Paraneoptera

Reconstruction of Palaeomantina pentamera, from Rasnitsyn (2002d).

Belongs within: Pterygota.
Contains: Hypoperlina, Psocodea, Thripida, Euhemiptera, Protopsyllidiidae, Sternorrhyncha.

The Paraneoptera are a group of insects characterised by a tendency to an elongate head, due to the enlargement of the postclypeus. Living paraneopterans are divided between three major clades, the Psocodea, Thripida and Hemiptera. The Middle Carboniferous to Permian Hypoperlida were regarded by Rasnitsyn (2002b) as stem-paraneopterans, citing the presence of a maxilla with detached, rod-like lacinia as a synapomorphy for this clade. Many hypoperlidans, however, are only known from wings. The Miomoptera, another group of mostly minute insects known from the Late Carboniferous and Permian (Grimaldi & Engel 2005), have variously been regarded as stem-representatives of the Paraneoptera or their sister group, the Holometabola. The Permian Permopsocida have been suggested to be related to the Psocodea but the presence of some primitive features such as small cerci and four- or five-segmented tarsi (versus no cerci and no more than three tarsomeres in living paraneopterans) suggests they may be stem-paraneopterans (Grimaldi & Engel 2005).

The Hemiptera, commonly known as bugs, are characterised by the possession of a sucking beak. 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).

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.

Characters (from Grimaldi & Engel 2005): Postclypeus large, with large cibarial dilator muscles; labial palps reduced or lost; asymmetrical mandibles; maxillary laciniae slender, long, detached from stipes, independently movable; legs with three or fewer tarsomeres; winged forms usually with abdominal trichobothria; ganglia in abdomen fused into one large mass; cerci lost.

<==Paraneoptera [Acercaria, Cimiciformes, Hypoperlidea]
    |  i. s.: Bruayaphis NR13
    |--Hypoperlida R02b
    |    |  i. s.: Rhynomaloptila R02b
    |    |         Rhipidioptera R02b
    |    |         Psoroptera [Psoropteridae] R02b
    |    |         Homoeodictyon [Homoeodictyidae] R02b
    |    |           `--H. elongatum R02a
    |    |--Hypoperlina R02b
    |    `--Perielytron Zalleskii 1948 R02b, KN13 [Perielytridae]
    |         `--P. mirabile R02b
    |--Miomoptera [Palaeomanteida, Palaeomanteidea] GE05
    |    |  i. s.: Saaromioptera R02d
    |    |--Metropator R02b [Metropatoridae GE05]
    |    |    `--M. pusillus R02b
    |    |--Archaemiopteridae R02d
    |    |    |--Archemioptera R02d
    |    |    `--Eodelopterum priscum Schmidt 1962 IB11
    |    |--Palaeomantiscidae R02d
    |    |    |--Palaeomantina pentamera R02d
    |    |    `--Sellardiopsis [incl. Palaeomantisca] R02d
    |    |         `--S. conspicua [incl. Palaeomantisca lata] R02d
    |    `--Palaeomanteidae R02d
    |         |--Epimastax [Epimastacidae] R02d
    |         |    `--E. soyanensis R02d
    |         |--Palaeomantis Handlirsch 1904 (see below for synonymy) R02d
    |         |    |--*P. schmidti R02d
    |         |    `--P. aestivus R02d
    |         `--Delopterum [incl. Miomatoneurella, Stephanomioptera] R02d
    |              |--D. kaltanicum R70
    |              |--D. rasnitsyni R02d
    |              `--D. sinuosum K-P91
    `--+--Permopsocida [Permopsocina] GE05
       |    |--Archipsyllidae R02c
       |    |--Permopsocus R70 [Permopsocidae GE05]
       |    |    `--P. latipennis R70
       |    `--Psocidiidae [Dichentomidae] GE05
       |         |--Eopsylla R02c
       |         |--Dichentomum tinctum K-P91
       |         `--Parapsocidium uralicum R02c
       `--+--+--Montanuraliidae N02
          |  `--+--Psocodea GE05
          |     `--Sojanopus Novokshonov 2002 [Sojanoperidae] N02
          |          `--*S. festivum Novokshonov 2002 N02
          `--Condylognatha [Aphidii] GE05
               |--Thripida Z02
               `--Hemiptera [Arthroidignatha, Cimicida, Homoptera, Rhynchota] GE05
                    |  i. s.: 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
                    |--Euhemiptera NR13
                    |--+--Protopsyllidiidae GE05
                    |  `--Sternorrhyncha GE05
                    `--Archescytinidae [Archescytinina, Archescytinoidea, Palaeohemiptera, Paleorrhyncha] GE05
                         |--Permothrips [Permothripidae] SP02
                         |--Maueria SP02
                         |--Permoscytina SP02
                         |--Archescytina permiana K-P91
                         `--Protopincombea obscura Evans 1943 RJ93, F71

Palaeomantis Handlirsch 1904 [incl. Miomatoneura Martynov 1927, Permodelopterum Kukalová 1963, Permonia Kukalová 1963, Permonikia Kukalová 1963, Perunopterum Kukalová 1963] R02d

*Type species of generic name indicated

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Last updated: 29 December 2021.