Paoliida

Paoliida is an extinct order of winged insects that lived in the late Paleozoic. Historically, both their systematic position and composition were controversial – for instance they had been considered as palaeodictyopterans, as basal Neoptera, or as stem-group of Pterygota – but recent studies have resolved them as the sister group of Dictyoptera.^[1][2][3]

Paoliida Temporal range: Carboniferous–Permian PreꞒ Ꞓ O S D C P T J K Pg N
Fossils of Paoliida. A: Zdenekia silesiensis; B: Darekia sanguinea; C, D1, D2: Paoliidae gen et sp. indet.
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Cohort:	Polyneoptera
Order:	†Paoliida Handlirsch, 1906
Families
†Anthracoptilidae †Blattinopsidae †Paoliidae
Synonyms
Protoptera Rasnitsyn, 1977

Families and genera

After Prokop et al. (2014), Guan et al. (2016) and Nel & Poschmann (2021):^[1][4][5]

• †Anthracoptilidae Handlirsch, 1922 (Synonyms: Strephocladidae Martynov, 1938; Strephoneuridae Martynov, 1940; ?Adiphlebiidae Handlirsch, 1906)
  • ?†Adiphlebia Scudder, 1885
  • †Afrocladus Nel, Garrouste & Prokop, 2015^[6]
  • †Anthracoptilus Lameere, 1917
  • †Carrizocladus Rasnitsyn in Rasnitsyn et al., 2004
  • †Graticladus Novokshonov and Aristov, 2004
  • †Homocladus Carpenter, 1966
  • †Jarmilacladus Rasnitsyn and Aristov, 2004
  • †Lodevocladus Prokop et al., 2015^[7]
  • †Mesoptilus Lameere, 1917
  • †Mycteroptila Rasnitsyn, 1977
  • †Paracladus Carpenter, 1966
  • †Pruvostia Bolton, 1921
  • †Pseudomesoptilus Guan et al., 2016
  • †Rhinomaloptila Rasnitsyn, 1977
  • †Spargoptilon Kukalová, 1965
  • †Strephocladus Scudder, 1885
  • †Strephoneura Martynov, 1940
  • †Strephoptilus Rasnitsyn and Aristov, 2013
  • †Westphaloptilus Guan et al., 2016
• †Blattinopsidae Bolton, 1925 (Synonym: Klebsiellidae Handlirsch, 1906)
  • †Alienus Handlirsch, 1906
  • †Avionblattinopsis Quispe et al., 2021^[8]
  • †Balduriella Meunier, 1925
  • †Blattinopsis Giebel, 1867
  • †Glaphyrophlebia Handlirsch, 1906
  • †Klebsiella Meunier, 1908
  • †Microblattina Scudder, 1895
  • †Protoblattiniella Meunier, 1912
  • †Rhipidioptera Brongniart, 1894
  • †Stephanopsis Kukalová, 1958
  • †Westphaloblattinopsis Béthoux and Jarzembowski, 2010
• †Paoliidae Handlirsch, 1906 (Synonyms: Ideliidae Zalessky, 1928; Herbstialidae Schmidt, 1953; Permulidae Aristov & Storozhenko, 2011; Permotermopsidae Martynov, 1937)
  • †Acropermula Aristov and Storozhenko, 2011
  • †Aenigmidelia Sharov, 1961
  • †Archidelia Sharov, 1961
  • †Carbonidelia Nel, Garrouste & Jouault, 2023^[9]
  • †Darekia Prokop et al., 2012
  • †Herbstiala Schmidt, 1952
  • †Holasicia Kukalová, 1958
  • †Idelina Storozhenko, 1992
  • †Kemperala Brauckmann, 1984
  • †Khosaridelia Storozhenko, 1992
  • †Kortshakolia Sharov, 1961
  • †Kochopteron Brauckmann, 1984
  • †Mertovia Prokop and Nel, 2007
  • †Micaidelia Aristov 2004
  • †Mongoloidelia Storozhenko, 1992
  • †Olinka Kukalová, 1958
  • ?†Paolekia Riek, 1976
  • †Paolia Smith, 1871
  • †Paoliola Handlirsch, 1919
  • †Permotermopsis Martynov, 1937
  • †Permula Handlirsch, 1919
  • †Protoblattina Meunier, 1909
  • †Pseudofouquea Handlirsch, 1906
  • †Silesiapteron Prokop et al., 2013
  • †Simplexpaolia Santos et al., 2023^[10]
  • †Sojanidelia Storozhenko, 1992
  • †Stenaropodites Martynov, 1928
  • †Sustaia Kukalová, 1958
  • †Sylvidelia Martynov, 1940
  • †Zdenekia Kukalová, 1958

References

1. Jakub Prokop; Wieslaw Krzemiński; Ewa Krzemińska; Thomas Hörnschemeyer; Jan-Michael Ilger; Carsten Brauckmann; Philippe Grandcolas; André Nel (2014). "Late Palaeozoic Paoliida is the sister group of Dictyoptera (Insecta: Neoptera)". Journal of Systematic Palaeontology. 12 (5): 601–622. doi:10.1080/14772019.2013.823468. S2CID 84407734.
2. Legendre, Frédéric; Nel, André; Svenson, Gavin J.; Robillard, Tony; Pellens, Roseli; Grandcolas, Philippe; Escriva, Hector (22 July 2015). "Phylogeny of Dictyoptera: Dating the Origin of Cockroaches, Praying Mantises and Termites with Molecular Data and Controlled Fossil Evidence". PLOS ONE. 10 (7): e0130127. Bibcode:2015PLoSO..1030127L. doi:10.1371/journal.pone.0130127. PMC 4511787. PMID 26200914.
3. Prokop, Jakub; Nel, André; Engel, Michael S. (2023). "Diversity, Form, and Postembryonic Development of Paleozoic Insects". Annual Review of Entomology. 68 (1): 401–429. doi:10.1146/annurev-ento-120220-022637.
4. Guan, Z.; Prokop, J.; Roques, P.; Lapeyrie, J.; Nel, A. (2016). "Revision of the enigmatic family Anthracoptilidae enlightens the evolution of Palaeozoic stem-dictyopterans". Acta Palaeontologica Polonica. 61 (1): 71–87. doi:10.4202/app.00051.2014.
5. André Nel; Markus J. Poschmann (2021). "Comparison of the recently described early Permian paoliid genus Permomertovia with the 'eoblattid' genera of the families Permulidae and Permotermopsidae". Palaeoentomology. 4 (1): 39–43. doi:10.11646/palaeoentomology.4.1.8.
6. Nel, A.; Garrouste, R.; Prokop, J. (2015). "The first African Anthracoptilidae (Insecta: Paoliida) near the Permian–Triassic boundary in Kenya". Zootaxa. 3925 (1): 145–150. doi:10.11646/zootaxa.3925.1.10.
7. Jakub Prokop; Jacek Szwedo; Jean Lapeyrie; Romain Garrouste; André Nel (2015). "New Middle Permian insects from Salagou Formation of the Lodève Basin in southern France (Insecta: Pterygota)". Annales de la Société Entomologique de France. New Series. 51 (1): 14–51. doi:10.1080/00379271.2015.1054645. S2CID 87501340.
8. Quispe, L.; Roques, P.; Garrouste, R.; Nel, A. (2021). "Carboniferous Blattinopsidae: revision of Klebsiella and new genus and species from Avion (Insecta, Paoliida)" (PDF). Historical Biology. 34 (3): 383–389. doi:10.1080/08912963.2021.1916817. S2CID 234869567.
9. Nel, A.; Garrouste, R.; Jouault, C. (2023). "New insects (Paoliida, Dictyoptera) from the Carboniferous outcrop of Tante Victoire in Var, France". Alcheringa: An Australasian Journal of Palaeontology. 47 (3): 305–314. doi:10.1080/03115518.2023.2258977.
10. Santos, A. A.; Hernández-Orúe, A.; Wappler, T.; Peñalver, E.; Diez, J. B.; Nel, A. (2022). "Late Carboniferous insects from the Iberian Peninsula: State of the art and new taxa". Palaeontographica Abteilung A. 326 (1–6): 1–27. doi:10.1127/pala/2022/0135. S2CID 251938467.