Big-eyed toad bug Gelastocoris sp., copyright Gail Hampshire.

Belongs within: Heteroptera.
Contains: Nepoidea, Corixidae, Naucoroidea, Notonectoidea, Leptopodomorpha, Cimicomorpha, AradoideaPentatomoidea, Coreoidea, LygaeidaeRhyparochromidae.

The Panheteroptera are a major clade of true bugs characterised by the division of the forewings between a sclerotised proximal corium and non-sclerotised distal membrane. Panheteropterans can be divided between three major clades, the mostly water-associated Nepomorpha and Leptopodomorpha and the more strongly terrestrial Geocorisae. The Geocorisae are in turn divided between the Cimicomorpha and Pentatomomorpha, the latter group lacking a costal fracture in the fore wing and having the R and M veins in the hind wing distally separated with the subcostal vein poorly developed (Carver et al. 1991). Pentatomomorphs with the exception of the Aradoidea are united in a clade Trichophora by the presence of more than two trichobothria on either side of the midline of abdominal sterna 3 to 7 (Yao et al. 2012).

The Nepomorpha are one of the few groups of insects including species that are aquatic through their entire life cycle. Members of the Ochteroidea are found mostly along margins of water bodies and differ from other nepomorphs in possessing ocelli. The Gelastocoridae, toad bugs, have raptorial fore legs with a broadened femur (Carver et al. 1991).

    |--Nepomorpha [Cryptocerata, Hydrocorisiae]GE05
    |    |--NepoideaGE05
    |    `--+--+--CorixidaeGE05
    |       |  `--+--NaucoroideaSP02
    |       |     `--+--ApnopidaeG75
    |       |        `--NotonectoideaGE05
    |       `--Ochteroidea [Gelastocoroidea]SP02
    |            |  i. s.: HeterochterusSP02
    |            |--Propreocoris [Propreocoridae, Propreocorinae]GE05
    |            `--+--Ochteridae [Pelogonidae]GE05
    |               |    |--Megochterus nasutusCGW91
    |               |    `--OchterusYS01
    |               |         |--O. australicusCGW91
    |               |         |--O. eurythoraxPH10
    |               |         `--O. marginatusWEE70 [=Pelogonus marginatusG20]
    |               `--Gelastocoridae [Mononychidae]GE05
    |                    |--Nerthra [Nerthrinae]T61b
    |                    |    |--N. asiatica (Horvarth 1892) [=Mononyx asiaticus; incl. M. grossus Montandon 1899]T61b
    |                    |    |--N. indica (Atkinson 1888) (see below for synonymy)T61b
    |                    |    |--N. luteovariaPH10
    |                    |    |--N. nudataCGW91
    |                    |    |--N. plautaCGW91
    |                    |    `--N. tuberculataT61a
    |                    `--Gelastocoris [incl. Montandonius Melin 1929; Gelastocorinae]T61a
    |                         |--G. angulatus [=Montandonius angulatus; incl. M. bridarollii, M. mansosotoi, M. willineri]T61a
    |                         |--G. flavusT61a
    |                         |--G. fuscus [incl. G. martinezi]T61a
    |                         |--G. majorT61a
    |                         |--G. nebulosus (see below for synonymy)T61a
    |                         |--G. quadrimaculatusT61a
    |                         `--G. vicinusT61a
       `--Geocorisae [Corisiae]GE05
                 |  i. s.: ProtocoridaeYR12
                 |           |--Protocoris indistinctusYR12
                 |           `--Pallicoris firmis Lin 1986YR12, RJ93
                 |         MesopentacoridaeYR12
                 |           |--Corienta transbaicalica Popov 1990RJ93
                 |           |--Pauropentacoris macrurataYR12
                 |           `--MesopentacorisGE05
                 |                |--M. costalisYR12
                 |                `--M. orientalisYR12
                      |    |--Venicoris Yao, Ren & Rider in Yao, Ren et al. 2012YR12
                      |    |    `--*V. solaris Yao, Ren & Rider in Yao, Ren et al. 2012YR12
                      |    `--Clavaticoris Yao, Ren & Cai in Yao, Ren et al. 2012YR12
                      |         `--*C. zhengi Yao, Ren & Cai in Yao, Ren et al. 2012YR12
                      `--+--Piesmatidae [Piesmatoidea, Piesmidae]SP02
                         |    |--Mcateella interiorisCGW91, G75
                         |    |--MiespaCGW91
                         |    |--Cretopisma suukyiaeYR12
                         |    `--PiesmaYR12
                         |         |--P. capitataH01
                         |         |--P. quadrataYR12
                         |         `--P. quadricornis [=Cantacader quadricornis]L49
                            |  `--CoreoideaYR12
                                 `--Artheneidae [Artheneinae]CGW91
                                      |--Nothochromus maoricus Slater, Woodward & Sweet 1962L02
                                      `--Dilompus robustusCGW91

Gelastocoris nebulosus [incl. G. bergi De Carlo 1954, G. bolivianus De Carlo 1954, G. monrosi De Carlo 1959, G. paraguayensis De Carlo 1954, G. vianai De Carlo 1954]T61a

Nerthra indica (Atkinson 1888) [=Mononyx indicus; incl. M. projectus Distant 1906, M. turgidulus Distant 1906, Nerthra turgidula]T61b

*Type species of generic name indicated


[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).

[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.

[H01] Horváth, G. 1901. Hemipterák [Hemiptera]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 245–274. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[L02] Larivière, M.-C. 2002. Primary types of Heteroptera in the New Zealand Arthropod Collection: Pentatomomorpha described from New Zealand before the year 2000. New Zealand Entomologist 25: 79–85.

[L49] Lucas, H. 1849. Exploration Scientifique de l’Algérie pendant les années 1840, 1841, 1842 publiée par ordre du gouvernement et avec le concours d’une commission académique. Sciences physiques. Zoologie. II. Histoire naturelle des animaux articulés. Troisième partie. Insectes (suite). Imprimerie Nationale: Paris.

[PH10] Pinder, A. M., S. A. Halse, R. J. Shiel & J. M. McRae. 2010. An arid zone awash with diversity: patterns in the distribution of aquatic invertebrates in the Pilbara region of Western Australia. Records of the Western Australian Museum Supplement 78 (1): 205–246.

[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.

[T61a] Todd, E. L. 1961a. Synonymical notes on some South American species of Gelastocoris Kirkaldy (Hemiptera: Gelastocoridae). Proceedings of the Biological Society of Washington 74: 57–64.

[T61b] Todd, E. L. 1961b. Notes on some toad bugs (Hemiptera: Gelastocoridae) from India. Proceedings of the Biological Society of Washington 74: 93–94.

[WEE70] Woodward, T. E., J. W. Evans & V. F. Eastop. 1970. Hemiptera (bugs, leafhoppers, etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 387–457. Melbourne University Press.

[YR12] Yao, Y., D. Ren, D. A. Rider & W. Cai. 2012. Phylogeny of the infraorder Pentatomomorpha based on fossil and extant morphology, with description of a new fossil family from China. PLoS One 7 (5): e37289.

[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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