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). 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] GE05
    |--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. 2013 NR13
    |    |--+--Heteropteroidea SP02
    |    |  `--Ingruidae SP02
    |    |       |--Kaltanospes SP02
    |    |       `--Scytoneurella major RJ93
    |    `--Auchenorrhyncha SP02
    |--+--Protopsyllidiidae GE05
    |  `--Sternorrhyncha GE05
    `--Archescytinidae [Archescytinina, Archescytinoidea, Palaeohemiptera, Paleorrhyncha] GE05
         |--Permothrips [Permothripidae] SP02
         |--Maueria SP02
         |--Permoscytina SP02
         |--Archescytina permiana SLN15, K-P91
         `--Protopincombea obscura Evans 1943 RJ93, F71

Hemiptera incertae sedis:
  Mesanthocoris ZBH03
  Pseudatomoscelis seriatus WS01
  Bagrada cruciferum G01
  Perignerus maidis G01
  Nasonovia ribisnigri A71
  Kikihia subalpina H05
  Heterotrephes admorsus Esaki & Miyamoto 1959 Iw92
  Suisha coreana (Matsumura 1927) Iw92
  Nipponosemia terminalis (Matsumura 1913) Iw92
  Asclepios shiranui (Esaki 1924) Iw92
  ‘Odius’ Stål 1867 non Lilljeborg 1866 C92
  Terioaphis trifolii C81
  Amphisalta zelandica H97
  Phera Stål 1864 K03
  Luederwaldtia Schmidt 1922 KO06
  Atopozelus pallens M07
  Poppea evelyna MG06
  Archiconiopteryx GE05
  Laodelphax striatellus R96
  Cephaloxys Signoret 1847 B83
  Anacornutipo lignosa S61
  Metastemma quinquemaculata Lucas 1847 E12
  Haematoloecha Stäl 1874 D56
  ‘Limonia’ Carvalho 1985 nec Meigen 1803 nec Agassiz 1846 KA-Z11
  Liops Fieber 1870 KA-Z11
  Umbronia spinosa Im92
  Sagotylus confluentus Im92
  Graphocephala BM76
    |--G. coccinea BM76
    `--G. versuta E66
  Proceiphilus tesselatus BM76
  Agrioaphis kuricola BM76
  Macropis viridis BM76
  Coenocorixa expleta WL09
  Laccogrypota grandis BMW13
  Prosapia bicincta BMW13
  Anacanthopus Montandon 1894 HW92
  Poekillopteridae K08
    |--Vanua vitiensis Kirkaldy 1907 K08
    |--Plestia marginata K08
    `--Euricania tristicula K08
  Amaurus Burmeister 1835 C96
  Naematopus albithorax Boisduval 1835 B35
  Archijassus [Archijassidae] RJ93
    `--A. plurinervis Zhang 1985 RJ93
  Chiliocyclidae RJ93
  Granulidae RJ93
  Magnacicadidae RJ93
  Prosbolecicada [Prosbolecicadidae] RJ93
    `--P. gondwanica Pinto 1987 RJ93
  Hotinus R13
    |--H. candelarius R13
    `--H. pyrorrhynchus R13
  Idiocorinae B50
    |--Paskia B50
    `--Idiocoris B50
  Plecophlebus Cockerell 1917 P92
    `--*P. nebulosus Cockerell 1917 P92
  Praecoris dominicana P92
  ‘Richteria’ Tode 1966 nec Jones 1874 nec Girault 1920 EH19
  Mevania Harold 1874 L90
  ‘Coryna’ Wolff 1811 non Bosc 1802 BP02

*Type species of generic name indicated


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