Panhexapoda

Tricholepidion gertschi, copyright Samuel DeGrey.

Belongs within: Allotriocarida.
Contains: Protura, Collembola, Diplura, Archaeognatha, Zygentoma, Pterygota.

The Panhexapoda include the modern Hexapoda (insects and related taxa) together with a small number of Devonian relatives. Members of this clade are united by the presence of only a single pair of antennae, and reduction of appendages posterior to the anterior three pairs of walking legs. In crown-group hexapods, the posterior appendages have been more or less lost though some basal lineages retain small remnants that do not function as walking legs (Grimaldi & Engel 2005). Functional post-thoracic legs remain present in the Devonian marine fossils Devonohexapodus, Wingertshellicus and Cambronatus. Leverhulmia mariae is a fragmentary fossil from the earlier Devonian that preserves at least five pairs of legs but which resembles modern Insecta in the possession of paired lateral claws and a fused median claw on the leg tarsi (Fayers & Trewin 2005). Modern hexapods are divided between four basal clades, the Protura, Collembola, Diplura and Insecta. The Insecta differ from other living hexapods in that the mandibles are not recessed within a pouch on the head capsule (entognathous); other synapomorphies of the insects include loss of musculature in the antenna beyond the scape, loss of articulations between the leg coxae and sternum, and division of the tarsi into tarsomeres (Grimaldi & Engel 2005). Authors have disagreed on whether the remaining entognathous hexapods form a single clade that is sister to the insects or whether they form a series of successive sister groups. Diplura may be united with Insecta by the presence of ancestrally filiform cerci (Kukalová-Peck 1991).

One of the most speciose of all groups of organisms, the insects dominate the terrestrial environment. The majority of insects belong to the winged clade Pterygota, but some basal clades retain the primitive wingless morphology. These latter groups, the Archaeognatha and Zygentoma, are superficially similar in having a more or less elongate body form with three long terminal filaments (the cerci and median filament), and so have been combined in the past under the name Thysanura. However, the Zygentoma share a number of synapomorphies with the Pterygota, including the presence of mandibles with a secondary articulation in addition to the single articulation found in Archaeognatha, and are now recognised a forming a clade with the winged insects (Grimaldi & Engel 2005). The northern Californian Tricholepidion gertschi is a silverfish-like species retaining a number of plesiomorphic features such as ocelli, large abdominal sterna, abdominal styli and eversible vesicles, and five-segmented tarsi (Grimaldi & Engel 2005), and some analyses have suggested it may represent a relict sister group to Zygentoma and Pterygota.

Panhexapoda
    |--Devonohexapodus bocksbergensis Haas, Waloszek & Haretenberger 2003 GE05, FT05
    |--+--Wingertshellicus backsei Briggs & Bartels 2001 GE05, FT05
    |  `--Cambronatus brasseli Briggs & Bartels 2001 GE05, FT05
    `--Hexapoda [Apterodicera, Apterygota, Ellipura, Entognatha, Entotropha, Nonoculata, Parainsecta, Thysanura] GE05
         |  i. s.: Leverhulmia Anderson & Trewin 2003 FT05
         |           `--*L. mariae Anderson & Trewin 2003 FT05
         |         Praemachilis [Praemachilidae] P92
         |--Protura LSE13
         `--+--Collembola LSE13
            `--+--Diplura LSE13
               `--Insecta [Ectognatha, Ectotropha, Lepismatona, Lepismenae, Scarabaeoda, Thysanuroidea, Triplura] LSE13
                    |  i. s.: Drosicha mangiferae G01
                    |         Neocolochelyna itoi Takeuchi 1951 I92
                    |         Paratlanticus tsushimensis Yamasaki 1986 I92
                    |         Cercomantispa shirozui Nakahara 1961 I92
                    |         Armatocillenus sumaoi (Morita 1981) I92
                    |         Heplochlaenius insularis Uéno 1964 I92
                    |         Ogasawarazo daitoensis Voss 1971 I92
                    |         Xyelecia japonica Tagashi 1972 I92
                    |         Pleroneura hikosana Togashi 1972 I92
                    |         Nipponorhynchus I92
                    |           |--N. bimaculatus Naito 1973 I92
                    |           `--N. mirabilis Takeuchi 1941 I92
                    |         Caenosclerogibba japonica Yasumatsu 1958 I92
                    |         Heliophobus texturatus (Alpheraky 1892) I92
                    |         Eldana saccharina H91
                    |         Tricopria drosophilae GJR97
                    |         Aramigus tesselatus GJR97
                    |         Leptophyllus Hope 1842 FT93
                    |         Amaurocoris laticeps G89
                    |         Aspongopus viduatus G89
                    |         Nariscus spinosus G89
                    |         Ischnopeza hirticornis G89
                    |         Tephrocerus gracilis G17
                    |         Eudamus proteus G18
                    |         Melitara junctolineella B88
                    |         Proceras sacchariphagum B88
                    |         Stomopteryx simplexella B88
                    |         Diplosis sorghicola B88
                    |         Adelidium cordatum Tillyard 1917 F71
                    |         Agetochoristella adscita Riek 1952 F71
                    |         Anomaloscytina F71
                    |           |--A. incompleta Evans 1943 F71
                    |           `--A. metapteryx Davis 1942 F71
                    |         Aphryganoneura anomala Tillyard 1926 F71
                    |         Archaeosmylus pectinatus Riek 1952 F71
                    |         Atelophlebis culleni (Etheridge & Olliff 1890) [=Ephemera culleni] F71
                    |         Austroprosbole maculata Evans 1943 F71
                    |         Austroscytina imperfecta Evans 1943 F71
                    |         Beaconiella F71
                    |           |--B. fennahi Evans 1963 F71
                    |           `--B. multivenata Evans 1963 F71
                    |         Belpsylla reticulata Evans 1943 F71
                    |         Elateridium angustius Tillyard 1917 F71
                    |         Eochiliocycla angusta Davis 1942 F71
                    |         Eupincombea postiva Davis 1942 F71
                    |         Homaloscytina plana Tillyard 1926 F71
                    |         Mesonotoperla sinuata Riek 1954 F71
                    |         Mesothyris westraliensis Riek 1968 F71
                    |         Metrorhynchites grandis Tillyard 1917 F71
                    |         Mitchelloneura permiana Tillyard 1921 F71
                    |         Neoageta elongata Riek 1953 F71
                    |         Neopetromantis australis Riek 1953 F71
                    |         Notoblattites F71
                    |           |--N. mitchelli Tillyard 1917 F71
                    |           `--N. wianamattensis Tillyard 1917 F71
                    |         Orthoscytina F71
                    |           |--O. belmontensis Tillyard 1926 F71
                    |           |--O. indistincta Tillyard 1926 F71
                    |           |--O. mitchelli Tillyard 1926 F71
                    |           |--O. obliqua Tillyard 1926 F71
                    |           |--O. pincombei Tillyard 1926 F71
                    |           |--O. quinquemedia Tillyard 1926 F71
                    |           |--O. subcostalis Tillyard 1926 F71
                    |           `--O. tetraneura Tillyard 1926 F71
                    |         Palaeovicia incerta Evans 1943 F71
                    |         Permagra distincta Evans 1943 F71
                    |         Permoberothella perplexa Riek 1953 F71
                    |         Permobrachus F71
                    |           |--P. dubia (Tillyard 1926) [=Permodiphthera dubia] F71
                    |           `--P. magnus Evans 1943 F71
                    |         Permocapitus globulus Evans 1943 F71
                    |         Permocephalus knighti Evans 1943 F71
                    |         Permodiphthera robusta Tillyard 1926 F71
                    |         Permojassus F71
                    |           |--P. australis Tillyard 1926 F71
                    |           `--P. dubius Tillyard 1926 F71
                    |         Permopsyllidops stanleyi Davis 1942 F71
                    |         Permopsylloides insolita Evans 1943 F71
                    |         Permorapisma biserialis Tillyard 1926 [incl. P. triserialis Tillyard 1926] F71
                    |         Permoscarta F71
                    |           |--P. mitchelli Tillyard 1917 F71
                    |           `--P. triventulata Tillyard 1926 F71
                    |         Permothea latipennis Tillyard 1926 F71
                    |         Permotheela scytinopteroides Davis 1942 F71
                    |         Permovicia obscura Evans 1943 F71
                    |         Phipoides elegans Riek 1953 F71
                    |         Protopsyllops minuta Evans 1943 F71
                    |         Psocopsyllidium media Davis 1942 F71
                    |         Psocoscytina bifida Davis 1942 F71
                    |         Psyllidiana davisia Evans 1943 F71
                    |         Stanleyana pulchra Evans 1943 F71
                    |         Stenoglyphis kimblensis Evans 1947 F71
                    |         Stenoscytina australiensis Tillyard 1926 F71
                    |         Triassophlebis stigmata Tillyard 1922 F71
                    |         Triassopsylla plecioides Tillyard 1917 F71
                    |         Triassocytinopsis paranotalis Evans 1956 F71
                    |         Tripsyllidium wadei Evans 1956 F71
                    |         Tychticoloides belmontensis Evans 1963 F71
                    |         Paleacrita vernata Z91
                    |         Itame argillacearia DC10
                    |         Biloba subsecivela VHK02
                    |         Lygropia quaternalis VHK02
                    |         Marasma suspicalis VHK02
                    |         Noorda moringa VHK02
                    |         Nephanteryx rhodobasalis VHK02
                    |         Anomalococcus indicus VHK02
                    |         Phidodonta modesta VHK02
                    |         Bissetia steniella VHK02
                    |         ‘Cheyletus’ nigripes Mola 1907 B49
                    |         Stephanostoma Danielsen 1880 D56
                    |         Phrynomorphus Curtis 1831 FG06
                    |         Catulus Kniphof 1759 C84
                    |         Fur Jones 1940 C84
                    |         Pterolamia Breuning 1942 C84
                    |         Griphologus lowei K-P91
                    |         Aethiocarenus Poinar & Brown 2017 [Aethiocarenidae, Aethiocarenodea] PB17
                    |           `--*A. burmanicus Poinar & Brown 2017 PB17
                    |         Ammoconia caecimacula F92
                    |         Brephos parthenias F92
                    |         Calocampa exoleta F92
                    |         Calymnia trapezina F92
                    |         Dianthoecia F92
                    |           |--D. capsincola F92
                    |           |--D. carpophaga F92
                    |           `--D. cucubali F92
                    |         Himera pennaria F92
                    |         Lygris poputata F92
                    |         Parasemia plantaginis F92
                    |         Phragmasobis fuliginosa F92
                    |         Porthetria dispar F92
                    |         Theria rubricapraria F92
                    |         Carcharodus alceae F92
                    |         Ochrostigma velitaris F92
                    |         Arsilonche alhovenosa F92
                    |         Brotolomia meticulosa F92
                    |         Hydroecia micacea F92
                    |         Triphosa dubitata F92
                    |         Hipocrita jacobaeae F92
                    |         Trichiocampus viminalis F92
                    |         Jaspidea celsia F92
                    |         Plecoptera reflexa F92
                    |         Perinephela lancealis F92
                    |         Diacerisia obliqua F92
                    |         Pterostoma palpina F92
                    |         Drepama harpogula F92
                    |         Bhima idiota F92
                    |         Ortholintha cervinata F92
                    |         Amphidasis betularia F92
                    |         Rhodophaea suavella F92
                    |         Psammotis hyalinalis F92
                    |         Polychrosis botrana F92
                    |         Rhopobota naevana F92
                    |         Notocelia uddmanniana F92
                    |         Sophronia humerella F92
                    |         Phalacropteryx praecellens F92
                    |         Plastenia retusa F92
                    |         Locastra muscosalis F92
                    |         Erocampa ovata F92
                    |         Rhyparia purpurata F92
                    |         Pyrrhidium sanguineum F92
                    |         Tephroclystia innotata F92
                    |         Thamnonoma wauaria F92
                    |         Epineuronia cespitis F92
                    |         Cyanis argiolus F92
                    |         Dasystoma salicella F92
                    |         Demas coryli F92
                    |         Alconeura Ball & DeLong 1925 DC37
                    |         Hyloidea McAtee 1926 DC37
                    |         Dikraneuroidea Lawson 1929 DC37
                    |         Forcipata DeLong & Caldwell 1936 DC37
                    |         Halidisota argentata M38
                    |         Euschausia ingens M38
                    |         Osprinchotus K18
                    |           |--O. capensis K18
                    |           `--O. (Xenodocon) seductor K18
                    |         ‘Gervaisia’ Robineau-Desvoidy 1863 nec Bonaparte 1854 nec Waga 1858 OF80
                    |         Cathilaria rigida G05
                    |         Doryscelis calcarata B28
                    |         Scopelosoma satellitia R13
                    |         Agrypnetes crassicornis R13
                    |         Agrypnia pagetana R13
                    |         Cheimatobia boreata R13
                    |         Sciaphila cellaria R13
                    |         Eugaster guyoni R13
                    |         Gloveria psidii R13
                    |         Jatrophobia brasiliensis M99
                    |         Ascidium Tode 1782 (n. d.) KC01
                    |         Caulogaster Corda 1831 (n. d.) KC01
                    |         Crateromyces Corda 1831 (n. d.) KC01
                    |         ‘Uralia’ Martynova 1976 nec Mulsant & Verreaux 1866 nec Likharev 1925 EH19
                    |--Archaeognatha TW05
                    `--Dicondylia [Cercofilata] GE05
                         |--Tricholepidion gertschi Wygodzinsky 1961 TW05, KZ02
                         `--+--Zygentoma TW05
                            `--Pterygota TW05

*Type species of generic name indicated

REFERENCES

[B49] Baker, E. W. 1949. A review of the mites of the family Cheyletidae in the United States National Museum. Proceedings of the United States National Museum 99 (3238): 267–320.

[B28] Betrem, J. G. 1928. Monographie der Indo-Australischen Scoliiden mit zoogeographischen Betrachtungen. H. Veenman & Zonen: Wageningen.

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[C84] Compagno, L. J. V. 1984. FAO Species Catalogue vol. 4. Sharks of the World: An annotated and illustrated catalogue of shark species known to date. Part 2—Carcharhiniformes. United Nations Development Programme, Food and Agriculture Organization of the United Nations: Rome.

[D56] Dawes, B. 1956. The Trematoda with special reference to British and other European forms. University Press: Cambridge.

[DC37] DeLong, D. M., & J. S. Caldwell. 1937. The genus Dikraneura—a study of the male genitalia—with descriptions of new species. Proceedings of the Entomological Society of Washington 39 (2): 17–34.

[DC10] Drummond, F. A., J. A. Collins, B. Choate, D. Woodman, D. T. Jennings, H. Y. Forsythe, Jr & J. C. Cokendolpher. 2010. Harvestman (Opiliones) fauna associated with Maine lowbush blueberry fields in the major production areas of Washington and Hancock counties. Environmental Entomology 39 (5): 1428–1440.

[EH19] Énay, R., & M. K. Howarth. 2019. Part L, revised, volume 3B, chapter 7: Systematic descriptions of the Perisphinctoidea. Treatise Online 120: 1–184.

[F92] Fan Z. 1992. Key to the Common Flies of China 2^nd ed. Science Press: Beijing.

[FT05] Fayers, S. R., & N. H. Trewin. 2005. A hexapod from the Early Devonian Windyfield Chert, Rhynie, Scotland. Palaeontology 48 (5): 1117–1130.

[FT93] Fensome, R. A., F. J. R. Taylor, G. Norris, W. A. S. Sarjeant, D. I. Wharton & G. L. Williams. 1993. A classification of living and fossil dinoflagellates. Micropaleontology Special Publication 7: i–viii, 1–351.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[FG06] Frost, D. R., T. Grant, J. Faivovich, R. H. Bain, A. Haas, C. F. B. Haddad, R. O. de Sá, A. Channing, M. Wilkinson, S. C. Donnellan, C. J. Raxworthy, J. A. Campbell, B. L. Blotto, P. Moler, R. C. Drewes, R. A. Nussbaum, J. D. Lynch, D. M. Green & W. C. Wheeler. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History 297: 1–370.

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso “Esploratore” dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 5–72.

[GJR97] Giordano, R., J. J. Jackson & H. M. Robertson. 1997. The role of Wolbachia bacteria in reproductive incompatibilities and hybrid zones of Diabrotica beetles and Gryllus crickets. Proceedings of the National Academy of Sciences of the USA 94: 11439–11444.

[G17] Girault, A. A. 1917. Descriptiones stellarum novarum. Privately published (reprinted: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 80–101).

[G18] Girault, A. A. 1918. North American Hymenoptera Trichogrammatidae. Privately published (reprinted: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 142–152).

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[G05] Grissell, E. E. 2005. A review of North American species of Microdontomerus Crawford (Torymidae: Hymenoptera). Journal of Hymenoptera Research 14 (1): 22–65.

[G01] Gupta, S. K. 2001. A conspectus of natural enemies of phytophagous mites and mites as potential biocontrol agents of agricultural pests in India. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10^th International Congress pp. 484–497. CSIRO Publishing: Melbourne.

[H91] Hokkanen, H. M. T. 1991. Trap cropping in pest management. Annual Review of Entomology 36: 119–138.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[KC01] Kirk, P. M., P. F. Cannon, J. C. David & J. A. Stalpers. 2001. Ainsworth & Bisby's Dictionary of the Fungi 9^th ed. CAB International: Wallingford (UK).

[KZ02] Klass, K.-D., O. Zompro, N. P. Kristensen & J. Adis. 2002. Mantophasmatodea: A new insect order with extant members in the Afrotropics. Science 296: 1456–1459.

[K18] Kohl, F. F. 1918. Die Hautflügergruppe "Sphecinae". IV. Teil. Die natürliche Gattung Sceliphron Klug (Pelopoeus Latr.). Annalen des Naturhistorischen Museum in Wien 32: 1–171.

[K-P91] Kukalová-Peck, J. 1991. Fossil history and the evolution of hexapod structures. In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 141–179. Melbourne University Press: Carlton (Victoria).

[LSE13] Legg, D. A., M. D. Sutton & G. D. Edgecombe. 2013. Arthropod fossil data increase congruence of morphological and molecular phylogenies. Nature Communications 4: 2485.

[M99] Moraes, G. J. de. 1999. Pest status of the cassava green mite in Brazil and strategies for its control. In: Needham, G. R., R. Mitchell, D. J. Horn & W. C. Welbourn (eds) Acarology IX vol. 2. Symposia pp. 287–291. Ohio Biological Survey: Columbus (Ohio).

[M38] Muesebeck, C. F. W. 1938. New species and synonymy in the genus Macrocentrus (Hymenoptera: Braconidae). Proceedings of the Entomological Society of Washington 40 (6): 170–172.

[OF80] Olson, S. L., & A. Feduccia. 1980. Relationships and evolution of flamingos (Aves: Phoenicopteridae). Smithsonian Contributions to Zoology 316: 1–73.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[PB17] Poinar, G., Jr & A. E. Brown. 2017. An exotic insect Aethiocarenus burmanicus gen. et sp. nov. (Aethiocarenodea ord. nov., Aethiocarenidae fam. nov.) from mid-Cretaceous Myanmar amber. Cretaceous Research 72: 100–104.

[R13] Reuter, O. M. 1913. Lebensgewohnheiten und Instinkte der Insekten bis zum Erwachen der sozialen Instinkte. R. Friedländer & Sohn: Berlin.

[TW05] Terry, M. D., & M. F. Whiting. 2005. Mantophasmatodea and phylogeny of the lower neopterous insects. Cladistics 21: 240–257.

[VHK02] Verma, M., M. Hayat & S. I. Kazmi. 2002. The species of Elasmus from India (Hymenoptera: Chalcidoidea: Eulophidae). Oriental Insects 36: 245–306.

[Z91] Zhang, Z. Q. 1991. Biology of mites of Allothrombiinae (Acari: Trombidiidae) and their potential role in pest control. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 513–520. SPB Academic Publishing: The Hague.

Last updated: 3 June 2022.