Paraneoptera

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

Belongs within: Pterygota.
Contains: Hypoperlina, Psocodea, Thripida, Hemiptera.

The Paraneoptera are a group of insects characterised by a tendency to an elongate head due to the enlargement of the postclypeus. This is in turn related to enlargement of the cibarial dilator muscles at the front of pharynx, allowing suction of fluids. Living paraneopterans are divided between three major clades, the Psocodea, Thripida and Hemiptera. Thripida and Hemiptera are commonly regarded as forming a clade Condylognatha based on mouthpart structures such as expanded hypopharyngeal apodemes, unicondylar mandibular and lacinial stylets, and a narrowed labrum, together with a dorsal shift of the anterior tentorial pits (Grimaldi & Engel 2005).

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. Where known, hypoperlidans had chewing mouthparts and often an elongate pronotum. Perielytron mirabile from the Early Permian of the Ural Mountains is distinguished from other Hypoperlida by the possession of broad, subelytrised fore wings (Rasnitsyn 2002b). The Miomoptera, another group of mostly minute insects known from the Late Carboniferous and Permian, have variously been regarded as stem-representatives of the Paraneoptera or their sister group, the Holometabola. Miomopterans had similar fore and hind wings, lacking cross-veins and lacking an anal fan in the hind wing. Where body fossils are known, they had mandibulate mouthparts, four-segmented tarsi and short cerci (Grimaldi & Engel 2005). 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).

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.: BruayaphisNR13
    |--HypoperlidaR02b
    |    |  i. s.: RhynomaloptilaR02b
    |    |         RhipidiopteraR02b
    |    |         Psoroptera [Psoropteridae]R02b
    |    |         Homoeodictyon [Homoeodictyidae]R02b
    |    |           `--H. elongatumR02a
    |    |--HypoperlinaR02b
    |    `--Perielytron Zalleskii 1948R02b, KN13 [Perielytridae]
    |         `--P. mirabileR02b
    |--Miomoptera [Palaeomanteida, Palaeomanteidea]GE05
    |    |  i. s.: SaaromiopteraR02d
    |    |--MetropatorR02b [MetropatoridaeGE05]
    |    |    `--M. pusillusR02b
    |    |--ArchaemiopteridaeR02d
    |    |    |--ArchemiopteraR02d
    |    |    `--Eodelopterum priscum Schmidt 1962IB11
    |    |--PalaeomantiscidaeR02d
    |    |    |--Palaeomantina pentameraR02d
    |    |    `--Sellardiopsis [incl. Palaeomantisca]R02d
    |    |         `--S. conspicua [incl. Palaeomantisca lata]R02d
    |    `--PalaeomanteidaeR02d
    |         |--Epimastax [Epimastacidae]R02d
    |         |    `--E. soyanensisR02d
    |         |--Palaeomantis Handlirsch 1904 (see below for synonymy)R02d
    |         |    |--*P. schmidtiR02d
    |         |    `--P. aestivusR02d
    |         `--Delopterum [incl. Miomatoneurella, Stephanomioptera]R02d
    |              |--D. kaltanicumR70
    |              |--D. rasnitsyniR02d
    |              `--D. sinuosumK-P91
    `--+--Permopsocida [Permopsocina]GE05
       |    |--ArchipsyllidaeR02c
       |    |--PermopsocusR70 [PermopsocidaeGE05]
       |    |    `--P. latipennisR70
       |    `--Psocidiidae [Dichentomidae]GE05
       |         |--EopsyllaR02c
       |         |--Dichentomum tinctumR02c, K-P91
       |         `--Parapsocidium uralicumR02c
       `--+--+--MontanuraliidaeN02
          |  `--+--PsocodeaGE05
          |     `--Sojanopus Novokshonov 2002 [Sojanoperidae]N02
          |          `--*S. festivum Novokshonov 2002N02
          `--Condylognatha [Aphidii]GE05
               |--ThripidaGE05
               `--HemipteraGE05

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

References

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[IB11] Ilger, J.-M., & C. Brauckmann. 2011. The smallest Neoptera (Baryshnyalidae fam. n.) from Hagen-Vorhalle (early Late Cretaceous: Namurian B; Germany). ZooKeys 130: 91–102.

[KN13] Kirejtshuk, A. G., & A. Nel. 2013. Skleroptera, a new order of holometabolous insects (Insecta) from the Carboniferous. Zoosystematica Rossica 22 (2): 247–257.

[K-P91] Kukalová-Peck, J. 1991. Fossil history and the evolution of hexapod structures. In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 1 pp. 141–179. Melbourne University Press: Carlton (Victoria).

[NR13] Nel, A., P. Roques, P. Nel, A. A. Prokin, T. Bourgoin, J. Prokop, J. Szwedo, D. Azar, L. Desutter-Grandcolas, T. Wappler, R. Garrouste, D. Coty, D. Huang, M. S. Engel & A. G. Kirejtshuk. 2013. The earliest known holometabolous insects. Nature 503: 257–261.

[N02] Novokshonov, V. G. 2002. New enigmatic insects (Insecta: Hypoperlidea?: Sojanoperidae) from the Upper Permian of northern Russia. Paleontologicheskii Zhurnal 2002 (1): 51–53 (translated: Paleontological Journal 36 (1): 48–49).

[R02a] Rasnitsyn, A. P. 2002a. Subclass Scarabaeona Laicharting, 1781. The winged insects (=Pterygota Lang, 1888). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 75–83. Kluwer Academic Publishers: Dordrecht.

[R02b] Rasnitsyn, A. P. 2002b. Cohors Cimiciformes Laicharting, 1781. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 104–115. Kluwer Academic Publishers: Dordrecht.

[R02c] Rasnitsyn, A. P. 2002c. Superorder Psocidea Leach, 1815. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 125–133. Kluwer Academic Publishers: Dordrecht.

[R02d] Rasnitsyn, A. P. 2002d. Cohors Scarabaeiformes Laicharting, 1781. The holometabolans (=Holometabola Burmeister, 1835, =Endopterygota Sharp, 1899, =Oligoneoptera Martynov, 1938). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 157–164. Kluwer Academic Publishers: Dordrecht.

[R70] Riek, E. F. 1970. Fossil history. In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 168–186. Melbourne University Press.

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