Heteroptera

Life-forms

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Paraneoptera
Superordo: Condylognatha
Ordo: Hemiptera
Subordo: Heteroptera
Infraordines (7): Cimicomorpha – Dipsocoromorpha – Enicocephalomorpha – Gerromorpha – Leptopodomorpha – Nepomorpha – Pentatomomorpha

Overview of families (94)

Acanthosomatidae – Aenictopecheidae – Aepophilidae – Alydidae – Anthocoridae – Aphelocheiridae – Aradidae – Artheneidae – Belostomatidae – Berytidae – Blissidae – Canopidae – Ceratocombidae – Cimicidae – Colobathristidae – Coreidae – Corixidae – Cryptorhamphidae – Curaliidae – Cydnidae – Cymidae – Dinidoridae – Dipsocoridae – ?†Ebboidae – Enicocephalidae – Gelastocoridae – Geocoridae – Gerridae – Hebridae – Helotrephidae – Henicocoridae – Hermatobatidae – Heterogastridae – Hydrometridae – Hyocephalidae – Hypsipterygidae – Idiostolidae – †Ignotingidae – Leptopodidae – Lestoniidae – Lygaeidae – Macroveliidae – Malcidae – Medocostidae – Megarididae – Meschiidae – Mesoveliidae – Micronectidae – Microphysidae – Miridae – Nabidae – Naucoridae – Nepidae – Ninidae – Notonectidae – Ochteridae – Omaniidae – Oxycarenidae – Pachygronthidae – †Pachymeridiidae – Pachynomidae – †Palaeoleptidae – Paraphrynoveliidae – Parastrachiidae – Pentatomidae – Phloeidae – Piesmatidae – Plataspididae – Pleidae – Plokiophilidae – Polyctenidae – Potamocoridae – Pyrrhocoridae – Reduviidae – Rhopalidae – Rhyparochromidae – Saldidae – Schizopteridae – Scutelleridae – Stemmocryptidae – Stenocephalidae – Termitaphididae – Tessaratomidae – Thaumastellidae – Thaumastocoridae – Thyreocoridae – Tingidae – †Trisegmentatidae – Urostylidae – Veliidae – Velocipedidae – †Vetanthocoridae – †Yuripopovinidae
Name

Heteroptera Latreille, 1810
References

Henry, T.J. 2009. Biodiversity of Heteroptera. Pp. 223–263 in: Foottit, R.G. & Adler, P.H. (eds.) 2009. Insect biodiversity: science and society. Blackwell Publishing Ltd. ISBN 978-1-4051-5142-9 PDF.
Schuh, R.T. & Slater, J.A. 1995. True bugs of the world (Hemiptera: Heteroptera). Classification and natural history. Ithaca and London: Cornell University Press. ISBN 0-8014-2066-0 Google books
Weirauch, C. & Schuh, R.T. 2011. Systematics and evolution of Heteroptera: 25 years of progress. Annual review of entomology 56: 487–510. DOI: 10.1146/annurev-ento-120709-144833

Additional references

Cassis, G. & Schuh, R.T. 2010. Systematic methods, fossils, and relationships within Heteroptera (Insecta). Cladistics 26(3): 262–280. DOI: 10.1111/j.1096-0031.2009.00283.x
Cordeiro, D.R.S. & Moreira, F.F.F. 2015. New distributional data on aquatic and semiaquatic bugs (Hemiptera: Heteroptera: Gerromorpha & Nepomorpha) from South America. Biodiversity Data Journal 3: e4913. DOI: 10.3897/BDJ.3.e4913 Reference page.
Coscarón, M. del C. 2017. A catalogue of the Heteroptera (Hemiptera) or true bugs of Argentina. Zootaxa 4295(1): 1–432. DOI: 10.11646/zootaxa.4295.1.1. Reference page.
Dursun, A. & Fent, M. 2017. Type Localities of Heteroptera (Insecta: Hemiptera) from Turkey. Zootaxa 4227(4): 451–494. DOI: 10.11646/zootaxa.4227.4.1. Reference page.
Esenbekova, P.A., Nurushev, M.Z. & Homziak, J. 2015. Aquatic Hemiptera (Heteroptera) of Kazakhstan, with notes on life history, ecology and distribution. Zootaxa 4013(2): 195–206. DOI: 10.11646/zootaxa.4013.2.2. Preview (PDF) Reference page.
Faúndez, E.I. 2017. Additions to Idiostolidae in Coscarón (2017) “A catalogue of the Heteroptera (Hemiptera) or true bugs from Argentina”. Zootaxa 4365(1): 99–100. DOI: 10.11646/zootaxa.4365.1.8 Reference page.
Fent, M. et al. 2011. Annotated catalogue of Enicocephalomorpha, Dipsocoromorpha, Nepomorpha, Gerromorpha, and Leptopodomorpha (Hemiptera: Heteroptera) of Turkey, with new records. Zootaxa 2856: 1–84. Preview (PDF).
Ghahari, H. et al. 2013: An annotated catalog of the Iranian Dipsocoromorpha, Enicocephalomorpha, Gerromorpha, Leptopodomorpha and Nepomorpha (Hemiptera: Heteroptera). Zootaxa 3641(4): 301–342. DOI: 10.11646/zootaxa.3641.4.1 Reference page.
Ishikawa, T., Saito, M.U., Kishimoto-Yamda, K., Kato, T., Kurashima, O. & Ito, M. 2015. Inventory of the Heteroptera (Insecta: Hemiptera) in Komaba Campus of the University of Tokyo, a highly urbanized area in Japan. Biodiversity Data Journal 3: e4981. DOI: 10.3897/BDJ.3.e4981. Reference page.
Jung, S-H., Kim, J-G., Oh, S. & Heiss, E. 2015: Type specimens of Heteroptera (Insecta: Hemiptera) collected from North Korea and adjacent regions deposited at Insect Collections of Chungnam National University (CNU) in Daejeon, Republic of Korea. Zootaxa 3981(3): 397–404. DOI: 10.11646/zootaxa.3981.3.5. Preview (PDF) Reference page.
Kanyukova, E.V. & Marusik, Y.M. 2006. A checklist of Heteroptera of the Kuril Islands and brief zoogeographical survey of the fauna. In: Takahashi, H. & Ôhara, M. (eds.) Biodiversity and biogeography of the Kuril Islands and Sakhalin, volume 2. Bulletin of the Hokkaido University Museum (3): 161–174.
Kment, P. & Jindra, Z. 2005. New and interesting records of true bugs (Heteroptera) from Turkey, southeastern Europe, Near and Middle East. Acta Entomologica Musei Nationalis Pragae 45: 3–16. PDF Reference page.
Krüger, A. & Deckert, J. 2016. True bugs (Hemiptera-Heteroptera) of Botswana—Bibliographical inventory and new records. Zootaxa 4114(1): 33–63. DOI: 10.11646/zootaxa.4114.1.2. Reference page.
Lamelas-López, L., Raposeiro, P.M., Borges, P.A.V. & Florencio, M. 2017. Annotated checklist of aquatic beetles (Coleoptera) and true bugs (Heteroptera) in the Azores Islands: new records and corrections of colonization status. Zootaxa 4353(1): 117–132. DOI: 10.11646/zootaxa.4353.1.7. Reference page.
Linnavuori, R.E. 2009. Studies on the Nepomorpha, Gerromorpha, Leptopodomorpha, and Miridae excluding Phylini (Hemiptera: Heteroptera) of Khuzestan and the adjacent provinces of Iran. Acta Entomologica Musei Nationalis Pragae 49: 1–32. Abstract (PDF).
Melo, M.C., Dellapé, P.M., Carpintero, D.L & Montemayor, S.I. 2011. Heteroptera (Hemiptera) from the Chaco National Park (Argentina). Zootaxa 2999: 1–19. Preview (PDF).
Montemayor, S.I.; Dellapé, P.M. 2013: An update of the types of Heteroptera (Hemiptera) housed at the Museo de La Plata Entomological Collection (Argentina). Zootaxa 3599(4): 343–360. DOI: 10.11646/zootaxa.3599.4.3 Reference page.
Pall, J.L. & Coscarón, M. del C. 2016. Contributions to the knowledge of the biodiversity of Heteroptera (Insecta) in the Southern Cone, Argentina. Zootaxa 4170(3): 553–566. DOI: 10.11646/zootaxa.4170.3.8. Reference page.
Rodrigues, J.M.S., Cordeiro, I.D.R.S. & Moreira, F.F.F. 2017. Additions and corrections to the type list of Heteroptera (Insecta: Hemiptera) of the Costa Lima Collection, Oswaldo Cruz Foundation. Zootaxa 4311(2): 183–210. DOI: 10.11646/zootaxa.4311.2.2 Reference page.
Rodrigues, H.D.D.; Ferreira-Keppler, R.L. 2013: Catalog of type specimens of invertebrates in the collection of the Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil. VI. Hexapoda: Hemiptera: Heteroptera. Zootaxa 3716(2): 192–206. DOI: 10.11646/zootaxa.3716.2.4 Reference page.
Weirauch, C. & Cassis, G. 2009. Frena and druckknopf: a synopsis of two fore wing-to-body coupling mechanisms in Heteropterodea (Hemiptera). [[ISSN 1399-560X|Insect Systematics & Evolution 40: 229–252. DOI: 10.1163/187631209X458349
2009: Zootaxa, 2311: 38–48.
Yasunaga, T., Yamada, K., Morakote, R., Taekul, C. & Duangthisan, J. 2016. Transferred depository for twenty-seven holotypes of the plant bug and flower bug species recently described from Thailand (Hemiptera: Heteroptera: Miridae and Anthocoridae). Zootaxa 4107(3): 444–446. DOI: 10.11646/zootaxa.4107.3.13. Reference page.
Ferrari, A., Barão, K.R., Bianchi, F.M., Campos, L.A. & Grazia, J. 2015. Classification and Biogeography of Neotropical True Bugs. In: Panizzi, A.R. & Grazia, J. (eds). True Bugs (Heteroptera) of the Neotropics. Entomology in Focus, vol 2. Springer, Dordrecht. Vol. 2: 57–87. ResearchGate Open access. DOI: 10.1007/978-94-017-9861-7_3Reference page.

Vernacular names
Deutsch: Wanzen
English: True bug
español: Heterópteros
suomi: Luteet
Nordfriisk: Wochlüs
français: Hétéroptères
magyar: Poloskák
日本語: カメムシ亜目 (異翅亜目)
한국어: 노린재류
Nederlands: Wantsen
polski: pluskwiaki różnoskrzydłe
português: Heterópteros/Percevejos
slovenščina: Stenice
Türkçe: Yarım kanatlılar
中文: 異翅亞目

The Heteroptera are a group of about 40,000 species of insects in the order Hemiptera. They are sometimes called "true bugs",[1] though that name more commonly refers to the Hemiptera as a whole. "Typical bugs" might be used as a more unequivocal alternative, since the heteropterans are most consistently and universally termed "bugs" among the Hemiptera. "Heteroptera" is Greek for "different wings": most species have forewings with both membranous and hardened portions (called hemelytra); members of the primitive sub-group Enicocephalomorpha have completely membranous wings.

The name "Heteroptera" is used in two very different ways in modern classifications. In Linnean nomenclature, it commonly appears as a suborder within the order Hemiptera, where it can be paraphyletic or monophyletic depending on its delimitation. In phylogenetic nomenclature, it is used as an unranked clade within the Prosorrhyncha clade, which in turn is in the Hemiptera clade. This results from the realization that the Coleorrhyncha are just "living fossil" relatives of the traditional Heteroptera, close enough to them to be united with that group.

The infraorders Leptopodomorpha, Gerromorpha, and Nepomorpha, comprise a significant component of the world’s aquatic and semiaquatic insects. There are 23 families, 343 genera and 4,810 species group taxa within these three infraorders.[2] Most of the remaining groups that are common and familiar are in the Cimicomorpha and Pentatomomorpha.

Classification
Anatomy of the dorsal aspect of a shield bug. A: head; B: thorax; C: abdomen. 1: claws; 2: tarsus; 3: tibia; 4: femur; 8: compound eye; 9: antenna; 10: clypeus; 23: laterotergites (connexivum); 25: pronotum; 26: scutellum; 27: clavus; 28: corium; 29: embolium; 30: hemelytral membrane.

The use of the name "Heteroptera" has had the rank of order, dating back to 1810 by Pierre André Latreille. Only recently has it been relegated to a subsidiary rank within a larger definition of Hemiptera, so many reference works still include it as an order. Whether to continue treating it as a suborder is still a subject of some controversy, as is whether the name itself should ever be used, although three basic approaches ranging from abolishing it entirely to maintaining the taxonomy with a slight change in systematics is proposed, two of which (but not the traditional one) agree with the phylogeny. The competing classifications call for a preference for two suborders versus one when the "living fossil" family Peloridiidae is taken into consideration:

In one revised classification proposed in 1995,[3] the name of the suborder is Prosorrhyncha, and Heteroptera is a rankless subgroup within it. The only difference between Heteroptera and Prosorrhyncha is that the latter includes the family Peloridiidae, which is a tiny relictual group that is in its own monotypic superfamily and infraorder. In other words, the Heteroptera and Prosorrhyncha sensu Sorensen et al. are identical except that Prosorrhyncha contains one additional infraorder, called Peloridiomorpha (comprising only 13 small genera). The ongoing conflict between traditional, Linnaean classifications and nontraditional classifications is exemplified by the problem inherent in continued usage of the name Heteroptera when it no longer can be matched to any standard Linnaean rank (as it falls below suborder but above infraorder). If this classification succeeds, then the "Heteroptera" grouping may be discarded, but in that case it is likely that no ranks will be used at all according to the standards of phylogenetic nomenclature.

In the traditional classification,[4] the Peloridiidae are retained as their own suborder, called Coleorrhyncha; "Heteroptera" is treated the same. Functionally, the only difference between this classification and the preceding is that the former uses the name Prosorrhyncha to refer to a particular clade, while the traditional approach divides this into the paraphyletic Heteroptera and the monophyletic Coleorrhyncha. Many believe it is preferable to use only one name because the characteristics of the two traditional suborders are too closely related to be treated as separate.

Alternatively,[5] the modified approach of placing Coleorrhyncha 'within' the Heteroptera can be used. Indeed, as that solution preserves the well-known Heteroptera at the taxonomic rank they traditionally hold while making them a good monophyletic group, it seems preferable to the paraphyletic "Heteroptera" used in older works. In that case, the "core" Heteroptera could be considered a section – as yet unnamed, mainly because the Prosorrhyncha were proposed earlier – within the "expanded" Heteroptera, or the latter could simply be described as consisting of a basal "living fossil" lineage and a more apomorphic main radiation. Whether the name "Coleorrhyncha" is to be retained for the basal lineage or whether the more consistent "Peloridiomorpha" is used instead is a matter of taste, as described below.

Separate from the question of the actual "closeness" of Heteroptera and Coleorrhyncha is the potential disruption to traditional construction of names; there seems to be reluctance among hemipterists to abandon the use of "Heteroptera". This can be seen by the name itself, as it is a violation of convention to use the ending "-ptera" for any rank above genus other than an order – though since it is a convention rather than a mandatory rule of Linnean nomenclature, taxonomists are technically free to violate it (which is why, for example, not all insect orders end in "-ptera", e.g., Odonata). However, in most cases when such conventions are violated, it does not create an internal conflict as in the present case (that is, the order Hemiptera has a suborder named Heteroptera, which is an internal conflict). At least some hemipterists argue that the name Heteroptera should be dropped entirely to eliminate this internal conflict, though the third possibility offers a workaround. In that case, to achieve full consistency of names "Coleorrhyncha" would probably be dropped in favor of "Peloridiomorpha".
References

Tree of Life Web Project (2005): Heteroptera. True bugs. Version of January 1, 2005. Retrieved July 28, 2008.
Polhemus, John T.; Polhemus, Dan A. (2008). "Global diversity of true bugs (Heteroptera; Insecta) in freshwater". Hydrobiologia. 595 (5): 379–91. doi:10.1007/s10750-007-9033-1. S2CID 45657091.
Sorensen, J. T., B. C. Campbell, R. J. Gill & J. D. Steffen-Campbell (1995): Non-monophyly of Auchenorrhyncha ("Homoptera"), based upon 18S rDNA phylogeny: eco-evolutionary and cladistic implications with pre-Heteropteroidea Hemiptera (s.l.) and a proposal for new monophyletic suborders. Pan-Pacific Entomologist 71 (1): 31–60.
Maddison, David R. (1995): Tree of Life Web Project – Hemiptera. True bugs, cicadas, leafhoppers, aphids, etc.. Version of January 1, 2005. Retrieved July 28, 2008.

Cassis, Gerasimos & Gross, Gordon (1995): Australian Biological Resources Study – Hemiptera: Heteroptera (Coleorrhyncha to Cimicomorpha). Gerrids, Reduviids, Water-striders. Version of June 30, 1995. Retrieved July 28, 2008.

Further reading
China, W.E.; Miller, N.C.E. (1959). "Check-list and keys to the families and subfamilies of the Hemiptera-Heteroptera". Bulletin of the British Museum (Natural History), Entomology. London: British Museum. 8 (1): 1–45.