Paleobiota of the Koshelevka Formation

The Koshelevka Formation is a Permian formation in Russia containing the Tshekarda lagerstätte, which preserves over 200 insect species.[1]

Insects

Holometabola

Holometabolans
Genus Species Higher taxon Notes Images
Sylvacoleus S. richteri, S. sharovi Tshekardocoleidae[2] Has a relatively long abdomen, named after the Sylva River[3]
Sylvacoleodes S. admirandus Tshekardocoleidae Resembles Sylvacoleus, but has a narrower head[4]
Tshekardocoleus T. magnus Tshekardocoleidae[2] Type genus of the family, wing venation quite similar to the hypothetical ancestral beetle[3]
Cavalarva C. caudata Holometabola incertae sedis Shares characteristics with larvae from numerous holometabolan orders[1]

Paraneoptera

Paraneopterans
Genus Species Higher taxon Notes Images
Graticladus G. apiatus[1] Ischnoneuridae (Hypoperlida) Placed in Hypoperlida, which was reclassified as stem-Paraneoptera in 2017[5]
Tshekardoclardus T. sparsus[1] Tococladidae Placed in Hypoperlida, which was reclassified as stem-Paraneoptera in 2017[5]

Polyneoptera (excluding Notoptera)

Non-Notopteran Polyneopterans
Genus Species Higher taxon Notes Images
Arachnophasma A. scurra Permophasmatidae Oldest stick insect known, has very long legs and a short body[1]
Uraloblatta U. insignis Archimylacridae ("Eoblattodea")[6] Has a relatively long oviscapt[7]

Notoptera

Notopterans
Genus Species Higher taxon Notes Images
Artinska A. infigurabilis[8] Lemmatophoridae
Bardapteron B. ovale Atactophlebiidae Formerly placed with paleodictyopterans, known from the less diverse Barda locality as opposed to Chekarda[8]
Chelopterum C. ultimum Chelopteridae[1]
Cucullistriga C. cucullata Idelinellidae Named after the hood-like pronotum covering its head[9]
Culiciforma C. formosa Lemmatophoridae Similar to Megorkhosa[8]
Czekardia C. blattoides Eoblattida incertae sedis Preserves stripes and spots on its tergites, only known from a nymph[10]
Depressopterum D. bardum Liomopteridae[8]
Euryptilodes E. commatulus Euryptilonidae Known from wings and exuviae of nymphs[8]
Iblatta I. attrepida Eoblattida incertae sedis Only known from a nymph, has quite short legs and a small head[10]
Jubala J. pectinata Euryptilonidae Formerly placed within Sylvardembia[1]
Khosarophlebia K. sylvaensis Liomopteridae Formerly known from a wing fragment, but had a complete specimen discovered later[8]
Kishertia K. tricubitalis Pinideliidae Distinguished from Pinidelia by wing venation and size[8]
Koshelevka K. megakhosaroides Doubraviidae Formerly included in Cerasopterum[11]
Kungurocauda K. spinosa Sylviaphlebiidae Second-smallest member of the family behind Sylvophenoptera[8]
Kungurolioma K. cancellata Liomopteridae
Lemmatonympha L. gracilis, L. gracilissima Lemmatophoridae Identifiable to family based on its tarsi and cerci[12]
Liomopterella L. kungurica Liomopteridae[8]
Micaidelia M. minutissima Ideliidae Similar to Anaidelia[8]
Neprotembia N. truncata Permulidae[13] Similar to Neraphidia[8]
Neraphidia N. mitis, N. rigida Neraphidiidae[14] Formerly only known from part of a forewing, but a complete specimen was discovered later[8]
Novokshonovus N. ignoratus Atactophlebiidae Related to Kirkorella[1]
Ornaticosta O. novokshonovi Liomopteridae[8]
Parakhosara P. coalita Megakhosaridae[8]
Parasheimia P. truncata, P. rotundata Sheimiidae Larger than Sheimia and has a wider pronotum[8]
Parastenaropodites P. stirps Mesorthopteridae[1]
Parasylvaella P. umbra, P. minor Liomopteridae[8]
Parasylviodes P. tetracladus Liomopteridae[8]
Paratillyardembia P. sepicolorata Protembiidae[15] Has an unusual pronotum[16]
Parmaptera P. permiana Parmapteridae Placed within its own monotypic family[1]
Pectinokhosara P. sylvardembiodes Megakhosaridae Has elongated wings[8]
Permedax P. effertus Tillyardembiidae Differs from all other known grylloblattid nymphs by its elongate body,[8] but this is also a character of adult tillyardembiids[12]
Permula P. tshekardensis, P. minor Permulidae Formerly placed within Sojanopermula,[8] with Permula itself being a replacement name for Allicula.[13]
Permoshurabia P. kungurica Geinitziidae One of the earliest geinitziids[1]
Permostriga P. augustalis Idelinellidae[9]
Pseudosheimia P. caudata Sheimiidae Has longer cerci than Sheimia[8]
Scutistriga S. scutata Idelinellidae Named after its shield-like pronotum[9]
Sheimia S. tshekardensis Sheimiidae Bears larger eyes than other Sheimia species[8]
Strigulla S. cuculiophoris Idelinellidae Formerly placed within Euryptilon[9]
Suksunus S. bicodex Cacurgidae Similar to Kochopteron[11]
Sylvaclinicus S. echinatus Grylloblattida incertae sedis Only known from nymphs[8]
Sylvafossor S. forcipatus Probnidae Has unusual earwig-like cerci[8]
Sylvamicropteron S. harpax Grylloblattida incertae sedis An unusual micropterous grylloblattid[8]
Sylvaphlebia S. tuberculata, S. fucata Sylvaphlebiidae Has a pigmented wing strip[8]
Sylvaprisca S. gravis, S. forta, S. focaleata[8] Lemmatophoridae Named after the lemmatophorid genus Paraprisca[17]
Sylvardembia S. matura, S. tamaena Sylvardembiidae[16]
Sylvastriga S. miranda[8] Idelinellidae
Sylvictor S. major Liomopteridae One of the largest notopterans from Koshelevka[8]
Sylviodes S. perloides Mesorthopteridae[18] Formerly placed in Ideliidae[8]
Sylvonympha S. tshekardensis Grylloblattida incertae sedis Unusually for a grylloblattid, the larva has gills[12]
Sylvophenoptera S. perlongata, S. fimbriata Sylvaphlebiidae Resembles Phenopterum but has a longer ovipositor[8]
Tillyardembia T. antennaeplana, T. ravisedorum[19] Tillyardembiidae Most common insects from Koshelevka alongside the earliest insects with preserved pollen attached to them[20]
Tshekardanympha T. lienterica, T. bardensis Grylloblattida incertae sedis Split from Tshekardites[12]
Tshekardeigma T. rasnitsyni Epideigmatidae[21]
Tshekardelia T. media Ideliidae One of the first ideliids known from a well-preserved body[22]
Tshekardites T. comitialis Grylloblattida incertae sedis[8] First established as a nymphal form taxon, before all other species were split out[12]
Tshekardomina T. maculatus, T. imbecilla, T. imbecillissima, T. subincurvata[8] Tshekardominidae (Grylloblattida) Type specimen preserves several eggs inside the abdomen, rest of specimens are all female (possibly suggesting flightless males?)[23]
Tshekardophlebia T. capitata Sylvaphlebiidae Resembles Sylvaphlebia but has a larger head[8]
Tshekardushka T. artenatis Reculida incertae sedis Only known from a nymph. Preserves an oval impression, likely representing a bolus of inorganic material[10]
Uralioma U. maxima Liomopteridae Incompletely preserved, may belong to a separate genus[8]
Uraloprisca U. uralica Lemmatophoridae Formerly included in Paraprisca[24]
Visherifera V. sylvaensis[8] Visheriferidae Originally described from a late Permian locality in the Solikamsk Formation[25]

Misc. Arthropoda

Miscellaneous arthropods
Genus Species Higher taxon Notes Images
Permarachne P. novokshonovi Uraraneida[26] Previously identified as a spider[27]
Permocrassacus P. novokshonovi Scolopendromorpha First described Permian centipede[28]

Plants

Seed plants

Conifers

Conifers
Genus Species Higher taxon Notes Images
Cyparissidium C. appressum Voltziales Formerly placed within Walchia[29]
Kungurodendron K. sharovii Voltziales Similar to Walchia, although differs in having seed stalks[29]
Taxodiella T. bardaeana Voltziales Formerly placed within Walchia, synonymous with several species[29]

References

  1. ^ a b c d e f g h i j k Aristov, Danil S.; Rasnitsyn, Alexandr P. (2015). "New insects from the Kungurian of Tshekarda fossil site in Permian Territory of Russia" (PDF). Russian Entomological Journal. 24 (1): 17–35. doi:10.15298/rusentj.24.1.03.
  2. ^ a b Ponomarenko, A. G. (2000). "New Beetles from the Permian of European Russia". Paleontological Journal. 34 (Suppl. 3): S312 – S316.
  3. ^ a b Ponomarenko, A. G. (1963). Палеозойские жуки Cupedidea европейской части СССР. Палеонтологический Журнал (in Russian). 1963 (1): 70–85.
  4. ^ Ponomarenko, A. G. (1969). Историческое Развитие Жесткокрылых-Архостемат (PDF). Proceedings of the Paleontological Institute (in Russian). Vol. 125. Moscow: Nauka. p. 56.
  5. ^ a b Prokop, Jakub; Pecharová, Martina; Garrouste, Romain; Beattie, Robert; Chintauan-Marquier, Ioana C.; Nel, André (December 2017). "Redefining the extinct orders Miomoptera and Hypoperlida as stem acercarian insects". BMC Evolutionary Biology. 17 (1) 205. Bibcode:2017BMCEE..17..205P. doi:10.1186/s12862-017-1039-3. PMC 5574135.
  6. ^ Ponomarovya, G. Yu.; Novokshonov, V. G.; Naugolnykh, S. V. (1998). Чекарда - местонахождение пермских ископаемых растений и насекомых [Chekarda - the locality of Permian fossil plants and insects] (PDF) (in Russian). Perm: Издательство Пермского университета. pp. 25–54. ISBN 5-8241-0165-5. Retrieved 15 November 2025.
  7. ^ Zalessky, Georges (1939). "Sur une nouvelle Blatte permienne portant un oviscapte". Annales de la Société géologique du Nord. 64: 85–94.
  8. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj Aristov, D. S. (2004). "The Fauna of Grylloblattid Insects (Grylloblattida) of the Lower Permian Locality of Tshekard". Paleontological Journal. 38 (2): S80 – S145.
  9. ^ a b c d Aristov, D. S.; Rasnitsyn, A. P. (January 2012). "Revision of the family Idelinellidae, with a review of the Permian Eoblattida (Insecta)". Paleontological Journal. 46 (1): 49–60. Bibcode:2012PalJ...46...49A. doi:10.1134/S0031030112010030.
  10. ^ a b c Aristov, D. S. (March 2018). "New and Little-known Nymphs of Gryllones (Insecta) from the Lower Permian of Russia". Paleontological Journal. 52 (2): 137–141. Bibcode:2018PalJ...52..137A. doi:10.1134/S0031030118020028.
  11. ^ a b Aristov, D. S. (November 2015). "Classification of the order eoblattida (insecta: Blattidea) with description of new taxa". Far Eastern Entomologist. 301: 1–56.
  12. ^ a b c d e Aristov, D. S.; Novokshonov, V. G.; Pan’kov, N. N. (January 2006). "Taxonomy of the fossil grylloblattid nymphs (Insecta: Grylloblattida)". Paleontological Journal. 40 (1): 79–89. Bibcode:2006PalJ...40...79A. doi:10.1134/S0031030106010096.
  13. ^ a b Aristov, Daniil; Storozhenko, Sergey (24 September 2011). "Review of the Permian family Permulidae nomen novum pro Aliculidae Storozhenko, 1997 (Grylloblattida)". ZooKeys (130): 111–130. Bibcode:2011ZooK..130..111A. doi:10.3897/zookeys.130.1489. PMC 3260753. PMID 22259270.
  14. ^ Aristov, D. S. (May 2014). "Classification of the order Cnemidolestida (Insecta; Perlidea) with description of new taxa". Far Eastern Entomologist. 277: 1–46.
  15. ^ Aristov, Danil S.; Rasnitsyn, Alexandr P. (2011). "A review of the family Protembiidae (Insecta: Eoblattida)" (PDF). Russian Entomological Journal. 20 (2): 119–127. doi:10.15298/rusentj.20.2.02.
  16. ^ a b Aristov, D. S. (September 2000). "New insects of the order Grylloblattida (Insecta) from the Lower Permian of the Middle Urals". Paleontological Journal. 34 (5): 48–50.
  17. ^ Aristov, D. S. (May 2001). "A new representative of the Lemmatophoridae (Insecta, Grylloblattida) from the Lower Permian of the Ural (Russia)". Neues Jahrbuch für Geologie und Paläontologie. 2001 (5): 257–264. doi:10.1127/njgpm/2001/2001/257.
  18. ^ Storozhenko, S.Yu.; Aristov, D.S. (7 September 2021). "The latest record of the genus Belmophenopterum (Eoblattida: Mesorthopteridae) from the Middle Triassic of Kyrgyzstan". Far Eastern Entomologist: 31–36. doi:10.25221/fee.437.6.
  19. ^ Aristov, Danil S.; Rasnitsyn, Alexandr P. (2009). "The family Tillyardembiidae Zalessky, 1938 and the system of the plecopteroid insects". Russian Entomological Journal. 18 (4): 257–264.
  20. ^ Khramov, Alexander V.; Foraponova, Tatiana; Węgierek, Piotr (March 2023). "The earliest pollen-loaded insects from the Lower Permian of Russia". Biology Letters. 19 (3) 20220523. doi:10.1098/rsbl.2022.0523. PMC 9975653. PMID 36855855.
  21. ^ Aristov, D. S. (July 2011). "Revision of the family Epideigmatidae (Insecta: Grylloblattida)". Paleontological Journal. 45 (4): 397–405. Bibcode:2011PalJ...45..397A. doi:10.1134/S0031030111040034.
  22. ^ Aristov, D. S. (2002). "Grylloblattids of the Family Ideliidae (Insecta: Grylloblattida) from the Lower Permian of the Ural Mountains". Paleontological Journal. 36 (1): 44–47.
  23. ^ Novokshonov, V. G.; Aristov, D. S. (November 2002). "New and little-known Permian insects (Insecta : Grylloblattida; Orthoptera) from the Chekarda locality, Central Ural Mountains". Paleontological Journal. 6 (6): 73–77.
  24. ^ Aristov, D. S. (May 2009). "New grylloblattids of the family Lemmatophoridae (Insecta: Grylloblattida) from the Permian of Russia". Paleontological Journal. 43 (3): 272–276. Bibcode:2009PalJ...43..272A. doi:10.1134/S0031030109030046.
  25. ^ Aristov, D. S. (2002). "New Insects from the Late Permian of the Ural Mountains". Paleontological Journal. 36 (2): 157–160.
  26. ^ Selden, Paul A.; Shear, William A.; Sutton, Mark D. (30 December 2008). "Fossil evidence for the origin of spider spinnerets, and a proposed arachnid order". Proceedings of the National Academy of Sciences. 105 (52): 20781–20785. doi:10.1073/pnas.0809174106. PMID 19104044.
  27. ^ Eskov, K. Y.; Selden, P. A. (2005). "First record of spiders from the Permian period (Araneae: Mesothelae)" (PDF). Bulletin of the British Arachnological Society. 13 (4): 111–116.
  28. ^ Khramov, Alexander; Shear, William; Mercurio, Randy; Kopylov, Dmitry (2018). "The first Permian centipedes from Russia". Acta Palaeontologica Polonica. 63. doi:10.4202/app.00482.2018.
  29. ^ a b c Meyen, Sergei V. (May 1997). "Permian conifers of Western Angaraland". Review of Palaeobotany and Palynology. 96 (3–4): 351–447. Bibcode:1997RPaPa..96..351M. doi:10.1016/S0034-6667(96)00046-2.
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