Permocupedidae

Permocupedidae Temporal range: Cisuralian–Anisian PreꞒ Ꞓ O S D C P T J K Pg N
Fossil and life restoration of Permocupes sojanensis
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Coleoptera
Suborder:	† Protocoleoptera
Superfamily:	† Permocupedoidea
Family:	† Permocupedidae Martynov, 1933 [1]
Genera
See text
Synonyms
Kaltanocoleidae Rohdendorf, 1961

Permocupedidae is an extinct family of Protocoleopteran stem group beetles. They first appeared during the Early Permian, and were one of the dominant groups of beetles during the Middle Permian. They became rare in the Late Permian, with only one species known from the Triassic, Frankencupes ultimus from the Middle Triassic (Anisian) Röt Formation of Germany. ^[2] They are thought to have been xylophagous (wood eating), which is presumed to be the ancestral ecology of beetles. ^[3]

Taxonomy

Kirejtshuk (2020) included the following genera in an expanded ( sensu lato ) family, with the traditional ( sensu stricto ) Permocupedidae and related Taldycupedidae as subfamilies Permocupedinae and Taldycupedinae respectively. ^[4] However, other studies have recovered Talycupedidae as more closely related to crown-group beetles than to Permocupedidae. ^[5]

• Afrocupes Geertsema & van den Heever, 1996 – South Africa: Western Cape (Kenmoore Farm), Whitehill Formation, Permian, Cisuralian, Sakmarian/Artinskian, −290.1–279.5 Ma.
  • Afrocupes firmaeGeertsema & van den Heever, 1996 (Type species)
• Archicupes Rohdendorf, 1961 (syn PalaeocupesRohdendorf, 1961) – Russia, Kemerovo Region (Kaltan), Mitina Formation, Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma
  • Archicupes jacobsoniRohdendorf, 1961 (Type species)
  • "Archicupes" reichardtiRohdendorf, 1961 – considered indeterminate to genus by Kirejtshuk (2020)
• Cytocupoides Ponomarenko, 1969 – Russia: Kemerovo Region (Kaltan), Mitina Formation, Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
  • Cytocupoides longatusPonomarenko, 1969 (Type species)
• Cytocupes Rohdendorf, 1961 – Russia: Kemerovo Region (Kaltan), Mitina Formation, Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
  • Cytocupes angustusRohdendorf, 1961 (Type species)
• Eocupes Rohdendorf, 1961 – Russia: Kemerovo Region (Kaltan), Mitina Formation, Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma).
  • Eocupes cellulosusPonomarenko, 1969
  • Eocupes lukjanovitshiRohdendorf, 1961 (Type species)
• Frankencupes Ponomarenko & Bashkuev, 2018 – Germany: Gambach am Main (Lower Franconia), Röt Formation Middle Triassic, Olenekian/Anisian, −247.2–242.0 Ma.
  • Frankencupes ultimusPonomarenko & Bashkuev, 2018 (Type species)
• Ichthyocupes Rohdendorf, 1961 – Russia: Kemerovo Region (Kaltan), Mitina Formation, Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma; Kemerovo Region, Gramoteinskaya Formation (Tykhta River, Sokolova, Kutsnetsk Basin), Permian, Lopingian, Wuchiapingian/Changhsingian (Tatarian, Severodvinian/Vyatkian), −259.0–252.3 Ma.
  • Ichthyocupes kuznetskiensisPonomarenko, 1969 – Mitina Formation
  • Ichthyocupes skoki(Rohdendorf, 1961) – Gramoteinskaya Formation
  • Ichthyocupes tyzhnoviRohdendorf, 1961 (Type species) – Mitina Formation
• Kaltanocoleus Rohdendorf, 1961 – Russia: Kemerovo Region (Kaltan), Mitina Formation, Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
  • Kaltanocoleus pospeloviRohdendorf, 1961 (Type species)
• Kaltanicupes Rohdendorf, 1961 – Russia: Kirov Region (Kityak), Belebeevo Formation, Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma; Orenburg Region (Kargala), Amanak Formation, Permian, Guadalupian/Lopingian, Capitanian/Wuchiapingian (Tatarian, Sweverodvinian), −265.0–259.0 Ma; Kemerovo Region (Kaltan), Mitina Formation, Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma. One undescribed species of this genus was recorded from Indian Barakar Formation of the Lower Permian.
  • Kaltanicupes acutusPonomarenko, 1963 – Belebeevo Formation
  • Kaltanicupes kargalensisPonomarenko, 1963 – Amanak Formation
  • Kaltanicupes kitjakensisPonomarenko, 1963 – Belebeevo Formation
  • Kaltanicupes richteriRohdendorf, 1961 (Type species) – Mitina Formation
• Linicupes Ponomarenko, Yan & Huang, 2014 ^[6] – China: Anhui (Houdong, Yinping Mountain, SW Chaohu City), Permian, Guadalupian/Lopingian, Capitanian/Wuchiapingian (Tatarian, Severodvinian/Vyatkian), −265.0–259.0 Ma.
  • Linicupes yinpinensisPonomarenko, Yan & Huang, 2014 (Type species)
• Maricoleus Shcherbakov in Shcherbakov, Makarkin, Aristov & Vasilenko, 2009 – Russia, Primorsky Krai (Cape, Novosilsky, Russky Island), Lower Permian, Cisuralian, Artinskian/Kungurian (Cisuralian/Biarmian, Kungurian/Kazanian), −279.5–272.5 Ma.
  • Maricoleus valentinaeShcherbakov in Shcherbakov, Makarkin, Aristov & Vasilenko, 2009 (Type species)
• Permocupes Martynov, 1933 (syn PermocupoidesMartynov, 1933) – Russia: Arkhangel’sk Region (Letopala River and Soyana), Iva-Gora Beds Formation, Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma; Udmurtia (Chepanikha, Rossokha River Valley), Permian, Guadalupian, Wordian (Biarmian, Urzhumian), −268.0–265.0 Ma: Tatarstan (Mendeleevsk District, Tikhie Gory), Baitugan Formation, Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
  • Permocupes distinctus(Martynov, 1933) – Baitugan Formation
  • Permocupes grandisPonomarenko, 1963 – Iva-Gora Beds Formation
  • Permocupes latusPonomarenko, 2013 – Chepanikha, Rossokha River Valley
  • Permocupes semenoviMartynov, 1933 (Type species) – Baitugan Formation
  • Permocupes sojanensisPonomarenko, 1963 – Iva-Gora Beds Formation
• Pintolla Kirejtshuk, 2020 – Brazil: Rio Grande do Sul (Road Cut on Road BR 290 near Minas do Leao), Irati Formation, Permian, Cisuralian, Artinskian/Kungurian (Cisuralian/Biarmian, Kungurian/Kazanian), −279.5–272.5 Ma.
  • Pintolla ponomarenkoi(Pinto, 1987) ^[7] (Type species)
• Protocupoides Rohdendorf, 1961 (syn TomiocupesRohdendorf, 1961, TricupesRohdendorf, 1961) – Russia: Vologda Region (Isady locality, Mutovino), Poldarsa Formation, Permian, Lopingian, Wuchiapingian/Changhsingian (Tatarian, Severodvinskian/Vyatkian), −259.0–254.0 Ma; Udmurtia (Chepanikha, Rossokha River Valley), Permian, Guadalupian, Wordian (Biarmian, Urzhumian), −268.0–265.0 Ma; Kemerovo Region (Sokolova, Kutsnetsk Basin), Gramoteinskaya Formation, Permian, Lopingian, Wuchiapingian/Changhsingian (Tatarian, Severodvinian/Vyatkian), −259.0–252.3 Ma; Kemerovo Region (Kaltan), Mitina Formation, Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
  • Protocupoides acer(Rohdendorf, 1961) – Gramoteinskaya Formation
  • Protocupoides carinatus(Rohdendorf, 1961) – Gramoteinskaya Formation
  • Protocupoides elongatusPonomarenko, 2013 – Chepanikha, Rossokha River Valley
  • Protocupoides esiniPonomarenko, 2013 – Poldarsa Formation
  • Protocupoides plavilstshikoviRohdendorf, 1961 – Mitina Formation
  • Protocupoides sharovi(Rohdendorf, 1961) – Mitina Formation
  • "Protocupoides" rohdendorfiPinto, 1987 ^[7] – Brazil (Rio Grande do Sul (Road Cut on Road BR 290 near Minas do Leao), Irati Formation, Permian, Kungurian, −279.5–272.5 Ma; considered indeterminate to genus by Kirejtshuk (2020)
• Tatarocupes Ponomarenko, 2004 – Russia: Orenburg Region (Novo-Aleksandrovka), Permian, Guadalupian/Lopingian, Capitanian/Wuchiapingian (Tatarian, Severodvinian), −265.0–259.0 Ma.
  • Tatarocupes granulatusPonomarenko, 2004 (Type species)
• Uralocupes Ponomarenko, 1969 – Russia: Belebeevo Formation Kirov Region (Kityak), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
  • Uralocupes major(Ponomarenko, 1963) (Type species)

References

1. Martynov, A.V. (1933). Пермские ископаемые насекомые Архангельского края. Часть II. Сетчатокрылые, вислокрылые и жуки, с приложением описания двух новых жуков из Тихих Гор [Permian fossil insects from the Arkhangelsk District. Part II. Neuroptera, Megaloptera and Coleoptera with the description of two new beetles from Tikhie Gory](DjVu). Trudy Paleozoologicheskogo Instituta Akademii Nauk SSSR (in Russian and English). 2: 63–96.
2. Ponomarenko, Alexandr G.; Bashkuev, Alexei S. (December 2018). "First Triassic record of the beetle family Permocupedidae (Insecta: Coleoptera): a peculiar example of a Lazarus taxon" . PalZ. 92 (4): 587–591. Bibcode:2018PalZ...92..587P. doi:10.1007/s12542-018-0426-4. ISSN   0031-0220. S2CID   133665417.
3. Zhao, Xianye; Yu, Yilun; Clapham, Matthew E; Yan, Evgeny; Chen, Jun; Jarzembowski, Edmund A; Zhao, Xiangdong; Wang, Bo (2021-11-08). Perry, George H; Fikacek, Martin (eds.). "Early evolution of beetles regulated by the end-Permian deforestation". eLife. 10: e72692. doi: 10.7554/eLife.72692 . ISSN   2050-084X. PMC   8585485 . PMID   34747694.
4. Kirejtshuk, Alexander G. (2020-02-17). "Taxonomic Review of Fossil Coleopterous Families (Insecta, Coleoptera). Suborder Archostemata: Superfamilies Coleopseoidea and Cupedoidea". Geosciences. 10 (2): 73. Bibcode:2020Geosc..10...73K. doi: 10.3390/geosciences10020073 . ISSN   2076-3263.
5. Boudinot, Brendon Elias; Yan, Evgeny Viktorovich; Prokop, Jakub; Luo, Xiao-Zhu; Beutel, Rolf Georg (2022-07-28). "Permian parallelisms: Reanalysis of †Tshekardocoleidae sheds light on the earliest evolution of the Coleoptera". Systematic Entomology. 48: 69–96. doi: 10.1111/syen.12562 . ISSN   0307-6970. S2CID   251171914.
6. Ponomarenko, A. G.; Yan, E. V.; Huang, Di-ying (2014). "New beetles (Coleoptera) from the terminal Middle Permian of China". Paleontological Journal. 48 (2): 191–200. Bibcode:2014PalJ...48..191P. doi:10.1134/S0031030114010109.
7. Pinto, I.D. (1987). "Permian Insects from Paraná Basin, South Brazil, IV Coleoptera". Pesquisas. 19 (19): 5–12. doi: 10.22456/1807-9806.21678 .