Paoliida

Extinct order of insects

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

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]

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

^ ^a ^b 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.
^ 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.
^ 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.
^ 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.
^ 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.
^ 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.
^ 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.
^ 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.
^ 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.
^ 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.

Insect orders

Kingdom: Animalia
Phylum: Arthropoda
(unranked): Pancrustacea
Subphylum: Hexapoda

Extant

Monocondylia	Archaeognatha (jumping bristletails)

D
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d
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Zygentoma (silverfish, firebrats)

P
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Palaeoptera

Ephemeropteroidea	Ephemeroptera (mayflies)
Odonatoptera	Odonata (dragonflies, damselflies)

N
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p
t
e
r
a

Polyneoptera

Plecoptera (stoneflies) Dermaptera (earwigs) Embioptera (webspinners) Phasmatodea (stick and leaf insects) Orthoptera (crickets, wetas, grasshoppers, locusts) Zoraptera (angel insects)
Notoptera (Xenonomia)	Grylloblattodea (ice-crawlers) Mantophasmatodea (gladiators)
Dictyoptera	Blattodea (cockroaches, termites) Mantodea (mantises)

E
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t
a
b
o
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a

Paraneoptera *	Psocodea (barklice, lice) Thysanoptera (thrips) Hemiptera (cicadas, aphids, true bugs)

H
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m
e
t
a
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Hymenoptera (sawflies, wasps, ants, bees)

Neuropteroidea

Coleopterida	Strepsiptera (twisted-winged parasites) Coleoptera (beetles)
Neuropterida	Raphidioptera (snakeflies) Megaloptera (alderflies, dobsonflies, fishflies) Neuroptera (net-winged insects: lacewings, mantidflies, antlions)

Panorpida
(Mecopterida)

Antliophora	Mecoptera (scorpionflies) + Siphonaptera (fleas) Diptera (gnats, mosquitoes, flies)
Amphiesmenoptera	Trichoptera (caddisflies) Lepidoptera (moths, butterflies)