Editing Termite (section)

==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 other 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]].<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 phylogenetic relationship, and is considered to be the sister-group to termites.<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><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>  It had 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> but subsequent 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">{{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> [[File:Isoptera.jpg|thumb|Macro image of a worker.]]The primitive [[giant northern termite]] (''Mastotermes darwiniensis'') exhibits numerous basal characteristics similar to other cockroaches 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> 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|bibcode=1997AnRES..28...27T |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 name=":8" /><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=[[Archotermopsidae]]
       |2={{clade
          |1=[[Hodotermopsidae]]
          |2=[[Hodotermitidae]]
       }}
    }}
 }}
 |targetB={SUBCLADE_B}
 |subcladeB={{clade
    |1=[[Kalotermitidae]]
    |2={{clade
       |label1=[[Neoisoptera]]
       |1={SUBCLADE_C}
    }}
 }}
 |targetC={SUBCLADE_C}
 |subcladeC={{clade
    |1=[[Stylotermitidae]]
    |2={{clade
       |1={{clade
          |1=[[Serritermitidae]]
          |2=[[Rhinotermitidae]]
       }}
       |2={{clade
          |1=[[Termitogetonidae]]
          |2={{clade
             |1=[[Psammotermitidae]]
             |2={{clade
                |label1=[[Geoisoptera]]
                |1={SUBCLADE_D}
             }}
          }}
       }}
    }}
 }}
 |targetD={SUBCLADE_D}
 |subcladeD={{clade
    |1=[[Heterotermitidae]]
    |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 [[Stylotermitidae]]: 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 name=":8">{{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>