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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
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
Superordines: Condylognatha - Psocodea
[list of superordines after Grimaldi & Engel (2005: 147, table 4.1)]
Overview of orders

Hemiptera - Phthiraptera - "Psocoptera" - Thysanoptera
[list of ordines after Grimaldi & Engel (2005: 147, table 4.1)]

Name

Paraneoptera
References

Grimaldi, D.; Engel, M.S. 2005: Evolution of the insects. Cambridge University Press, New York, USA. limited preview on Google books
Lambkin, K.J. 2018. A new genus and species of the family Archipsyllidae (Insecta: Paraneoptera: Permopsocida) from the Late Triassic of Queensland. Zootaxa 4382(1): 192–194. DOI: 10.11646/zootaxa.4382.1.9 Reference page.
Yoshizawa, K. & Lienhard, C. 2016. Bridging the gap between chewing and sucking in the hemipteroid insects: new insights from Cretaceous amber. Zootaxa 4079(2): 229–245. DOI: 10.11646/zootaxa.4079.2.5 Full article (PDF).Reference page.
Yoshizawa, K.; Saigusa, T. 2001: Phylogenetic analysis of paraneopteran orders (Insecta: Neoptera) based on forewing base structure, with comments on monophyly of Auchenorrhyncha (Hemiptera). Systematic entomology, 26: 1–13.
Liang, F-Y., Zhang, W-W. & Liu, X-Y. 2016. A new genus and species of the paraneopteran family Archipsyllidae in mid-Cretaceous amber of Myanmar. Zootaxa 4105(5): 483–490. DOI: 10.11646/zootaxa.4105.5.4. Reference page.

Vernacular names
français: Paranéoptère
日本語: 準新翅類 (じゅんしんしるい)
中文: 副新翅總目

Paraneoptera or Acercaria[1] is a superorder of insects which includes lice (bark lice and true lice), thrips, and hemipterans, the true bugs.[2] It also includes the extinct order Permopsocida, known from fossils dating from the Early Permian to the mid-Cretaceous.

All of the insects classified here exhibit various “reductions” or “simplifications” from the primitive body-plan found in typical polyneopterans. Cerci, for example, are entirely absent in all living paraneopterans (Acercaria meaning without cerci). Other “reductions” occur in wing venation, in the number of tarsal segments (not more than three), in the number of Malpighian tubules (not more than six), and in the number of ganglia present in the ventral nerve cord (not more than five).

The mouthparts of the Paraneoptera reflect diverse feeding habits. Basal groups are microbial surface feeders, whereas more advanced groups feed on plant or animal fluids.[2]

Phylogeny

Paraneoptera consists of Psocodea {lice}, along with their sister clade, the monophyletic grouping Condylognatha that contains Hemiptera (true bugs) and Thysanoptera (thrips). However, analysis has shown that Psocodea could instead be the sister taxon to Holometabola, which would render Paraneoptera as paraphyletic.[3]

Here is a simple cladogram showing the traditional relationships with a monophyletic Paraneoptera:[3]

Neoptera

Polyneoptera

Eumetabola

Holometabola

Paraneoptera

Psocodea

Condylognatha

Thysanoptera (thrips)

Hemiptera (true bugs)



Here is an alternative cladogram showing Paraneoptera as paraphyletic, with Psocodea as sister taxon to Holometabola:[3]

Neoptera

Polyneoptera

Eumetabola

Holometabola

Psocodea

Condylognatha

Thysanoptera (thrips)

Hemiptera (true bugs)


Within Paraneoptera, Psocodea contains the two orders Phthiraptera (lice) and Psocoptera (booklice, barklice or barkflies). However, studies have shown that Phthiraptera is in fact nested deep within Psocoptera, making Psocoptera paraphyletic and an invalid grouping.[4][3]

Assuming Paraneoptera is monophyletic, here is a more detailed cladogram showing the internal relationships, and how Phthiraptera falls within Psocodea:[3]

Other insects Annualreportofag1119021903univ 0052AA2 Figure 1.jpg

Paraneoptera
Psocodea

Trogiomorpha (barklice) Psocoptera icon.png

Psocomorpha (barklice) Psocoptera icon.png

Troctomorpha (paraphyletic with respect to Phthiraptera)

Amphientometae

Sphaeropsocidae

Liposcelididae (booklice) Liposcelis icon.png

Phthiraptera (lice) Lipeurus forficulatus f.png

Condylognatha

Thysanoptera (thrips) Taeniothrips inconsequens.jpg

Hemiptera (true bugs)

Sternorrhyncha (aphids) Aphid icon.png

Heteroptera (shield bugs, assassin bugs, etc) Bug (PSF).jpg

Coleorrhyncha (moss bugs) HEMI Peloridiidae Oiophysa distincta 1.png

Auchenorrhyncha

Fulgoromorpha (planthoppers) Acanaloniidae.jpg

Cicadomorpha (cicadas, leafhoppers, spittlebugs, etc) Marbaarus bubalus Distant.jpg



Taxonomy
Hemiptera
Main article: Hemiptera

Hemiptera /hɛˈmɪptərə/ is an order of insects most often known as the true bugs (cf. bug), comprising around 50,000–80,000 species of cicadas, aphids, planthoppers, leafhoppers, shield bugs, bed bugs and others. They range in size from 1 millimetre (0.039 in) to around 15 centimetres (5.9 in), and share a common arrangement of sucking mouthparts.
Thrips
Main article: thrips

Order Thysanoptera includes 5,500 species classified into two suborders distinguished by the ovipositor. Terebrantia have a well-developed conical ovipositor, while the Tubulifera do not. Instead the abdomen is drawn out in the shape of a tube. These insects are called thrips.
Psocoptera
Main article: Psocoptera

Psocoptera, the bark lice, include 4,400 described species arranged in 3 suborders, Trogiomorpha, Troctomorpha, and Psocomorpha. There are 50 families of bark lice with over 200 genera. This is the first insect order to show the beginnings of a transition to sucking mouthparts. Recent studies have found that Psocoptera is paraphyletic, with Phthiraptera nested deep within Psocoptera, within the now-paraphyletic suborder Troctomorpha, making Psocoptera an invalid grouping.[4][3]
Phthiraptera
Main article: louse

Phthiraptera, the lice, includes 5,000 described species divided into 4 suborders. The Amblycera is the most basal group and parasitize birds and mammals. The Ischnocera is the largest suborder and parasitize mostly birds and some groups of mammals. The Rhynchophthirina, the elephant lice, consists of only 3 species that parasitize elephants and wild pigs in Africa. The Anoplura (sucking lice) parasitize only mammals. Phthiraptera has been found to be contained within the order Psocoptera.[4][3]
Permopsocida
Permopsocida fossil Psocorrhyncha burmitica
Main article: Permopsocida

The extinct order Permopsocida includes 18 genera divided into 3 families, dating from the Early Permian (Asselian) to the early Late Cretaceous (Cenomanian), Permopsocida are more closely related to thrips and bugs than to lice.[5]
References

Hu Li; Renfu Shao; Nan Song; Fan Song; Pei Jiang; Zhihong Li; Wanzhi Cai (23 February 2015). "Higher-level phylogeny of paraneopteran insects inferred from mitochondrial genome sequences". Scientific Reports. 5 (8527): 8527. Bibcode:2015NatSR...5E8527L. doi:10.1038/srep08527. PMC 4336943. PMID 25704094.
David A. Grimaldi & Michael S. Engel (2005). "The Paraneopteran Orders". Evolution of the Insects. Volume 1 of Cambridge Evolution Series. Cambridge University Press. pp. 261–330. ISBN 978-0-521-82149-0.
Kevin P. Johnson; Christopher H. Dietrich; Frank Friedrich; Rolf G. Beutel; Benjamin Wipfler; Ralph S. Peters; Julie M. Allen; Malte Petersen; Alexander Donath; Kimberly K. O. Walden; Alexey M. Kozlov; Lars Podsiadlowski; Christoph Mayer; Karen Meusemann; Alexandros Vasilikopoulos; Robert M. Waterhouse; Stephen L. Cameron; Christiane Weirauch; Daniel R. Swanson; Diana M. Percy; Nate B. Hardy; Irene Terry; Shanlin Liu; Xin Zhou; Bernhard Misof; Hugh M. Robertson; Kazunori Yoshizawa (Dec 2018). "Phylogenomics and the evolution of hemipteroid insects". Proceedings of the National Academy of Sciences. 115 (50): 12775–12780. doi:10.1073/pnas.1815820115. PMC 6294958. PMID 30478043.
C. H. C. Lyal (1985). "Phylogeny and classification of the Psocodea, with particular reference to the lice (Psocodea: Phthiraptera)". Systematic Entomology. 10 (2): 145–165. doi:10.1111/j.1365-3113.1985.tb00525.x. S2CID 86331606.
Huang, Di-Ying; Bechly, Günter; Nel, Patricia; Engel, Michael S.; Prokop, Jakub; Azar, Dany; Cai, Chen-Yang; van de Kamp, Thomas; Staniczek, Arnold H.; Garrouste, Romain; Krogmann, Lars (March 2016). "New fossil insect order Permopsocida elucidates major radiation and evolution of suction feeding in hemimetabolous insects (Hexapoda: Acercaria)". Scientific Reports. 6 (1): 23004. Bibcode:2016NatSR...623004H. doi:10.1038/srep23004. ISSN 2045-2322. PMC 4785345. PMID 26961785.

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