Reconstruction of generalised Archescytinidae species, from Grimaldi & Engel (2005).

Belongs within: Paraneoptera.
Contains: Heteropteroidea, Auchenorrhyncha, Protopsyllidiidae, Sternorrhyncha.

The Hemiptera, bugs, are a major group of insects in which the mouthparts have become modified into a tubular rostrum for sucking fluids. The mandibles and maxillae form two pairs of elongate stylets lying in a grooved labium (Grimaldi & Engel 2005). The maxillary and labial palps have been lost. Earlier authors divided the Hemiptera between the suborders Heteroptera, with forewings often divided between a leathery corium and distal membrane, and Homoptera, with undivided wings. The ‘Homoptera’ are now recognised as paraphyletic, and are instead divided between the Sternorrhyncha, Auchenorrhyncha and Coleorrhyncha with the latter two groups forming the Euhemiptera with the Heteroptera.

The earliest hemipterans, the Archescytinidae, are known from the early Permian to the Triassic; members of this group had a long, thin and coiled ovipositor held underneath the abdomen that may have been used to insert eggs into plants (Grimaldi & Engel 2005). Members of more derived groups are known from the Late Permian, indicating that the major lineages of the Sternorrhyncha, Auchenorrhyncha and Heteropteroidea had diverged by that point (Shcherbakov & Popov 2002).

Characters (from Shcherbakov & Popov 2002): Head from hypognathous and nearly immobile to prognathous and freely movable. Antennae moderately long to short, usually of fewer than ten segments. Mouthparts transformed into a sucking proboscis, ensheathed with 1–4-segmented labium and consisting of stylet-like mandibles and maxillae (with bases invaginated into cranium) forming bundle and leaving food and salivary canals medially. Palps lost. Cibarial dilator muscles originating from enlarged postclypeus. Lateral ocelli plesiomorphically close to eyes. Wings homonomous and uncoupled in flight (in Archescytinidae) or heteronomous and coupled at flight by means of (sub)marginal devices, usually with both pairs well developed (sometime subdipterous, (sub)brachypterous with hind wings usually lost, or apterous). Accordingly, flight functionally four-winged, probably anteromotoric in Archescytinidae, functionally (sometimes even morphologically) two-winged, in-phase, anteromotoric in others. Fore wing usually more sclerotised than hind one and fixed on the thorax in repose, with at least its posterior margin inserted into lateral mesoscutellar groove. Hind wing either diminished or with anal area expanded. SC closely associated with or fused to R, its apical portion simulating a branch of the latter. Venation basically simple with two anal veins and two cross-veins, secondarily either prolific or reduced up to simple (SC+)R in fore wings of some coccid males. Hind leg often modified and armed for jumping, or similar to middle one. Tarsi 3–1-segmented. Ovipositor laciniate (with cutting blades), or variously modified, or reduced. Male genitalia variable, always including a penis and usually a pair of parameres. Cerci lost. 10th and 11th abdominal segments forming small anal tube, sometimes modified or reduced. Ovarioles telotrophic. Chromosomes holokinetic. Water-shunting filter chambers of various kinds and symbiotic bacteria in mycetomes often present. Nymphs dorsoventrally flattened, oval, cryptic, unable to jump, secondarily often more adult-like; last instar(s) occasionally transformed into non-feeding sedentary metamorphic (pupal) stage(s).

<==Hemiptera [Arthroidignatha, Cimicida, Homoptera, Rhynchota]
    |--Euhemiptera [Hemelytrata]NR13
    |    |  i. s.: Aviorrhyncha Nel, Bourgoin et al. in Nel, Roques et al. 2013 [Aviorrhynchidae]NR13
    |    |           `--*A. magnifica Nel, Bourgoin et al. in Nel, Roques et al. 2013NR13
    |    |--+--HeteropteroideaSP02
    |    |  `--IngruidaeSP02
    |    |       |--KaltanospesSP02
    |    |       `--Scytoneurella majorRJ93
    |    `--AuchenorrhynchaSP02
    |  `--SternorrhynchaGE05
    `--Archescytinidae [Archescytinina, Archescytinoidea, Palaeohemiptera, Paleorrhyncha]GE05
         |--Permothrips [Permothripidae]SP02
         |--Archescytina permianaSLN15, K-P91
         `--Protopincombea obscura Evans 1943RJ93, F71
Hemiptera incertae sedis:
  Pseudatomoscelis seriatusWS01
  Bagrada cruciferumG01
  Perignerus maidisG01
  Nasonovia ribisnigriA71
  Kikihia subalpinaH05
  Heterotrephes admorsus Esaki & Miyamoto 1959Iw92
  Suisha coreana (Matsumura 1927)Iw92
  Nipponosemia terminalis (Matsumura 1913)Iw92
  Asclepios shiranui (Esaki 1924)Iw92
  ‘Odius’ Stål 1867 non Lilljeborg 1866C92
  Terioaphis trifoliiC81
  Amphisalta zelandicaH97
  Phera Stål 1864K03
  Luederwaldtia Schmidt 1922KO06
  Atopozelus pallensM07
  Poppea evelynaMG06
  Laodelphax striatellusR96
  Cephaloxys Signoret 1847B83
  Anacornutipo lignosaS61
  Metastemma quinquemaculata Lucas 1847E12
  Haematoloecha Stäl 1874D56
  ‘Limonia’ Carvalho 1985 nec Meigen 1803 nec Agassiz 1846KA-Z11
  Liops Fieber 1870KA-Z11
  Umbronia spinosaIm92
  Sagotylus confluentusIm92
    |--G. coccineaBM76
    `--G. versutaE66
  Proceiphilus tesselatusBM76
  Agrioaphis kuricolaBM76
  Macropis viridisBM76
  Coenocorixa expletaWL09
  Laccogrypota grandisBMW13
  Prosapia bicinctaBMW13
    |--Vanua vitiensis Kirkaldy 1906K08
    |--Plestia marginataK08
    `--Euricania tristiculaK08
  Amaurus Burmeister 1835C96
  Naematopus albithorax Boisduval 1835B35
  Archijassus [Archijassidae]RJ93
    `--A. plurinervis Zhang 1985RJ93
  Prosbolecicada [Prosbolecicadidae]RJ93
    `--P. gondwanica Pinto 1987RJ93
    |--H. candelariusR13
    `--H. pyrorrhynchusR13
  Plecophlebus Cockerell 1917P92
    `--*P. nebulosus Cockerell 1917P92
  Praecoris dominicanaP92
  ‘Richteria’ Tode 1966 nec Jones 1874 nec Girault 1920EH19
  Mevania Harold 1874L90
  ‘Coryna’ Wolff 1811 non Bosc 1802BP02

*Type species of generic name indicated


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