Permocupedidae Explained

Permocupedidae is a 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.^[1] They are thought to have been xylophagous (wood eating), which is presumed to be the ancestral ecology of beetles.^[2]

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.^[3] However, other studies have recovered Talycupedidae as more closely related to crown-group beetles than to Permocupedidae.^[4]

• Afrocupes – South Africa: Western Cape (Kenmoore Farm), Whitehill Formation, Permian, Cisuralian, Sakmarian/Artinskian, −290.1–279.5 Ma.
  • Afrocupes firmae (Type species)
• Archicupes (syn Palaeocupes) – Russia, Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma
  • Archicupes jacobsoni (Type species)
  • "Archicupes" reichardti – considered indeterminate to genus by Kirejtshuk (2020)
• Cytocupoides – Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
  • Cytocupoides longatus (Type species)
• Cytocupes – Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
  • Cytocupes angustus (Type species)
• Eocupes – Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma).
  • Eocupes cellulosus
  • Eocupes lukjanovitshi (Type species)
• Frankencupes – Germany: Gambach am Main (Lower Franconia), Röt Formation Middle Triassic, Olenekian/Anisian, −247.2–242.0 Ma.
  • Frankencupes ultimus (Type species)
• Ichthyocupes – Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma; Kemerovo Region (Tykhta River, Sokolova, Kutsnetsk Basin), Permian, Lopingian, Wuchiapingian/Changhsingian (Tatarian, Severodvinian/Vyatkian), −259.0–252.3 Ma.
  • Ichthyocupes kuznetskiensis
  • Ichthyocupes skoki
  • Ichthyocupes tyzhnovi (Type species)
• Kaltanocoleus – Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
  • Kaltanocoleus pospelovi (Type species)
• Kaltanicupes – Russia: Kirov Region (Kityak), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma; Orenburg Region (Kargala), Permian, Guadalupian/Lopingian, Capitanian/Wuchiapingian (Tatarian, Sweverodvinian), −265.0–259.0 Ma; Kemerovo Region (Kaltan), 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 acutus
  • Kaltanicupes kargalensis
  • Kaltanicupes kitjakensis
  • Kaltanicupes richteri (Type species)
• Linicupes ^[5] – China: Anhui (Houdong, Yinping Mountain, SW Chaohu City), Permian, Guadalupian/Lopingian, Capitanian/Wuchiapingian (Tatarian, Severodvinian/Vyatkian), −265.0–259.0 Ma.
  • Linicupes yinpinensis (Type species)
• Maricoleus – Russia, Primorsky Krai (Cape, Novosilsky, Russky Island), Lower Permian, Cisuralian, Artinskian/Kungurian (Cisuralian/Biarmian, Kungurian/Kazanian), −279.5–272.5 Ma.
  • Maricoleus valentinae (Type species)
• Permocupes (syn Permocupoides) – Russia: Arkhangel’sk Region (Letopala River and Soyana), 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), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
  • Permocupes distinctus
  • Permocupes grandis
  • Permocupes latus
  • Permocupes semenovi (Type species)
  • Permocupes sojanensis
• Pintolla – 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 ^[6] (Type species)
• Protocupoides (syn Tomiocupes, Tricupes) – Russia: Vologda Region (Isady locality, Mutovino), 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), Permian, Lopingian, Wuchiapingian/Changhsingian (Tatarian, Severodvinian/Vyatkian), −259.0–252.3 Ma; Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
  • Protocupoides acer
  • Protocupoides carinatus
  • Protocupoides elongatus
  • Protocupoides esini
  • Protocupoides plavilstshikovi
  • Protocupoides sharovi
  • "Protocupoides" rohdendorfi ^[6] – 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 – Russia: Orenburg Region (Novo-Aleksandrovka), Permian, Guadalupian/Lopingian, Capitanian/Wuchiapingian (Tatarian, Severodvinian), −265.0–259.0 Ma.
  • Tatarocupes granulatus (Type species)
• Uralocupes – Russia: Belebeevo Formation Kirov Region (Kityak), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
  • Uralocupes major (Type species)

Notes and References

1. 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. en. 92. 4. 587–591. 10.1007/s12542-018-0426-4. 2018PalZ...92..587P . 133665417 . 0031-0220.
2. 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. Early evolution of beetles regulated by the end-Permian deforestation. eLife. 10. e72692. 10.7554/eLife.72692. 34747694 . 8585485 . 2050-084X. free.
3. Kirejtshuk. Alexander G.. 2020-02-17. Taxonomic Review of Fossil Coleopterous Families (Insecta, Coleoptera). Suborder Archostemata: Superfamilies Coleopseoidea and Cupedoidea. Geosciences. 10. 2. 73. 2020Geosc..10...73K. 10.3390/geosciences10020073. 2076-3263. free.
4. 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 . en . 69–96 . 10.1111/syen.12562 . 251171914 . 0307-6970. free .
5. A. G.. Ponomarenko. E. V.. Yan. Di-ying. Huang. 2014. New beetles (Coleoptera) from the terminal Middle Permian of China. Paleontological Journal. 48. 2. 191–200. 10.1134/S0031030114010109. 2014PalJ...48..191P .
6. Pinto. I.D.. 1987. Permian Insects from Paraná Basin, South Brazil, IV Coleoptera. Pesquisas. 19. 19 . 5–12. 10.22456/1807-9806.21678. free.