Paraneoptera

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Paraneoptera
Temporal range: Late Carboniferous–Recent
Magicicada septendecim.jpg
Magicicada septendecim , a cicada (Hemiptera)
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
(unranked): Eumetabola
(unranked): Paraneoptera
Orders

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.

Contents

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 (no more than three), only four Malpighian tubules, and only one complex of abdominal ganglia. [3]

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. [4]

Here is a simple cladogram showing the traditional relationships with a monophyletic Paraneoptera: [4]

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: [4]

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. [5] [4]

Assuming Paraneoptera is monophyletic, here is a more detailed cladogram showing the internal relationships, and how Phthiraptera falls within Psocodea: [4]

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

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

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

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. [5] [4]

Phthiraptera

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. [5] [4]

Permopsocida

Permopsocida fossil Psocorrhyncha burmitica Permopsocida.jpg
Permopsocida fossil Psocorrhyncha burmitica

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. [6]

Related Research Articles

<span class="mw-page-title-main">Louse</span> Order of insects

Louse is the common name for any member of the clade Phthiraptera, which contains nearly 5,000 species of wingless parasitic insects. Phthiraptera has variously been recognized as an order, infraorder, or a parvorder, as a result of developments in phylogenetic research.

<span class="mw-page-title-main">Hemiptera</span> Order of insects often called true bugs

Hemiptera is an order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from 1 mm (0.04 in) to around 15 cm (6 in), and share a common arrangement of piercing-sucking mouthparts. The name "true bugs" is often limited to the suborder Heteroptera.

<span class="mw-page-title-main">Homoptera</span> Suborder of true bugs

Homoptera is a suborder of order Hemiptera that is considered by some taxonomists to be paraphyletic, and therefore deprecated (obsolete). It was therefore split into the suborders Sternorrhyncha, Auchenorrhyncha, and Coleorrhyncha. The earlier work was based on nuclear DNA, but more recent phylogenetic analysis using mitochondrial DNA suggest that Homoptera may be a monophyletic group after all, a sister group of Heteroptera. The cause of the disparity in the analyses is suggested to be the long branch attraction effect in phylogenetic analysis, due to rapidly evolving DNA regions.

<span class="mw-page-title-main">Thrips</span> Order of insects

Thrips are minute, slender insects with fringed wings and unique asymmetrical mouthparts. Entomologists have described approximately 7,700 species. They fly only weakly and their feathery wings are unsuitable for conventional flight; instead, thrips exploit an unusual mechanism, clap and fling, to create lift using an unsteady circulation pattern with transient vortices near the wings.

<span class="mw-page-title-main">Pterygota</span> Subclass of insects

The Pterygota are a subclass of insects that includes all winged insects and the orders that are secondarily wingless.

<span class="mw-page-title-main">Exopterygota</span> Superorder of insects

The Exopterygota, also known as Hemipterodea, are a superorder of insects of the subclass Pterygota in the infraclass Neoptera, in which the young resemble adults but have externally developing wings. They undergo a modest change between immature and adult, without going through a pupal stage. The nymphs develop gradually into adults through a process of moulting.

<span class="mw-page-title-main">Psocoptera</span> Order of insects

Psocoptera are a paraphyletic group of insects that are commonly known as booklice, barklice or barkflies. The name Psocoptera has been replaced with Psocodea in recent literature, with the inclusion of the former order Phthiraptera into Psocodea.

A bird louse is any chewing louse of order Phthiraptera which parasitizes warm-blooded animals, especially birds. Bird lice may feed on feathers, skin, or blood. They have no wings, and their biting mouth parts distinguish them from true lice, which suck blood.

<span class="mw-page-title-main">Mandible (insect mouthpart)</span>

Insect mandibles are a pair of appendages near the insect's mouth, and the most anterior of the three pairs of oral appendages. Their function is typically to grasp, crush, or cut the insect's food, or to defend against predators or rivals. Insect mandibles, which appear to be evolutionarily derived from legs, move in the horizontal plane unlike those of vertebrates, which appear to be derived from gill arches and move vertically.

<span class="mw-page-title-main">Psocomorpha</span> Suborder of booklice

Psocomorpha is a suborder of barklice, booklice, and parasitic lice in the order Psocodea. There are more than 20 families and 5,300 described species in Psocomorpha.

<span class="mw-page-title-main">Trogiomorpha</span> Group of booklice

Trogiomorpha is one of the three major suborders of barklice, booklice, and parasitic lice in the order Psocodea, alongside Troctomorpha and Psocomorpha. There are about 8 families and more than 430 described species in Trogiomorpha. Trogiomorpha is widely agreed to be the earliest diverging of the three suborders, and retains the most primitive characteristics.

<span class="mw-page-title-main">Troctomorpha</span> Suborder of booklice

Troctomorpha is one of the three major suborders of Psocodea (barklice, booklice, and parasitic lice), alongside Psocomorpha and Trogiomorpha. There are more than 30 families and 5,800 described species in Troctomorpha. The order includes parasitic lice, which are most closely related to the booklice family Liposcelididae.

Pachytroctidae is a family of thick barklice in the order Psocodea. There are about 15 genera and at least 90 described species in Pachytroctidae.

<span class="mw-page-title-main">Ectopsocidae</span> Family of booklice

Ectopsocidae is a family of Psocodea belonging to the suborder Psocomorpha. The family includes fewer than 200 species, most of them in the genus Ectopsocus.

<span class="mw-page-title-main">Homilopsocidea</span> Group of booklice

Homilopscocidea is an infraorder of Psocodea. It is probably a paraphyletic group, still in use for lack of a better solution. There are about 7 families and more than 1,200 described species in Homilopsocidea.

<span class="mw-page-title-main">Psocodea</span> Order of insects

Psocodea is a taxonomic group of insects comprising the bark lice, book lice and parasitic lice. It was formerly considered a superorder, but is now generally considered by entomologists as an order. Despite the greatly differing appearance of parasitic lice (Phthiraptera), they are believed to have evolved from within the former order Psocoptera, which contained the bark lice and book lice, now found to be paraphyletic. They are often regarded as the most primitive of the hemipteroids. Psocodea contains around 11,000 species, divided among four suborders and more than 70 families. They range in size from 1–10 millimetres (0.04–0.4 in) in length.

<span class="mw-page-title-main">Epipsocetae</span> Infraorder of booklice

Epipsocetae is an infraorder of psocids in the order Psocodea. There are about 5 families and more than 480 described species in Epipsocetae.

<span class="mw-page-title-main">Permopsocida</span> Extinct genus of insects

Permopsocida is an extinct order of insects known from the Early Permian to the Mid-Cretaceous. It is part of Paraneoptera, alongside bark lice, bugs and thrips. Within Paraneoptera it is considered to be closer to the clade containing bugs and thrips rather than bark lice, with an estimated divergence during the Late Carboniferous. The group was first named as a suborder by Robert John Tillyard in 1926, and was raised to a full order by Huang et al. in 2016. It is currently divided up into three families, Psocidiidae which is known from the Permian to Liassic. Permopsocidae which is only known from the Permian, and Archipsyllidae, which is known from the Late Triassic to mid-Cretaceous (Cenomanian). While most members of the group are known from compression fossils, several members of Archipsyllidae are 3 dimensionally preserved in Burmese amber, which has helped clarify the morphology and phylogenetic position of the group. The morphology of the mouthparts suggests that they were capable of suction feeding and chewing, with preserved angiosperm pollen grains in the gut of Psocorrhyncha suggesting that at least some members of the group were pollenivorous.

<span class="mw-page-title-main">Condylognatha</span> Superorder of insects

Condylognatha or Panhemiptera is a monophyletic grouping (superorder) that contains Hemiptera and Thysanoptera (thrips). Condylognatha belongs to Paraneoptera, which include its sister group, lice (Psocodea).

References

  1. 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.
  2. 1 2 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.
  3. Letsch, H. O.; Meusemann, K.; Wipfler, B.; Schütte, K.; Beutel, R.; Misof, B. (2012). "Insect phylogenomics: Results, problems and the impact of matrix composition". Proceedings. Biological Sciences. 279 (1741): 3282–3290. doi:10.1098/rspb.2012.0744. PMC   3385742 . PMID   22628473.
  4. 1 2 3 4 5 6 7 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.
  5. 1 2 3 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.
  6. 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.