Permopsocida

Permopsocida is an extinct order of insects known from the Early Permian to Mid-Cretaceous. It is part of Paraneoptera, alongside bark lice (including 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.[1] 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[2] 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 phyogenetic position of the group.[1][3][4][5] 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.[1]

Comparison of the head morphology of Permospocida (b) with that of Psocodea (a) Thysanoptera (c) and Hemiptera (d)
Head of Psocorrhyncha

Permopsocida
Temporal range: Early Permian–Cenomanian
Psocorrhyncha burmitica
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
(unranked): Paraneoptera
Order: †Permopsocida
Tillyard, 1926
Families
  • †Psocidiidae
  • †Permopsocidae
  • †Archipsyllidae

Taxonomy

From Huang et al 2016 and subsequent literature

  • †Psocidiidae Tillyard, 1926
    • †Dichentomum Tillyard, 1926 Koshelevka Formation, Russia, Kungurian, Bursum Formation, New Mexico, Asselian, Wellington Formation, Kansas, Oklahoma, Artinskian
    • †Hypopsylla Prokop, Garrouste & Nel, 2016 Croudace Bay Formation, Australia, Changhsingian
    • †Liassopsocus Ansorge, 1996 "Green Series", Germany, Toarcian
    • †Austropsocidium Tillyard 1935 Croudace Bay Formation, Australia, Changhsingian
    • †Stenopsocidium Tillyard 1935 Croudace Bay Formation, Australia, Changhsingian
  • †Permopsocidae Tillyard, 1926
    • †Permopsocus Tillyard, 1926 Wellington Formation, Kansas, Artinskian
    • †Lithopsocidium Carpenter 1932 Wellington Formation, Kansas, Artinskian
    • †Orthopsocus Carpenter 1932 Wellington Formation, Kansas, Artinskian
    • †Progonopsocus Tillyard 1926 Wellington Formation, Kansas, Artinskian
  • †Archipsyllidae Handlirsch, 1906
    • †Eopsylla Vishnyakova 1976 Iva-Gora Beds Formation, Russia, Roadian
    • †Archipsylla Handlirsch, 1908 "Green Series", Germany, Toarcian Daohugou, China, Callovian, Karabastau Formation, Kazakhstan, Oxfordian
    • †Archipsyllodes Vishniakova, 1976 Zaza Formation, Russia, Aptian
    • †Archipsyllopsis Vishniakova, 1976 Zaza Formation, Russia, Aptian
    • †Bittacopsocus Beutel et al. 2019 Burmese amber, Myanmar, Cenomanian
    • †Burmopsylla Liang, Zhang & Liu, 2016 Burmese amber, Myanmar, Cenomanian
    • †Dinmopsylla Lambkin, 2018 Blackstone Formation, Australia, Norian
    • †Mydiognathus Yoshizawa & Lienhard, 2016 Burmese amber, Myanmar, Cenomanian
    • †Psocorrhyncha Huang et al. 2016 Burmese amber, Myanmar, Cenomanian

References
  1. 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.
  2. Lambkin, Kevin J. (2018-02-20). "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. ISSN 1175-5334. PMID 29689945.
  3. Yoshizawa, K.; Lienhard, C. (2016). "Bridging the gap between chewing and sucking in the hemipteroid insects:new insights from Cretaceous amber". Zootaxa. 4079 (2): 229–45. doi:10.11646/zootaxa.4079.2.5. ISSN 1175-5334. PMID 27396002.
  4. Liang, F.; Zhang, W.; Liu, X. (2016). "A new genus and species of the paraneopteran family Archipsyllidae in mid-Cretaceous amber of Myanmar". Zootaxa. 4105 (5): 483–90. doi:10.11646/zootaxa.4105.5.4. ISSN 1175-5334. PMID 27394792.
  5. Beutel, R. G.; Prokop, J.; Müller, P.; Pohl, H. (2019). "†Bittacopsocus—a new bizarre genus of †Permopsocida (Insecta) from Burmese Cretaceous amber". Zootaxa. 4576 (2): 357–366. doi:10.11646/zootaxa.4576.2.9. PMID 31715767.
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