| Psocodea Temporal range: Late Jurassic – Recent | |
|---|---|
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| An unidentified bark louse in the family Stenopsocidae | |
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| Human body louse | |
Scientific classification  | |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Clade: | Pancrustacea |
| Class: | Insecta |
| Clade: | Eumetabola |
| Order: | Psocodea Hennig, 1966 |
| Suborders[1] | |
Psocodea is a taxonomic group of insects comprising the bark lice, book lice and parasitic lice.[2] It was formerly considered a superorder, but is now generally considered by entomologists as an order.[1][3][4] 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.[5][6]
Psocodea contains around 11,000 species, divided among three suborders and more than 70 families.[1][2][7] They range in size from 1–10 millimetres (0.04–0.4 in) in length.
The species known as booklice received their common name because they are commonly found amongst old books—they feed upon the paste used in binding. The barklice are found on trees, feeding on algae and lichen.
External phylogeny
Psocodea has been recovered as a monophyletic group in recent studies. Their next closest relatives are traditionally recognized as the monophyletic grouping Condylognatha that contains Hemiptera (true bugs) and Thysanoptera (thrips), which all combined form the group Paraneoptera. However, this is somewhat unclear, as analysis has shown that Psocodea could instead be the sister taxon to Holometabola, which would render Paraneoptera as paraphyletic.[6]
Here is a simple cladogram showing the traditional relationships with a monophyletic Paraneoptera:[6]
| Neoptera |
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Here is an alternative cladogram showing Paraneoptera as paraphyletic, with Psocodea as sister taxon to Holometabola:[6]
| Neoptera |
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Paraneoptera | ||||||||||||||||||||||||
Internal phylogeny
Here is a cladogram showing the relationships within Psocodea:[3]
| Psocodea | |
Classification
The order Psocodea (formerly 'Psocoptera') is divided into three extant suborders.
Suborder Trogiomorpha
Trogiomorpha have antennae with many segments (22–50 antennomeres) and always three-segmented tarsi.[8]
Trogiomorpha is the smallest suborder of the Psocoptera sensu stricto (i.e., excluding Phthiraptera), with about 340 species in 7 families, ranging from the fossil family Archaeatropidae with only a handful of species to the speciose Lepidopsocidae (over 200 species). Trogiomorpha comprises infraorder Atropetae (extant families Lepidopsocidae, Psoquillidae and Trogiidae, and fossil families Archaeatropidae and Empheriidae) and infraorder Psocathropetae (families Psyllipsocidae and Prionoglarididae).
Suborder Troctomorpha
Troctomorpha have antennae with 15–17 segments and two-segmented tarsi.
Troctomorpha comprises the Infraorder Amphientometae (families Amphientomidae, Compsocidae, Electrentomidae, Musapsocidae, Protroctopsocidae and Troctopsocidae) and Infraorder Nanopsocetae (families Liposcelididae, Pachytroctidae and Sphaeropsocidae). Troctomorpha are now known to also contain the order Phthiraptera (lice), and are therefore paraphyletic, as are Psocoptera as a whole.
Some Troctomorpha, such as Liposcelis (which are similar to lice in morphology), are often found in birds' nests, and it is possible that a similar behavior in the ancestors of lice is at the origin of the parasitism seen today.[8]
Suborder Psocomorpha
Psocomorpha are notable for having antennae with 13 segments. They have two- or three-segmented tarsi, this condition being constant (e.g., Psocidae) or variable (e.g., Pseudocaeciliidae) within families. Their wing venation is variable, the most common type being that found in the genus Caecilius (rounded, free areola postica, thickened, free pterostigma, r+s two-branched, m three-branched). Additional veins are found in some families and genera (Dicropsocus and Goja in Epipsocidae, many Calopsocidae, etc.)
Psocomorpha is the largest suborder of the Psocoptera sensu stricto (i.e., excluding Phthiraptera), with about 3,600 species in 24 families, ranging from the species-poor Bryopsocidae (2 spp.) to the speciose Psocidae (about 900 spp).[8] Psocomorpha comprises Infraorder Epipsocetae (families Cladiopsocidae, Dolabellopsocidae, Epipsocidae, Neurostigmatidae and Ptiloneuridae), Infraorder Caeciliusetae (families Amphipsocidae, Asiopsocidae, Caeciliusidae, Dasydemellidae and Stenopsocidae), Infraorder Homilopsocidea (families Archipsocidae, Bryopsocidae, Calopsocidae, Ectopsocidae, Elipsocidae, Lachesillidae, Mesopsocidae, Peripsocidae, Philotarsidae, Pseudocaeciliidae and Trichopsocidae) and Infraorder Psocetae (families Hemipsocidae, Myopsocidae, Psilopsocidae and Psocidae).
References
- 1 2 3 Johnson, Kevin P.; Smith, Vincent S. (2021). "Psocodea species file online, Version 5.0". Retrieved 2021-11-01.
- 1 2 P. J. Gullan & P. S. Cranston (2010). "Taxobox 17 – Psocodea: "Psocoptera" (bark lice and book lice)". The Insects: an Outline of Entomology (4th ed.). John Wiley & Sons. p. 479. ISBN 9781444317671.
- 1 2 De Moya, Robert S.; Yoshizawa, Kazunori; Walden, Kimberly K. O.; Sweet, Andrew D.; et al. (2021). "Phylogenomics of Parasitic and Nonparasitic Lice (Insecta: Psocodea): Combining Sequence Data and Exploring Compositional Bias Solutions in Next Generation Data Sets". Systematic Biology. 70 (4): 719–738. doi:10.1093/sysbio/syaa075. PMID 32979270.
- ↑ "Psocodea". GBIF. Retrieved 2021-11-01.
- ↑ 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. Bibcode:1985SysEn..10..145L. doi:10.1111/j.1365-3113.1985.tb00525.x. S2CID 86331606.
- 1 2 3 4 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. Bibcode:2018PNAS..11512775J. doi:10.1073/pnas.1815820115. PMC 6294958. PMID 30478043.
- ↑ David Grimaldi & Michael S. Engel (2005). "The paraneopteran orders". Evolution of the Insects. Cambridge Evolution Series. Cambridge University Press. pp. 216–330. ISBN 9780521821490.
- 1 2 3 C. Lienhard & C. N. Smithers (2002). "Psocoptera (Insecta): World Catalogue and Bibliography". Instrumenta Biodiversitatis. 5.
External links
- National Barkfly Recording Scheme
- Psoco Net
- Tree of Life: Psocodea
- Archipsocus nomas, a webbing barklouse on the UF / IFAS Featured Creatures Web site
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