Gamostolus subantarcticus, from Michigan Science Art.

Belongs within: Heteropteroidea.
Contains: Panheteroptera, Gerromorpha.

The Heteroptera include the so-called 'true bugs', united by the reduction of the tentorium and the presence in nymphs of scent glands on tergites 3 to 7 (Grimaldi & Engel 2005). The name 'Heteroptera' derives from the differentiated wings of many true bugs but this feature is actually characteristic of a derived subgroup of the Heteroptera (the Panheteroptera) whereas other basal lineages retain undifferentiated wings.

Living bugs other than the Enicocephalidae are united as a clade, the Euheteroptera, by mostly reductional synapomorphies including the loss of the proximal crossvein in the fore wing and reduction of the ambient vein, especially posteroapically, and migration of the ventral scent gland openings anteriorly to the thorax. The Enicocephalidae are a primarily tropical and subtropical group of bugs with raptorial fore legs, and the head transversely bilobed with the ocelli on the hind lobe (Carver et al. 1991). Some authors have divided the enicocephalids between distinct families Enicocephalidae and Aenictopecheidae with the former having the pronotum divided by transverse sulci into three distinct laterally constricted areas.

<==Heteroptera [Cimicides, Cimicina, Gymnocerata, Hydrocorisae]
    |--Actinoscytinidae [Actinescytinidae] G75
    |    |--Actinoscytina belmontensis Tillyard 1926 G75, F71
    |    |--Heteroscytina G75
    |    |--Triassodoecus chinai Evans 1963 G75, F71
    |    `--Triscytina G75
    `--+--Euheteroptera CGW91
       |    |--+--Panheteroptera GE05
       |    |  `--+--Ceresopseidae SP02
       |    |     `--Gerromorpha GE05
       |    `--Dipsocoromorpha GE05
       |         |  i. s.: Boreostolus SP02
       |         |--Stemmocryptidae [Stemmocryptoidea] SP02
       |         `--Dipsocoroidea [Trichotelocera] SP02
       |              |--Cuneocoridae GE05
       |              |--Hypsipterygidae SP02
       |              |--Cryptostemma [Dipsocoridae] CGW91
       |              |    `--C. monga CGW91
       |              |--Ceratocombus G75 [Ceratocombidae SP02]
       |              |    |--C. (Xylonannus) australiensis G75
       |              |    `--C. hurdi Wygodzinsky 1959 P92
       |              `--Schizopteridae SP02
       |                   |  i. s.: Dictyonannus flavus CGW91, G75
       |                   |--Macromannus [Hypselosomatinae] CGW91
       |                   |    `--M. webbicus CGW91
       |                   |--Ogeriinae CGW91
       |                   |    |--Pachyplagia australia CGW91, G75
       |                   |    `--Ogeria CGW91
       |                   `--Schizopterinae CGW91
       |                        |--Pachyplagioides reginae CGW91, G75
       |                        |--Pinochius CGW91
       |                        `--Schizoptera RD77
       `--Enicocephalidae [Aenictopecheidae, Enicocephaloidea, Enicocephalomorpha, Henicocephalidae] CGW91
            |  i. s.: Sphigmocephalus MW01
            |         Hoplitocoris YS01
            |         Enicocephalinus acragrimaldii GE05
            |         Disphaerocephalus constrictus GE05
            |         Paenicotechys fossilis GE05
            |         Gamostolus Bergroth 1927 B27
            |           |--*G. subantarcticus [=Henicocephalus subantarcticus] B27
            |           `--G. tonnoiri Bergroth 1927 B27
            |--Monteithostolini [Phthirocorinae] CGW91
            |--Phallopiratinae CGW91
            |--Enicocephalinae GE05
            |    |--Systelloderes CGW91
            |    |--Henschiella CGW91
            |    |--Oncylocotis tasmanicus CGW91
            |    |--Usingeriella CGW91
            |    |--Didymocephalus WEE70
            |    `--Enicocephalus B27
            |         |--E. corticicola Kirkaldy 1908 K08
            |         |--E. fungicola Kirkaldy 1908 K08
            |         |--E. maclachlani B27
            |         `--E. telescopicus K08
            |--Nymphocoris [Nymphocorinae] CGW91
            |    `--N. hilli CGW91
            `--Aenictopecheinae GE05
                 |--Australostolus monteithi CGW91
                 `--Aenictopechys RD77 [incl. Aerorchestes Bergroth 1927 B27]
                      |--A. alluaudi RD77 [=*Aerorchestes alluaudi B27]
                      `--A. necopinatus B27

Heteroptera incertae sedis:
  Malacocoris chlorizans HS01
  Eocimicidae G75
  Probascanion RJ93 [Probascaniidae, Probascanionidae G75]
    `--P. megacephalum RJ93
  Acanthia L02
    |--A. littoralis L02
    |--A. saltatoria G20
    `--A. zosterae L02
  Galgulus Latreille 1802 L02
    `--*G. oculatus [=Naucoris oculata] L02
  Halosalda lateralis RD77
  Macrocephalus cymicoides G20

*Type species of generic name indicated


[B27] Bergroth, E. 1927. Hemiptera Heteroptera from New Zealand. Transactions and Proceedings of the New Zealand Institute 57: 671–684.

[CGW91] Carver, M., G. F. Gross & T. E. Woodward. 1991. Hemiptera (bugs, leafhoppers, cicadas, aphids, scale insects etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 1 pp. 429–509. Melbourne University Press: Carlton (Victoria).

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

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

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

[G75] Gross, G. F. 1975. Plant-feeding and Other Bugs (Hemiptera) of South Australia. Heteroptera—Part I. Handbook of the Flora and Fauna of South Australia.

[HS01] Hossain, M. S., & T. Solhøy. 2001. Densities of Panonychus ulmi (Acari: Tetranychidae) and Aculus schlechtendali (Acari: Eriophyidae) on three apple cultivars in western Norway. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10th International Congress pp. 300–305. CSIRO Publishing: Melbourne.

[K08] Kirkaldy, G. W. 1908. A catalogue of the Hemiptera of Fiji. Proceedings of the Linnean Society of New South Wales 33: 345–391, pl. 4.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[MW01] Molleman, F., & D. E. Walter. 2001. Niche segregation and can-openers: scydmaenid beetles as predators of armoured mites in Australia. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10th International Congress pp. 283–288 Publishing: Melbourne.

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

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[SP02] Shcherbakov, D. E., & Yu. A. Popov. 2002. Superorder Cimicidea Laicharting, 1781: Order Hemiptera Linné, 1758. The bugs, cicadas, plantlice, scale insects, etc. [=Cimicida Laicharting, 1781, =Homoptera Leach, 1815 + Heteroptera Latreille, 1810). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 143–157. Kluwer Academic Publishers: Dordrecht.

[YS01] Yoshizawa, K., & T. Saigusa. 2001. Phylogenetic analysis of paraneopteran orders (Insecta: Neoptera) based on forewing base structure with comments on monophyly of Auchenorrhycha (Hemiptera). Systematic Entomology 26: 1–13.

Last updated: 15 May 2022.

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