Protanisoptera

Protanisoptera Temporal range: Artinskian–Changhsingian PreꞒ Ꞓ O S D C P T J K Pg N
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Superorder:	Odonatoptera
Clade:	† Protanisoptera Carpenter, 1931 [1]
Families
See text.
Synonyms
Permanisoptera Martynov, 1931

Protanisoptera is an extinct clade of winged insects that lived during the Permian period. They were originally established as a suborder of Odonata (dragonflies and damselflies), but are now are generally considered a member of the superorder Odonatoptera. They are the sister group to the Discoidalia, a clade including modern Odonata. Their wings have a structure resembling the pterostigmata of modern Odonata due to convergent evolution, though it is crossed by the anterior radial vein (RA) unlike in true pterostigmata which are limited to the area between the costa (vein C) and RA. ^[2]

Classification

The following taxa are included in Protanisoptera. ^{[3] [4] [5]}

• Clade Protanisoptera Carpenter, 1931
  • Family †Pholidoptilidae Zalessky, 1931
    • †PholidoptilonZalessky, 1931 – Baitugan Formation, Russia, Middle Permian (Roadian) (known from a forewing)
  • Family †Polytaxineuridae Tillyard, 1935
    • †PolytaxineuraTillyard, 1935 – Warner's Bay, Croudace Bay Formation, Australia, Late Permian (Changhsingian) (known from a forewing and fragments of hindwings)
  • Clade †DitaxineuromorphaPritykina, 1980
    • Family †Permaeschnidae Martynov, 1931
      • †GondvanoptilonRösler et al., 1981 – Irati Formation, Brazil, Early Permian (Kungurian) (known from a nearly complete wing)
      • †PermaeschnaMartynov, 1931 – Iva-Gora Beds Formation, Russia, Middle Permian (Roadian) (known from wing fragments and a nearly complete wing)
    • Clade †DitaxineuridaBechly in Huguet et al., 2002
      • Family †Callimokaltaniidae Zalessky, 1955
        • †CallimokaltaniaZalessky, 1955 – Mitina Formation, Russia, Middle Permian (Roadian) (known from nearly complete wings)
      • Superfamily †DitaxineuroideaTillyard, 1926
        • Family †Ditaxineuridae Tillyard, 1926
          • †DitaxineuraTillyard, 1926 – Wellington Formation, Kansas, United States, Early Permian (Artinskian) (known from wing fragments and some complete wings)
          • †ProditaxineuraHuguet et al., 2002 – Chekarda site, Koshelevka Formation, Russia, Early Permian (Kungurian) (known from incomplete body fossils)
        • Family †Hemizygopteridae Zalessky, 1955
          • †HemizygopteronZalessky, 1955 – Barda locality, Koshelevka Formation, Russia, Early Permian (Kungurian) (known from wing fragments)
          • (?)†BansheepteronGarrouste et al., 2017 – Gonfaron, Pélitique Formation, France, Early–Middle Permian (Artinskian–Wordian) (known from a wing fragment)
  • incertae sedis
    • †DitaxineurellaMartynov, 1940 – Chekarda site, Koshelevka Formation, Russia, Early Permian (Kungurian) (known from wing fragments)

The families Camptotaxineuridae (from the Early Permian of Kansas) and Kaltanoneuridae (from the Late Permian of Russia) were originally included in Protanisoptera, but were excluded from the group by Huguet et al. (2002). The former family possibly belongs to the Palaeodictyoptera, while the latter is now included in the Protozygoptera. ^[4]

References

1. Carpenter, F.M. (1931). "The Lower Permian Insects of Kansas. Part 2. The Orders Paleodictyoptera, Protodonata and Odonata". American Journal of Science. 5. 21: 97–139. doi: 10.2475/ajs.s5-21.122.97 .
2. Nel, A.; Piney, B. (2023). "Odonatopteran Approaches to the Challenges of Flight: Convergence of Responses Subject to a Common Set of Morphological Constraints". In Bels, V.L.; Russell, A.P. (eds.). Convergent Evolution. Fascinating Life Sciences. Cham: Springer. doi:10.1007/978-3-031-11441-0_2.
3. Bechly, Günter (27 November 2007). "Phylogenetic Systematics of Odonata" . Retrieved 16 November 2025.
4. Huguet, A.; Nel, A.; Martínez-Delclòs, X.; Bechly, G.; Martins-Neto, R. (2002). "Preliminary phylogenetic analysis of the Protanisoptera (Insecta: Odonatoptera)". Geobios. 35 (5): 537–560. doi:10.1016/S0016-6995(02)00071-2 . Retrieved 16 November 2025.
5. Garrouste, R.; Lapeyrie, J.; Steyer, J.-S.; Giner, S.; Nel, A. (2017). "Insects in the Red Middle Permian of Southern France: first Protanisoptera (Odonatoptera) and new Caloneurodea (Panorthoptera), with biostratigraphical implications". Historical Biology. 30 (4): 546–553. doi:10.1080/08912963.2017.1301448.