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. The Actinoscytinidae are a fossil family known from the Late Permian to Early Jurassic that may represent stem-group Panheteroptera (Gross 1975).

<==Heteroptera [Cimicides, Cimicina, Gymnocerata, Hydrocorisae]
    |--Actinoscytinidae [Actinescytinidae]G75
    |    |--Actinoscytina belmontensis Tillyard 1926G75, F71
    |    |--HeteroscytinaG75
    |    |--Triassodoecus chinai Evans 1963G75, F71
    |    `--TriscytinaG75
       |    |--+--PanheteropteraGE05
       |    |  `--+--CeresopseidaeSP02
       |    |     `--GerromorphaGE05
       |    `--DipsocoromorphaGE05
       |         |  i. s.: BoreostolusSP02
       |         |--Stemmocryptidae [Stemmocryptoidea]SP02
       |         `--Dipsocoroidea [Trichotelocera]SP02
       |              |--CuneocoridaeGE05
       |              |--HypsipterygidaeSP02
       |              |--Cryptostemma [Dipsocoridae]CGW91
       |              |    `--C. mongaCGW91
       |              |--CeratocombusG75 [CeratocombidaeSP02]
       |              |    |--C. (Xylonannus) australiensisG75
       |              |    `--C. hurdi Wygodzinsky 1959P92
       |              `--SchizopteridaeSP02
       |                   |  i. s.: Dictyonannus flavusCGW91, G75
       |                   |--Macromannus [Hypselosomatinae]CGW91
       |                   |    `--M. webbicusCGW91
       |                   |--OgeriinaeCGW91
       |                   |    |--Pachyplagia australiaCGW91, G75
       |                   |    `--OgeriaCGW91
       |                   `--SchizopterinaeCGW91
       |                        |--Pachyplagioides reginaeCGW91, G75
       |                        |--PinochiusCGW91
       |                        `--SchizopteraRD77
       `--Enicocephalidae [Aenictopecheidae, Enicocephaloidea, Enicocephalomorpha, Henicocephalidae]GE05
            |  i. s.: SphigmocephalusMW01
            |         HoplitocorisYS01
            |         Enicocephalinus acragrimaldiiGE05
            |         Disphaerocephalus constrictusGE05
            |         Paenicotechys fossilisGE05
            |         Gamostolus Bergroth 1927B27
            |           |--*G. subantarcticus [=Henicocephalus subantarcticus]B27
            |           `--G. tonnoiri Bergroth 1927B27
            |--Monteithostolini [Phthirocorinae]CGW91
            |    |--SystelloderesCGW91
            |    |--HenschiellaCGW91
            |    |--Oncylocotis tasmanicusCGW91
            |    |--UsingeriellaCGW91
            |    |--DidymocephalusWEE70
            |    `--EnicocephalusB27
            |         |--E. corticicola Kirkaldy 1908K08
            |         |--E. fungicola Kirkaldy 1908K08
            |         |--E. maclachlaniB27
            |         `--E. telescopicusK08
            |--Nymphocoris [Nymphocorinae]CGW91
            |    `--N. hilliCGW91
                 |--Australostolus monteithiCGW91
                 `--AenictopechysRD77 [incl. Aerorchestes Bergroth 1927B27]
                      |--A. alluaudiRD77 [=*Aerorchestes alluaudiB27]
                      `--A. necopinatusB27
Heteroptera incertae sedis:
  Malacocoris chlorizansHS01
  ProbascanionRJ93 [Probascaniidae, ProbascanionidaeG75]
    `--P. megacephalumRJ93
    |--A. littoralisL02
    |--A. saltatoriaG20
    `--A. zosteraeL02
  Galgulus Latreille 1802L02
    `--*G. oculatus [=Naucoris oculata]L02
  Halosalda lateralisRD77
  Macrocephalus cymicoidesG20

*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.

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