many lights. The time when nuptial flight begins depends on the environmental conditions, the time of day, moisture, wind speed and precipitation.<ref name=Gouge/> The number of termites in a colony also varies, with the larger species typically having 100–1,000 individuals. However, some termite colonies, including those with many individuals, can number in the millions.<ref name=styria/>
The queen only lays 10–20 eggs in the very early stages of the colony, but lays as many as 1,000 a day when the colony is several years old.<ref name=aus/> At maturity, a primary queen has a great capacity to lay eggs. In some species, the mature queen has a greatly distended abdomen and may produce 40,000 eggs a day.<ref>{{cite journal |last1=Kaib |first1=M. |last2=Hacker |first2=M. |last3=Brandl |first3=R. |title=Egg-laying in monogynous and polygynous colonies of the termite ''Macrotermes michaelseni'' (Isoptera, Macrotermitidae)|journal=Insectes Sociaux |date=2001 |volume=48 |issue=3 |pages=231–237 |doi=10.1007/PL00001771|s2cid=35656795 }}</ref> The two mature ovaries may have some 2,000 [[ovariole]]s each.<ref>{{cite book|last1=Gilbert|first1=executive editors, G.A. Kerkut, L.I.|title=Comprehensive insect physiology, biochemistry, and pharmacology |date=1985 |publisher=Pergamon Press |location=Oxford |isbn=978-0-08-026850-7 |page=167 |edition=1st |url=https://books.google.com/books?id=bkTgBAAAQBAJ&q=termite+queen+ovarioles&pg=PA167}}</ref> The abdomen increases the queen's body length to several times more than before mating and reduces her ability to move freely; attendant workers provide assistance.
[[File:Egg grooming behaviour of Reticulitermes speratus workers in a nursery cell - pone.0000813.s006.ogv|thumb|alt=In this movie, workers pick eggs from the egg pile, groom them and then return them into the egg pile. They coat the eggs with their saliva to protect them against drying out and to protect them against infection: the saliva contains lyzozyme which has antibacterial properties. The lysozyme also acts as an egg recognition pheromone.|Egg grooming behaviour of ''[[Reticulitermes|Reticulitermes speratus]]'' workers in a nursery cell]]The king grows only slightly larger after initial mating and continues to mate with the queen for life (a termite queen can live between 30&nbsp;and 50&nbsp;years); this is very different from ant colonies, in which a queen mates once with the males and stores the gametes for life, as the male ants die shortly after mating.<ref name=Keller1998>{{cite journal |last1=Keller|first1=L.|year=1998 |title=Queen lifespan and colony characteristics in ants and termites |journal=Insectes Sociaux|volume=45|pages=235–246|doi=10.1007/s000400050084|issue=3|s2cid=24541087}}</ref><ref name=Michael/> If a queen is absent, a termite king produces pheromones which encourage the development of replacement termite queens.<ref>{{cite book|last1=Wyatt|first1=T.D.|title=Pheromones and animal behaviour: communication by smell and taste|url=https://archive.org/details/pheromonesanimal0000wyat|url-access=registration|date=2003|publisher=Cambridge University Press|location=Cambridge|isbn=978-0-521-48526-5|page=[https://archive.org/details/pheromonesanimal0000wyat/page/119 119]|edition=Repr. with corrections 2004.}}</ref> As the queen and king are monogamous, sperm competition does not occur.<ref>{{cite journal|last1=Morrow|first1=E.H.|title=How the sperm lost its tail: the evolution of aflagellate sperm.|journal=Biological Reviews of the Cambridge Philosophical Society|date=2004|volume=79|issue=4|pages=795–814|doi=10.1017/S1464793104006451|pmid=15682871|s2cid=25878093}}</ref>
Termites going through incomplete metamorphosis on the path to becoming alates form a subcaste in certain species of termite, functioning as potential supplementary reproductives. These supplementary reproductives only mature into primary reproductives upon the death of a king or queen, or when the primary reproductives are separated from the colony.<ref name="uflorida">{{cite web |title=Native subterranean termites |url=http://entnemdept.ufl.edu/creatures/urban/termites/native_subterraneans.htm |access-date=8 January 2015 |publisher=University of Florida}}</ref><ref>{{cite web|title=Supplementary reproductive|url=http://www2.hawaii.edu/~entomol/glossary/definition_supplementary_reproductive.htm|archive-url=https://web.archive.org/web/20141030112432/http://www2.hawaii.edu/~entomol/glossary/definition_supplementary_reproductive.htm |archive-date=30 October 2014|publisher=University of Hawaii|access-date=16 September 2015}}</ref> Supplementaries have the ability to replace a dead primary reproductive, and there may also be more than a single supplementary within a colony.<ref name=aus/> Some queens have the ability to switch from sexual reproduction to [[asexual reproduction]]. Studies show that while termite queens mate with the king to produce colony workers, the queens reproduce their replacements ([[Neoteny|neotenic]] queens) [[Parthenogenesis|parthenogenetically]].<ref>{{cite journal|last1=Yashiro|first1=T.|last2=Matsuura|first2=K.|title=Termite queens close the sperm gates of eggs to switch from sexual to asexual reproduction|journal=Proceedings of the National Academy of Sciences |date=2014 |volume=111 |issue=48 |pages=17212–17217 |doi=10.1073/pnas.1412481111 |pmid=25404335 |pmc=4260566 |bibcode=2014PNAS..11117212Y|doi-access=free}}</ref><ref>{{cite journal |last1=Matsuura |first1=K.|last2=Vargo |first2=E.L.|last3=Kawatsu |first3=K. |last4=Labadie |first4=P. E. |last5=Nakano |first5=H.|last6=Yashiro |first6=T. |last7=Tsuji |first7=K. |title=Queen Succession Through Asexual Reproduction in Termites |journal=Science |date=2009 |volume=323 |issue=5922 |pages=1687 |doi=10.1126/science.1169702 |pmid=19325106 |bibcode=2009Sci...323.1687M|s2cid=21785886}}</ref>
The neotropical termite ''Embiratermes neotenicus'' and several other related species produce colonies that contain a primary king accompanied by a primary queen or by up to 200 [[neoteny|neotenic]] queens that had originated through [[thelytoky|thelytokous parthenogenesis]] of a founding primary queen.<ref name="pmid26019158">{{cite journal |vauthors=Fougeyrollas R, Dolejšová K, Sillam-Dussès D, Roy V, Poteaux C, Hanus R, Roisin Y |title=Asexual queen succession in the higher termite Embiratermes neotenicus |journal=Proc. Biol. Sci. |volume=282 |issue=1809 |pages=20150260 |date=June 2015 |pmid=26019158 |pmc=4590441 |doi=10.1098/rspb.2015.0260 }}</ref> The form of [[parthenogenesis]] likely employed maintains [[zygosity|heterozygosity]] in the passage of the [[genome]] from mother to daughter, thus avoiding [[inbreeding depression]].
==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 |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 |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...3E1650S}}</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.
==Nests==
[[File:Termite workers at work.webm|thumb|Termite workers at work]]
[[File:Termite-nest-Tulum-Mexico.jpg|thumb|upright|alt=Photograph of an arboreal termite nest built on a tree trunk high above ground. It has an ovoid shape and appears to be larger than a basketball. It is dark brown in colour, and it is made of carton, a mixture of digested wood and termite faeces that is strong and resistant to rain. Covered tunnels constructed of carton can be seen leading down the shaded side of the tree from the nest to the ground.|An arboreal termite nest in Mexico]]
[[File:Termitenest33189014241 9f0022ebfe o.jpg|thumb|Termite nest in a [[Banksia]], [[Palm Beach, New South Wales|Palm Beach, Sydney.]]]]
A termite nest can be considered as being composed of two parts, the inanimate and the animate. The animate is all of the termites living inside the colony, and the inanimate part is the structure itself, which is constructed by the termites.{{sfn|Bignell|Roisin|Lo|2010|p=3}} Nests can be broadly separated into three main categories: hypogeal, i.e subterranean (completely below ground), epigeal (protruding above the soil surface), and arboreal (built above ground, but always connected to the ground via [[#Shelter tubes|shelter tubes]]).<ref name=woodregulation/> Epigeal nests (mounds) protrude from the earth with ground contact and are made out of earth and mud.{{sfn|Bignell|Roisin|Lo|2010|p=20}} A nest has many functions such as providing a protected living space and providing shelter against predators. Most termites construct underground colonies rather than multifunctional nests and mounds.<ref name=divterm>{{cite journal |last1=Eggleton |first1=P. |last2=Bignell |first2=D.E.|last3=Sands |first3=W.A. |last4=Mawdsley |first4=N. A. |last5=Lawton|first5=J. H.|last6=Wood |first6=T.G. |last7=Bignell |first7=N.C. |title=The Diversity, Abundance and Biomass of Termites under Differing Levels of Disturbance in the Mbalmayo Forest Reserve, Southern Cameroon |journal=Philosophical Transactions of the Royal Society B: Biological Sciences|date=1996 |volume=351 |issue=1335 |pages=51–68 |doi=10.1098/rstb.1996.0004|bibcode=1996RSPTB.351...51E }}</ref> Primitive termites of today nest in wooden structures such as logs, stumps and the dead parts of trees, as did termites millions of years ago.<ref name=woodregulation>{{cite book |last1=Noirot|first1=C. |last2=Darlington |first2=J.P.E.C. |title=''Termite Nests: Architecture, Regulation and Defence'' in Termites: Evolution, Sociality, Symbioses, Ecology |date=2000 |publisher=Springer |doi=10.1007/978-94-017-3223-9_6 |isbn=978-94-017-3223-9 |pages=121–139}}</ref>
To build their nests, termites use a variety of resources such as faeces which have many desirable properties as a construction material.{{sfn|Bignell|Roisin|Lo|2010|p=21}} Other building materials include partly digested plant material, used in carton nests (arboreal nests built from faecal elements and wood), and soil, used in subterranean nest and mound construction. Not all nests are visible, as many nests in tropical forests are located underground.<ref name=divterm/> Species in the subfamily [[Apicotermitinae]] are good examples of subterranean nest builders, as they only dwell inside tunnels.{{sfn|Bignell|Roisin|Lo|2010|p=21}} Other termites live in wood, and tunnels are constructed as they feed on the wood. Nests and mounds protect the termites' soft bodies against desiccation, light, pathogens and parasites, as well as providing a fortification against predators.<ref>{{cite journal|last1=De Visse|first1=S.N.|last2=Freymann|first2=B.P. |last3=Schnyder |first3=H. |title=Trophic interactions among invertebrates in termitaria in the African savanna: a stable isotope approach |journal=Ecological Entomology |date=2008 |volume=33 |issue=6 |pages=758–764 |doi=10.1111/j.1365-2311.2008.01029.x|bibcode=2008EcoEn..33..758D |s2cid=33877331|url=http://mediatum.ub.tum.de/node?id=681224}}</ref> Nests made out of carton are particularly weak, and so the inhabitants use counter-attack strategies against invading predators.{{sfn|Bignell|Roisin|Lo|2010|p=22}}
Arboreal carton nests of [[mangrove swamp]]-dwelling ''Nasutitermes'' are enriched in [[lignin]] and depleted in cellulose and xylans. This change is caused by bacterial decay in the gut of the termites: they use their faeces as a carton building material. Arboreal termites nests can account for as much as 2% of above ground carbon storage in [[Puerto Rica]]n mangrove swamps. These ''Nasutitermes'' nests are mainly composed of partially biodegraded wood material from the stems and branches of mangrove trees, namely, ''[[Rhizophora mangle]]'' (red mangrove), ''[[Avicennia germinans]]'' (black mangrove) and ''[[Laguncularia racemosa]]'' (white mangrove).<ref name="Vane_et_al_2013">{{cite journal|last1=Vane|first1=C.H.|last2=Kim|first2=A.W.|last3=Moss-Hayes|first3=V.|last4=Snape|first4=C.E.|last5=Diaz|first5=M.C.|last6=Khan|first6=N.S.|last7=Engelhart|first7=S.E.|last8=Horton|first8=B.P.|title=Degradation of mangrove tissues by arboreal termites (''Nasutitermes acajutlae'') and their role in the mangrove C cycle (Puerto Rico): Chemical characterization and organic matter provenance using bulk δ13C, C/N, alkaline CuO oxidation-GC/MS, and solid-state|journal=Geochemistry, Geophysics, Geosystems|date=2013|volume=14|issue=8|pages=3176–3191|doi=10.1002/ggge.20194|url=http://nora.nerc.ac.uk/503347/1/ggge20194.pdf|bibcode=2013GGG....14.3176V|s2cid=130782273 }}</ref>
Some species build complex nests called polycalic nests; this habitat is called polycalism. Polycalic species of termites form multiple nests, or calies, connected by subterranean chambers.<ref name=wilson2014/> The termite genera ''[[Apicotermes]]'' and ''[[Trinervitermes]]'' are known to have polycalic species.<ref name=nasupol>{{cite journal |last1=Roisin |first1=Y. |last2=Pasteels |first2=J. M. |title=Reproductive mechanisms in termites: Polycalism and polygyny in ''Nasutitermes polygynus'' and ''N. costalis'' |journal=Insectes Sociaux |date=1986 |volume=33 |issue=2 |pages=149–167 |doi=10.1007/BF02224595|s2cid=41799894 }}</ref> Polycalic nests appear to be less frequent in mound-building species although polycalic arboreal nests have been observed in a few species of ''Nasutitermes''.<ref name=nasupol/>
===Mounds<!--"termite mounds" in Ant-hill article links here -->===
{{See also|Mound-building termites}}
{{Commons category|Termite mounds}}
Nests are considered mounds if they protrude from the earth's surface.{{sfn|Bignell|Roisin|Lo|2010|p=21}} A mound provides termites the same protection as a nest but is stronger.{{sfn|Bignell|Roisin|Lo|2010|p=22}} Mounds located in areas with torrential and continuous rainfall are at risk of mound erosion due to their clay-rich construction. Those made from carton can provide protection from the rain, and in fact can withstand high precipitation.{{sfn|Bignell|Roisin|Lo|2010|p=21}} Certain areas in mounds are used as strong points in case of a breach. For example, ''[[Cubitermes]]'' colonies build narrow tunnels used as strong points, as the diameter of the tunnels is small enough for soldiers to block.<ref>{{cite journal|last1=Perna|first1=A.|last2=Jost|first2=C.|last3=Couturier|first3=E.|last4=Valverde|first4=S.|last5=Douady|first5=S.|last6=Theraulaz|first6=G.|title=The structure of gallery networks in the nests of termite ''Cubitermes'' spp. revealed by X-ray tomography.|journal=Die Naturwissenschaften|date=2008|volume=95|issue=9|pages=877–884|doi=10.1007/s00114-008-0388-6|pmid=18493731|bibcode = 2008NW.....95..877P |s2cid=15326313}}</ref> A highly protected chamber, known as the "queen's cell", houses the queen and king and is used as a last line of defence.{{sfn|Bignell|Roisin|Lo|2010|p=22}}
Species in the genus ''Macrotermes'' arguably build the most complex structures in the insect world, constructing enormous mounds.{{sfn|Bignell|Roisin|Lo|2010|p=21}} These mounds are among the largest in the world, reaching a height of 8 to 9 metres (26 to 29 feet), and consist of chimneys, pinnacles and ridges.<ref name=Britannica/> Another termite species, ''[[Amitermes meridionalis]]'', can build nests 3&nbsp;to 4&nbsp;metres (9&nbsp;to 13 feet) high and 2.5 metres (8&nbsp;feet) wide. The tallest mound ever recorded was 12.8 metres (42&nbsp;ft) long found in the Democratic Republic of the Congo.<ref>{{Cite book|title=Guinness World Records 2014|last=Glenday|first=Craig|year=2014|isbn=978-1-908843-15-9|pages=[https://archive.org/details/guinnessworldrec0000unse_r3e7/page/33 33]|publisher=Guinness World Records Limited |url=https://archive.org/details/guinnessworldrec0000unse_r3e7/page/33}}</ref>
The sculptured mounds sometimes have elaborate and distinctive forms, such as those of the compass termite (''Amitermes meridionalis'' and ''A. laurensis''), which builds tall, wedge-shaped mounds with the long axis oriented approximately north–south, which gives them their common name.<ref name=jacklyn1991>{{cite journal |last1=Jacklyn |first1=P. |title=Evidence for Adaptive Variation in the Orientation of ''Amitermes'' (Isoptera, Termitinae) Mounds From Northern Australia |journal=Australian Journal of Zoology |date=1991 |volume=39 |issue=5 |pages=569 |doi=10.1071/ZO9910569}}</ref><ref name=jacklyn2002>{{cite journal |last1=Jacklyn |first1=P.M. |last2=Munro |first2=U. |title=Evidence for the use of magnetic cues in mound construction by the termite ''Amitermes meridionalis'' (Isoptera : Termitinae) |journal=Australian Journal of Zoology |date=2002 |volume=50 |issue=4 |pages=357 |doi=10.1071/ZO01061}}</ref> This orientation has been experimentally shown to assist [[thermoregulation]]. The north–south orientation causes the internal temperature of a mound to increase rapidly during the morning while avoiding overheating from the midday sun. The temperature then remains at a plateau for the rest of the day until the evening.<ref>{{cite journal |last1=Grigg |first1=G.C. |title=Some Consequences of the Shape and Orientation of 'magnetic' Termite Mounds |journal=Australian Journal of Zoology |date=1973 |volume=21 |issue=2 |pages=231–237 |doi=10.1071/ZO9730231|url=http://espace.library.uq.edu.au/view/UQ:10143/gg_ajz_21_73.pdf }}</ref>
<gallery widths="200px" class="center">
<!-- Please note that punctuation in the alt text is intended to assist recitation by a screen reader and is not necessarily intended to be grammatical. Please use a screen reader to validate any changes in alt text. -->
File:RayNorris termite cathedral mounds.jpg|alt=. These termite mounds have a base shaped like the base of a tree, about two meters wide and a meter high. From this base, rounded chimneys from half a meter to a meter in diameter rise to a total height of about four or five meters. The chimneys are fused together with ridges between, and terminate in rounded pinnacles at the top.|[[Cathedral termite|Cathedral mound]]s in the [[Northern Territory]], [[Australia]]
File:Termite Magnetic DSC03613.jpg|alt=. Hundreds of compass termite mounds are visible in this photo of a field in northern Australia. The chisel-shaped mounds range from several centimeters to several meters in height.|Mounds of "compass" or "magnetic" termites (''Amitermes'') oriented north–south, thereby avoiding mid-day heat
File:Termitenhuegel.jpg|alt=. This termite mound is about three meters in height and four meters across. The mound chimneys are about a meter in diameter and fuse together to form a rounded top.|Termite mound in [[Queensland]], Australia
File:Termites in a mound.jpg|alt=. The photographer has broken off a piece of a mound to show the mound's interior. Dozens of tunnels have been exposed, and hundreds of soldiers have emerged to guard the breech in the wall.|Termites in a mound, [[Analamazoatra Reserve]], [[Madagascar]]
<!--File:Termite_mound_with_comparison.jpg|Humans and trees for size comparison-->
File:Termitenhügel Namibia.jpg|Termite mound in [[Namibia]]
</gallery>
===Shelter tubes===
[[File:Termite-nest-tunnels.jpg|thumb|upright|alt=Photo taken upwards from ground level of shelter tubes going up the shaded side of a tree. Where the main trunk of the tree divides into separate major branches, the shelter tube also branches. Although the nests are not visible in this photo, the branches of the shelter tube presumably lead up to polycalic sister colonies of the arboreal termites that built these tubes.|Nasutiterminae shelter tubes on a tree trunk provide cover for the trail from nest to forest floor.]]
Termites construct shelter tubes, also known as earthen tubes or mud tubes, that start from the ground. These shelter tubes can be found on walls and other structures.<ref name=term1996>{{cite book |last1=Hadlington |first1=P. |title=Australian Termites and Other Common Timber Pests |date=1996 |publisher=New South Wales University Press |location=Kensington, NSW, Australia |isbn=978-0-86840-399-1|pages=28–30 |edition=2nd}}</ref> Constructed by termites during the night, a time of higher humidity, these tubes provide protection to termites from potential predators, especially ants.<ref name=bolinas>{{cite book|last1=Kahn|first1=L.|last2=Easton|first2=B.|title=Shelter II|date=2010|publisher=Shelter Publications|location=Bolinas, California|isbn=978-0-936070-49-0|page=198}}</ref> Shelter tubes also provide high humidity and darkness and allow workers to collect food sources that cannot be accessed in any other way.<ref name=term1996/> These passageways are made from soil and faeces and are normally brown in colour. The size of these shelter tubes depends on the number of food sources that are available. They range from less than 1&nbsp;cm to several cm in width, but may be dozens of metres in length.<ref name=bolinas/>
== Relationship with humans ==
=== As pests ===
[[File:Termite mound on runway at Khorixas (2018).jpg|thumb|Termite mound as an obstacle on a runway at [[Khorixas]] ([[Namibia]])]]
[[File:Termite damage.JPG|thumb|200px|left|Termite damage on external structure]]
Owing to their wood-eating habits, many termite species can do significant damage to unprotected buildings and other wooden structures.<ref name=pests2000/> Termites play an important role as decomposers of wood and vegetative material, and the conflict with humans occurs where structures and landscapes containing structural wood components, cellulose derived structural materials and ornamental vegetation provide termites with a reliable source of food and moisture.<ref>{{cite book|last=Thorne, Ph.D|first=Barbara L. |date=1999 |title=NPMA Research Report On Subterranean Termites |location=Dunn Loring, VA |publisher=NPMA|page=22|url=https://entomology.umd.edu/thorne-barbara-l.html}}</ref> Their habit of remaining concealed often results in their presence being undetected until the timbers are severely damaged, with only a thin exterior layer of wood remaining, which protects them from the environment.<ref>{{cite web|title=Termites|url=http://www.vba.vic.gov.au/consumer-resources/building/pages/termites|work=Victorian Building Authority|publisher=Government of Victoria|access-date=20 September 2015|date=2014|archive-date=3 February 2018|archive-url=https://web.archive.org/web/20180203051240/http://www.vba.vic.gov.au/consumer-resources/building/pages/termites|url-status=dead}}</ref> Of the 3,106 species known, only 183 species cause damage; 83 species cause significant damage to wooden structures.<ref name=pests2000>{{cite book|last1=Su|first1=N.Y.|last2=Scheffrahn|first2=R.H.|title=''Termites as Pests of Buildings'' in Termites: Evolution, Sociality, Symbioses, Ecology|date=2000|publisher=Springer Netherlands|doi=10.1007/978-94-017-3223-9_20|isbn=978-94-017-3223-9|pages=437–453}}</ref> In North America, 18 subterranean species are pests;<ref>{{cite book|last=Thorne, Ph.D|first=Barbara L. |date=1999 |title=NPMA Research Report On Subterranean Termites |location=Dunn Loring, VA |publisher=NPMA|page=2|url=https://entomology.umd.edu/thorne-barbara-l.html}}</ref> in Australia, 16 species have an economic impact; in the Indian subcontinent 26 species are considered pests, and in tropical Africa, 24. In Central America and the West Indies, there are 17 pest species.<ref name=pests2000/> Among the termite genera, ''Coptotermes'' has the highest number of pest species of any genus, with 28 species known to cause damage.<ref name=pests2000/> Less than 10% of drywood termites are pests, but they infect wooden structures and furniture in tropical, subtropical and other regions. Dampwood termites only attack lumber material exposed to rainfall or soil.<ref name=pests2000/>
Drywood termites thrive in warm climates, and human activities can enable them to invade homes since they can be transported through contaminated goods, containers and ships.<ref name=pests2000/> Colonies of termites have been seen thriving in warm buildings located in cold regions.<ref>{{cite journal|last1=Grace|first1=J.K.|last2=Cutten|first2=G.M.|last3=Scheffrahn|first3=R.H.|last4=McEkevan|first4=D.K.|title=First infestation by ''Incisitermes minor'' of a Canadian building (Isoptera: Kalotermitidae)|journal=Sociobiology|date=1991|volume=18|pages=299–304}}</ref> Some termites are considered invasive species. ''Cryptotermes brevis'', the most widely introduced invasive termite species in the world, has been introduced to all the islands in the West Indies and to Australia.<ref name=Heather1971/><ref name=pests2000/>
[[File:House stumps eaten by termites.jpg|thumb|right|200px|Termite damage in wooden house stumps]]
In addition to causing damage to buildings, termites can also damage food crops.<ref name=Sands1973>{{cite journal|last1=Sands|first1=W.A.|title=Termites as Pests of Tropical Food Crops|journal=Tropical Pest Management|date=1973|volume=19|issue=2|pages=167–177|doi=10.1080/09670877309412751}}</ref> Termites may attack trees whose resistance to damage is low but generally ignore fast-growing plants. Most attacks occur at harvest time; crops and trees are attacked during the dry season.<ref name=Sands1973/>
In Australia, at a cost of more than {{AUD|1.5 billion}} per year,<ref>[https://www.uts.edu.au/research-and-teaching/our-research/sustainability/our-research/termites-quietly-reveal-their-secrets Termites quietly reveal their secrets] [[University of Technology Sydney]]. Retrieved 3 April 2023.</ref> termites cause more damage to houses than fire, floods and storms combined.<ref>[https://www.vba.vic.gov.au/consumers/guides/termites Termites] Victorian Building Authority. Retrieved 3 April 2023.</ref> In Malaysia, it is estimated that termites caused about RM400 million of damages to properties and buildings.<ref>{{Cite web|url=https://www.freemalaysiatoday.com/category/leisure/property/2021/10/02/a-guide-to-termite-infestations-in-malaysia/|title=A guide to termite infestations in Malaysia &#124; Free Malaysia Today (FMT)|date=2 October 2021 }}</ref> The damage caused by termites costs the southwestern United States approximately $1.5&nbsp;billion each year in wood structure damage, but the true cost of damage worldwide cannot be determined.<ref name=pests2000/><ref name=agricultural2010/> Drywood termites are responsible for a large proportion of the damage caused by termites.<ref>{{cite journal|last1=Su|first1=N.Y.|last2=Scheffrahn|first2=R.H.|title=Economically important termites in the United States and their control|journal=Sociobiology|date=1990|volume=17|pages=77–94|url=http://flrec.ifas.ufl.edu/pdfs/su_pub/su045_ecotmt.pdf|url-status=dead|archive-url=https://web.archive.org/web/20110812214933/http://flrec.ifas.ufl.edu/pdfs/Su_pub/Su045_EcoTMT.pdf|archive-date=2011-08-12}}</ref>  The goal of termite control is to keep structures and susceptible ornamental plants free from termites.;<ref>{{cite book|last=Thorne, Ph.D|first=Barbara L. |date=1999 |title=NPMA Research Report On Subterranean Termites |location=Dunn Loring, VA |publisher=NPMA|page=40|url=https://entomology.umd.edu/thorne-barbara-l.html}}</ref>  Structures may be homes or business, or elements such as wooden fence posts and telephone poles.  Regular and thorough inspections by a trained professional may be necessary to detect termite activity in the absence of more obvious signs like termite swarmers or alates inside or adjacent to a structure.  Termite monitors made of wood or cellulose adjacent to a structure may also provide indication of termite foraging activity where it will be in conflict with humans. Termites can be controlled by application of [[Bordeaux mixture]] or other substances that contain [[copper]] such as [[chromated copper arsenate]].<ref name="cus">{{cite web |last1=Elliott |first1=Sara |title=How can copper keep termites at bay? |url=https://home.howstuffworks.com/home-improvement/household-safety/copper-stop-termites2.htm |publisher=HowStuffWorks |date=26 May 2009}}</ref>  In the United states, application of a soil termiticide with the active ingredient [[Fipronil]], such as Termidor SC or Taurus SC, by a licensed professional,<ref>{{Cite web|title=Questions and Answers About Termites|url=https://www.pestboard.ca.gov/forms/termites.pdf|access-date=19 April 2021|website=Department of Consumer Affairs, Structural Pest Control Board of California}}</ref> is a common remedy approved by the Environmental Protection Agency for economically significant subterranean termites.<ref>{{Cite web|title=EPA Registration and Label for Taurus SC Termiticide|url=https://www3.epa.gov/pesticides/chem_search/ppls/053883-00279-20110915.pdf|website=EPA.gov}}</ref><ref>{{Cite web|title=EPA Registration and Label for Termidor SC|url=https://www3.epa.gov/pesticides/chem_search/ppls/007969-00210-20041015.pdf|access-date=19 April 2021|website=EPA.gov}}</ref>  A growing demand for alternative, green, and "more natural" extermination methods has increased demand for mechanical and biological control methods such as [[orange oil]].
To better control the population of termites, various methods have been developed to track termite movements.<ref name=agricultural2010>{{cite news|last1=Flores|first1=A.|title=New Assay Helps Track Termites, Other Insects|url=http://www.ars.usda.gov/is/pr/2010/100217.htm|access-date=15 January 2015|work=Agricultural Research Service|publisher=United States Department of Agriculture|date=17 February 2010}}</ref> One early method involved distributing termite bait laced with [[immunoglobulin G]] (IgG) marker proteins from rabbits or chickens. Termites collected from the field could be tested for the rabbit-IgG markers using a rabbit-IgG-specific [[assay]]. More recently developed, less expensive alternatives include tracking the termites using egg white, cow milk, or soy milk proteins, which can be sprayed on termites in the field. Termites bearing these proteins can be traced using a protein-specific [[ELISA]] test.<ref name=agricultural2010/> [[RNAi]] insecticides specific to termites [[insecticide development|are in development]].<ref name="Assay for RNAi resistance in termites"/> One factor reducing [[investment]] in its [[research and development]] is concern about high potential for [[Pesticide resistance|resistance evolution]].<ref name="Assay for RNAi resistance in termites">{{Cite journal|number=1|volume=43|year=2023|department=Review paper|pages=55–68|last1=Mogilicherla|first1=Kanakachari|last2=Chakraborty|first2=Amrita|last3=Taning|first3=Clauvis Nji Tizi|last4=Smagghe|first4=Guy|last5=Roy|first5=Amit|doi=10.1127/entomologia/2022/1636|title=RNAi in termites (Isoptera): current status and prospects for pest management|journal=Entomologia Generalis |hdl=1854/LU-01H7T2H1DB5XMEKN7APN3SEPYR|hdl-access=free}}</ref>
In 1994, termites, of the species ''[[Reticulitermes]] grassei'', were identified in two bungalows in [[Saunton]], [[Devon]]. Anecdotal evidence suggests the infestation could date back 70 years before the official identification. There are reports that gardeners had seen white ants and that a greenhouse had had to be replaced in the past. The Saunton infestation was the first and only colony ever recorded in the UK. In 1998, Termite Eradication Programme was set-up, with the intention of containing and eradicating the colony. The TEP was managed by the Ministry of Housing, Communities & Local Government (now the [[Department for Levelling Up, Housing and Communities]].) The TEP used "insect growth regulators" to prevent the termites from reaching maturity and reproducing. In 2021, the UK's Termite Eradication Programme announced the eradication of the colony, the first time a country has eradicated termites.<ref>{{Cite web|last=Pidd|first=Helen|date=21 December 2021|title='A world first': Devon calls victory in 27-year war on termites|url=https://www.theguardian.com/uk-news/2021/dec/21/a-world-first-devon-calls-victory-in-27-year-war-on-termites|url-status=live|archive-url=https://web.archive.org/web/20211221234612/https://www.theguardian.com/uk-news/2021/dec/21/a-world-first-devon-calls-victory-in-27-year-war-on-termites|archive-date=21 December 2021|access-date=22 December 2021|website=The Guardian|language=en}}</ref>
===As food===
{{See also|Entomophagy in humans}}
[[File:Collecting Ngumbi.jpg|thumb|Mozambican boys from the Yawo tribe collecting flying termites]]
[[File:Ngumbi.jpg|thumb|These flying alates were collected as they came out of their nests in the ground during the early days of the rainy season.]]
43 termite species are used as food by humans or are fed to livestock.<ref name=greview>{{cite journal|last1=Figueirêdo|first1=R.E.C.R.|last2=Vasconcellos|first2=A.|last3=Policarpo|first3=I.S.|last4=Alves|first4=R.R.N.|title=Edible and medicinal termites: a global overview|journal=Journal of Ethnobiology and Ethnomedicine|date=2015|volume=11|issue=1|pages=29|doi=10.1186/s13002-015-0016-4|pmid=25925503|pmc=4427943 |doi-access=free }}</ref> These insects are particularly important in impoverished countries where malnutrition is common, as the [[protein]] from termites can help improve the human diet. Termites are consumed in many regions globally, but this practice has only become popular in developed nations in recent years.<ref name=greview/>
Termites are consumed by people in many different cultures around the world. In many parts of Africa, the [[alate]]s are an important factor in the diets of native populations.<ref name=delicacy>{{cite book|last1=Nyakupfuka|first1=A.|title=Global Delicacies: Discover Missing Links from Ancient Hawaiian Teachings to Clean the Plaque of your Soul and Reach Your Higher Self.|date=2013|publisher=BalboaPress|location=Bloomington, Indiana|isbn=978-1-4525-6791-4|pages=40–41}}</ref> Groups have different ways of collecting or cultivating insects; sometimes collecting soldiers from several species. Though harder to acquire, queens are regarded as a delicacy.<ref name=B1951>{{cite book|last1=Bodenheimer|first1=F.S.|title=Insects as Human Food: A Chapter of the Ecology of Man|date=1951|publisher=[[Springer Science+Business Media|Springer]]|location=[[Netherlands]]|isbn=978-94-017-6159-8|pages=331–350}}</ref> Termite alates are high in nutrition with adequate levels of [[fat]] and protein. They are regarded as pleasant in taste, having a nut-like flavour after they are cooked.<ref name=delicacy/>
Alates are collected when the rainy season begins. During a nuptial flight, they are typically seen around lights to which they are attracted, and so nets are set up on lamps and captured alates are later collected. The wings are removed through a technique that is similar to [[winnowing]]. The best result comes when they are lightly roasted on a hot plate or fried until crisp. [[cooking oil|Oil]] is not required as their bodies usually contain sufficient amounts of oil. Termites are typically eaten when livestock is lean and tribal crops have not yet developed or produced any food, or if food stocks from a previous growing season are limited.<ref name=delicacy/>
In addition to Africa, termites are consumed in local or tribal areas in Asia and North and South America. In Australia, [[Indigenous Australians]] are aware that termites are edible but do not consume them even in times of scarcity; there are few explanations as to why.<ref name=delicacy/><ref name=B1951/> Termite mounds are the main sources of soil consumption ([[geophagy]]) in many countries including [[Kenya]], [[Tanzania]], [[Zambia]], [[Zimbabwe]] and [[South Africa]].<ref>{{cite journal|last1=Geissler|first1=P.W.|title=The significance of earth-eating: social and cultural aspects of geophagy among Luo children|journal=Africa|date=2011|volume=70|issue=4|pages=653–682|doi=10.3366/afr.2000.70.4.653|s2cid=145754470}}</ref><ref>{{cite journal|last1=Knudsen|first1=J.W.|title=Akula udongo (earth eating habit): a social and cultural practice among Chagga women on the slopes of Mount Kilimanjaro|journal=African Journal of Indigenous Knowledge Systems|date=2002|volume=1|issue=1|pages=19–26|issn=1683-0296|oclc=145403765|doi=10.4314/indilinga.v1i1.26322}}</ref><ref>{{cite journal|last1=Nchito|first1=M.|last2=Wenzel Geissler|first2=P.|last3=Mubila|first3=L.|last4=Friis|first4=H.|last5=Olsen|first5=A.|title=Effects of iron and multimicronutrient supplementation on geophagy: a two-by-two factorial study among Zambian schoolchildren in Lusaka|journal=Transactions of the Royal Society of Tropical Medicine and Hygiene|date=2004|volume=98|issue=4|pages=218–227|doi=10.1016/S0035-9203(03)00045-2|pmid=15049460}}</ref><ref>{{cite journal|last1=Saathoff|first1=E.|last2=Olsen|first2=A.|last3=Kvalsvig|first3=J.D.|last4=Geissler|first4=P.W.|title=Geophagy and its association with geohelminth infection in rural schoolchildren from northern KwaZulu-Natal, South Africa|journal=Transactions of the Royal Society of Tropical Medicine and Hygiene|date=2002|volume=96|issue=5|pages=485–490|doi=10.1016/S0035-9203(02)90413-X|pmid=12474473}}</ref> Researchers have suggested that termites are suitable candidates for human consumption and [[Space farming|space agriculture]], as they are high in protein and can be used to convert inedible waste to consumable products for humans.<ref>{{cite journal|last1=Katayama|first1=N.|last2=Ishikawa|first2=Y.|last3=Takaoki|first3=M.|last4=Yamashita|first4=M.|last5=Nakayama|first5=S.|last6=Kiguchi|first6=K.|last7=Kok|first7=R.|last8=Wada|first8=H.|last9=Mitsuhashi|first9=J.|title=Entomophagy: A key to space agriculture|journal=Advances in Space Research|date=2008|volume=41|issue=5|pages=701–705|doi=10.1016/j.asr.2007.01.027|bibcode=2008AdSpR..41..701S|url=http://fr.khepri.eu/wp-content/uploads/sites/3/2013/10/Entomophagy-A-key-to-space-agriculture.pdf}}</ref>
===In agriculture===
[[File:Termites marked with traceable protiens.jpg|thumb|left|Scientists have developed a more affordable method of tracing the movement of termites using traceable proteins.<ref name="agricultural2010"/>]]
Termites can be major agricultural pests, particularly in East Africa and North Asia, where crop losses can be severe (3–100% in crop loss in Africa).<ref>{{cite journal|last1=Mitchell|first1=J.D.|title=Termites as pests of crops, forestry, rangeland and structures in Southern Africa and their control|journal=Sociobiology|date=2002|volume=40|issue=1|pages=47–69|url=http://cat.inist.fr/?aModele=afficheN&cpsidt=13648716|issn=0361-6525}}</ref> Counterbalancing this is the greatly improved water infiltration where termite tunnels in the soil allow rainwater to soak in deeply, which helps reduce runoff and consequent soil erosion through [[bioturbation#Terrestrial|bioturbation]].<ref name=aseanbiodiversity.info>{{cite journal|last1=Löffler|first1=E.|last2=Kubiniok|first2=J.|title=Landform development and bioturbation on the Khorat plateau, Northeast Thailand|journal=Natural History Bulletin of the Siam Society|date=1996|volume=44|pages=199–216|url=http://www.siamese-heritage.org/nhbsspdf/vol041-050/NHBSS_044_2h_Loffler_LandformDevelopme.pdf}}</ref> In South America, cultivated plants such as eucalyptus, upland rice and [[sugarcane]] can be severely damaged by termite infestations, with attacks on leaves, roots and woody tissue. Termites can also attack other plants, including [[cassava]], [[Coffea|coffee]], [[Gossypium|cotton]], fruit trees, [[maize]], [[peanut]]s, soybeans and vegetables.<ref name=cap2008/> Mounds can disrupt farming activities, making it difficult for farmers to operate farming machinery; however, despite farmers' dislike of the mounds, it is often the case that no net loss of production occurs.<ref name=cap2008/> Termites can be beneficial to agriculture, such as by boosting [[crop yield]]s and enriching the soil. Termites and ants can re-colonise untilled land that contains crop stubble, which colonies use for nourishment when they establish their nests. The presence of nests in fields enables larger amounts of rainwater to soak into the ground and increases the amount of nitrogen in the soil, both essential for the growth of crops.<ref>{{cite journal |last1=Evans |first1=T.A. |last2=Dawes |first2=T.Z. |last3=Ward |first3=P.R. |last4=Lo |first4=N. |title=Ants and termites increase crop yield in a dry climate |journal=Nature Communications |date=2011 |volume=2 |pages=262 |bibcode=2011NatCo...2..262E|doi=10.1038/ncomms1257|pmid=21448161|pmc=3072065}}</ref>
{{Clear}}
===In science and technology===
{{See also|Renewable energy|Termite-inspired robots|Sustainable architecture}}
The termite gut has inspired various research efforts aimed at replacing [[fossil fuels]] with cleaner, renewable energy sources.<ref name=doeinstitute/> Termites are efficient [[bioreactor]]s, theoretically capable of producing two litres of [[hydrogen]] from a single sheet of paper.<ref>{{cite news|last1=Hirschler|first1=B.|title=Termites' gut reaction set for biofuels|url=http://www.abc.net.au/science/articles/2007/11/22/2097855.htm|access-date=8 January 2015|work=ABC News|date=22 November 2007}}</ref> Approximately 200 species of microbes live inside the termite hindgut, releasing the hydrogen that was trapped inside wood and plants that they digest.<ref name=doeinstitute>{{cite web|title=Termite Power |url=http://www.jgi.doe.gov/education/bioenergy/bioenergy_4.html |publisher=United States Department of Energy |access-date=11 September 2015 |archive-url=https://web.archive.org/web/20060922180946/http://www.jgi.doe.gov/education/bioenergy/bioenergy_4.html|archive-date=22 September 2006 |date=14 August 2006 |work=DOE Joint Genome Institute |url-status=unfit }}</ref><ref>{{cite news|last1=Roach|first1=J.|title=Termite Power: Can Pests' Guts Create New Fuel?|url=http://news.nationalgeographic.com/news/2006/03/0314_060314_termite.html|archive-url=https://web.archive.org/web/20060316155632/http://news.nationalgeographic.com/news/2006/03/0314_060314_termite.html|url-status=dead|archive-date=March 16, 2006|access-date=11 September 2015|work=National Geographic News|date=14 March 2006}}</ref> Through the action of unidentified enzymes in the termite gut, [[lignocellulose]] [[polymer]]s are broken down into sugars and are transformed into hydrogen. The bacteria within the gut turns the sugar and hydrogen into [[cellulose acetate]], an [[acetate]] [[ester]] of cellulose on which termites rely for energy.<ref name=doeinstitute/> [[Microbiota#Metagenomic sequencing|Community DNA sequencing]] of the microbes in the termite hindgut has been employed to provide a better understanding of the [[metabolic pathway]].<ref name=doeinstitute/> Genetic engineering may enable hydrogen to be generated in bioreactors from woody biomass.<ref name=doeinstitute/>
The development of [[autonomous robot]]s capable of constructing intricate structures without human assistance has been inspired by the complex mounds that termites build.<ref name=Terminspired>{{cite journal |last1=Werfel|first1=J.|last2=Petersen|first2=K.|last3=Nagpal|first3=R. |title=Designing Collective Behavior in a Termite-Inspired Robot Construction Team |journal=Science|date=2014|volume=343|issue=6172|pages=754–758|doi=10.1126/science.1245842|pmid=24531967|bibcode= 2014Sci...343..754W|s2cid=38776920}}</ref> These robots work independently and can move by themselves on a tracked grid, capable of climbing and lifting up bricks. Such robots may be useful for future projects on Mars, or for building [[levee]]s to prevent flooding.<ref>{{cite journal|last1=Gibney|first1=E.|title=Termite-inspired robots build castles|journal=Nature|date=2014|doi=10.1038/nature.2014.14713|s2cid=112117767|url=http://www.nature.com/news/termite-inspired-robots-build-castles-1.14713}}</ref>
Termites use sophisticated means to control the temperatures of their mounds. [[#Mounds|As discussed above]], the shape and orientation of the mounds of the Australian compass termite stabilises their internal temperatures during the day. As the towers heat up, the [[solar chimney]] effect ([[stack effect]]) creates an updraft of air within the mound.<ref name=AAK2013>{{cite web|title=Termites Green Architecture in the Tropics|url=http://www.thearchitectmagazine.com/termites-green-architecture-in-the-tropics/|website=The Architect|publisher=Architectural Association of Kenya|access-date=17 October 2015|archive-date=22 March 2016|archive-url=https://web.archive.org/web/20160322132820/http://www.thearchitectmagazine.com/termites-green-architecture-in-the-tropics/|url-status=dead}}</ref> Wind blowing across the tops of the towers enhances the circulation of air through the mounds, which also include side vents in their construction. The solar chimney effect has been in use for centuries in the [[Middle East]] and [[Near East]] for passive cooling, as well as in Europe by the [[Ancient Rome|Romans]].<ref>{{cite journal|last1=Tan|first1=A.|last2=Wong|first2=N.|title=Parameterization Studies of Solar Chimneys in the Tropics|journal=Energies|date=2013|volume=6|issue=1|pages=145–163|doi=10.3390/en6010145|doi-access=free}}</ref> It is only relatively recently, however, that climate responsive construction techniques have become incorporated into modern architecture. Especially in Africa, the stack effect has become a popular means to achieve natural ventilation and passive cooling in modern buildings.<ref name=AAK2013/>
===In culture===
[[File:Eastgate Centre, Harare, Zimbabwe.jpg|thumb|right|250px|The pink-hued Eastgate Centre]]
The [[Eastgate Centre, Harare|Eastgate Centre]] is a shopping centre and office block in central [[Harare]], Zimbabwe, whose architect, [[Mick Pearce]], used [[passive cooling]] inspired by that used by the local termites.<ref name=eastgate>{{cite news|last1=Tsoroti|first1=S.|title=What's that building? Eastgate Mall|url=http://www.hararenews.co.zw/2014/05/whats-that-building-eastgate-mall/|access-date=8 January 2015|work=Harare News|date=15 May 2014|archive-date=11 April 2021|archive-url=https://web.archive.org/web/20210411182543/http://www.hararenews.co.zw/2014/05/whats-that-building-eastgate-mall/|url-status=dead}}</ref> It was the first major building exploiting termite-inspired cooling techniques to attract international attention. Other such buildings include the Learning Resource Center at the [[Catholic University of Eastern Africa]] and the [[Council House 2]] building in [[Melbourne]], Australia.<ref name=AAK2013/>
Few zoos hold termites, due to the difficulty in keeping them captive and to the reluctance of authorities to permit potential pests. One of the few that do, the [[Zoo Basel]] in [[Switzerland]], has two thriving ''Macrotermes bellicosus'' populations&nbsp;– resulting in an event very rare in captivity: the mass migrations of young flying termites. This happened in September 2008, when thousands of male termites left their mound each night, died, and covered the floors and water pits of the house holding their exhibit.<ref>
{{cite news
|url=http://www.nzz.ch/nachrichten/panorama/im_zoo_basel_fliegen_die_termiten_aus__1.848530.html
|title=Im Zoo Basel fliegen die Termiten aus |newspaper=Neue Zürcher Zeitung |date=8 February 2014 |access-date=21 May 2011 |language=de}}</ref>
African tribes in several countries have termites as [[totem]]s, and for this reason tribe members are forbidden to eat the reproductive alates.<ref>{{cite journal|last1=Van-Huis|first1=H.|title=Insects as food in Sub-Saharan Africa|journal=Insect Science and Its Application|date=2003|volume=23|issue=3|pages=163–185|url=http://ag.udel.edu/delpha/4434.pdf|doi=10.1017/s1742758400023572|s2cid=198497332|access-date=2015-09-20|archive-date=2017-07-13|archive-url=https://web.archive.org/web/20170713120108/http://ag.udel.edu/delpha/4434.pdf|url-status=dead}}</ref> Termites are widely used in traditional popular medicine; they are used as treatments for diseases and other conditions such as asthma, [[bronchitis]], [[hoarseness]], influenza, [[sinusitis]], [[tonsillitis]] and whooping cough.<ref name=greview/> In Nigeria, ''[[Macrotermes nigeriensis]]'' is used for spiritual protection and to treat wounds and sick pregnant women. In Southeast Asia, termites are used in ritual practices. In Malaysia, Singapore and Thailand, termite mounds are commonly worshiped among the populace.<ref name=Neoh2013>{{cite journal|last1=Neoh|first1=K.B.|title=Termites and human society in Southeast Asia|journal=The Newsletter|date=2013|volume=30|issue=66|pages=1–2|url=http://www.iias.nl/sites/default/files/IIAS_NL66_3031.pdf}}</ref> Abandoned mounds are viewed as structures created by spirits, believing a local guardian dwells within the mound; this is known as [[Keramat]] and Datok Kong.<ref>{{Cite web |last=Pratama |first=Rian |date=2024-04-26 |title=Tips Termite Prevention untuk Bisnis Anda |url=https://umas.co.id/tips-termite-prevention-untuk-bisnis-anda/ |access-date=2024-07-30 |website=UMAS Pest Control |language=id}}</ref> In urban areas, local residents construct red-painted shrines over mounds that have been abandoned, where they pray for good health, protection and luck.<ref name=Neoh2013/>
==See also==
* [[Mound-building termites]]
* [[Stigmergy]]
* [[Termite shield]]
* [[Xylophagy]]
==Notes==
{{Reflist|group=note}}
==References==
{{Reflist|30em|refs=
<ref name="fossilworks">{{Cite web
| url = https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=220276
| title = Fossilworks, Gateway to the Paleobiology Database
| last1 = Behrensmeyer
| first1 = A. K.
| last2 = Turner
| first2 = A.
| access-date = 2021-12-17
| archive-date = 2021-12-13
| archive-url = https://web.archive.org/web/20211213003706/http://www.fossilworks.org/cgi-bin/bridge.pl?a=taxonInfo&taxon_no=220276
| url-status = live
}}</ref>
<ref name=Engel2009>
{{Cite journal
| title = Termites (Isoptera): their phylogeny, classification, and rise to ecological dominance
| date = 2009
| last1 = Engel | first1 = M.S.
| last2 = Grimaldi | first2 = D.A.
| last3 = Krishna | first3 = K.
| journal = American Museum Novitates
| issue = 3650
| pages = 1–27
| issn = 0003-0082
| doi = 10.1206/651.1
| hdl = 2246/5969
| s2cid = 56166416
| url = https://www.biodiversitylibrary.org/item/288387
| hdl-access = free
}}</ref>
}}
===Cited literature===
*{{cite book|last1=Bignell|first1=D.E.|last2=Roisin|first2=Y.|last3=Lo|first3=N.|title=Biology of Termites: a Modern Synthesis|date=2010|publisher=Springer|location=Dordrecht|isbn=978-90-481-3977-4|edition=1st}}
*{{cite book|last1=Schmid-Hempel|first1=P.|title=Parasites in Social Insects|date=1998|publisher=Princeton University Press|location=New Jersey|isbn=978-0-691-05924-2}}
[https://www.rdfm.com.au/services/pest-control-sydney/ pest control Sydney]
==External links==
{{wiktionary|termite}}
{{Commons category|Isoptera}}
{{Wikispecies|Isoptera|Isoptera}}
* {{wsPSM2|The White Ant: A Theory|27|October 1885}}
* [http://www.ento.csiro.au/education/insects/isoptera.html Isoptera: termites] at CSIRO Australia Entomology
* [https://www.ibiology.org/ecology/termite-gut/ Jared Leadbetter seminar: Termites and Their Symbiotic Gut Microbes]
{{Blattodea}}
{{Eusociality}}
{{Insects in culture}}
{{Taxonbar|from1=Q546583|from2=Q21069206}}
{{Authority control}}
[[Category:Termites| ]]
[[Category:Articles containing video clips]]
[[Category:Building defects]]
[[Category:Household pest insects]]
[[Category:Insects in culture]]
[[Category:Extant Early Cretaceous first appearances]]
[[Category:Symbiosis]]
[[Category:Superorganisms]]