Permocupedidae

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]

Permocupedidae
Temporal range:
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
Genera

See text

Synonyms
  • Taldycupedidae

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]

Subfamily Permocupedinae Martynov, 1932

  • Afrocupes Geertsema & van den Heever, 1996 South Africa: Western Cape (Kenmoore Farm), Whitehill Formation, Permian, Cisuralian, Sakmarian/Artinskian, −290.1–279.5 Ma.
    • Afrocupes firmae Geertsema & van den Heever, 1996 (Type species)
  • Archicupes Rohdendorf, 1961 (syn Palaeocupes Rohdendorf, 1961) Russia, Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma
    • Archicupes jacobsoni Rohdendorf, 1961 (Type species)
  • Cytocupoides Ponomarenko, 1969 Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
    • Cytocupoides longatus Ponomarenko, 1969 (Type species)
  • Cytocupes Rohdendorf, 1961Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
    • Cytocupes angustus Rohdendorf, 1961 (Type species)
  • Eocupes Rohdendorf, 1961Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma).
    • Eocupes cellulosus Ponomarenko, 1969
    • Eocupes lukjanovitshi Rohdendorf, 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 ultimus Ponomarenko & Bashkuev, 2018 (Type species)
  • Ichthyocupes Rohdendorf, 1961 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 Ponomarenko, 1969
    • Ichthyocupes skoki (Rohdendorf, 1961)
    • Ichthyocupes tyzhnovi Rohdendorf, 1961 (Type species)
  • Kaltanocoleus Rohdendorf, 1961. Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5–268.0 Ma.
    • Kaltanocoleus pospelovi Rohdendorf, 1961 (Type species)
  • Kaltanicupes Rohdendorf, 1961Russia: 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 Ponomarenko, 1963
    • Kaltanicupes kargalensis Ponomarenko, 1963
    • Kaltanicupes kitjakensis Ponomarenko, 1963
    • Kaltanicupes richteri Rohdendorf, 1961 (Type species)
    • Kaltanicupes reichardti Rohdendorf, 1961
  • Linicupes Ponomarenko, Yan & Huang, 2014[4] China: Anhui (Houdong, Yinping Mountain, SW Chaohu City), Permian, Guadalupian/Lopingian, Capitanian/Wuchiapingian (Tatarian, Severodvinian/Vyatkian), −265.0–259.0 Ma.
    • Linicupes yinpinensis Ponomarenko, 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 valentinae Shcherbakov in Shcherbakov, Makarkin, Aristov & Vasilenko, 2009 (Type species)
  • Permocupes Martynov, 1932 (syn Permocupoides Martynov, 1932) 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 (Ponomarenko, 1963
    • Permocupes latus Ponomarenko, 2013
    • Permocupes semenovi Martynov, 1932
    • Permocupes sojanensis (Ponomarenko, 1963
  • 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) (Type species)
  • Protocupoides Rohdendorf, 1961 (syn Tomiocupes Rohdendorf, 1961, Tricupes Rohdendorf, 1961) 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 (Rohdendorf, 1961)
    • Protocupoides carinatus (Rohdendorf, 1961)
    • Protocupoides elongatus Ponomarenko, 2013
    • Protocupoides esini Ponomarenko, 2013
    • Protocupoides plavilstshikovi Rohdendorf, 1961
    • Protocupoides sharoviRohdendorf, 1961
    • “Permocupes” rohdendorfi Pinto, 1987 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 granulatus Ponomarenko, 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)

Subfamily Taldycupedinae Rohdendorf, 1961

  • Afrotaldycupes Kirejtshuk, 2020 Beaufort Group, South Africa, Late Permian (Changhsingian)
    • Afrotaldycupes africanus (Ponomarenko, 2005) (Type species)
    • Afrotaldycupes lidgettoniensis (Ponomarenko, 2005)
  • Mesothoris Tillyard, 1916 Blackstone Formation, Australia Late Triassic (Norian)
    • Mesothoris clathrata Tillyard, 1916 (Type species)
    • Mesothoris grandis Dunstan, 1923
    • Mesothoris punctomarginum Dunstan, 1923
  • Proterocupes Ponomarenko, 2015 (syn Lobanovia Kirejtshuk, 2020)
    • Proterocupes major Ponomarenko, 2015 Vokhma Formation, Vologda, Russia, Late Permian (Changhsingian)
    • Proterocupes nedubrovensis Ponomarenko, 2015 Vokhma Formation, Vologda, Russia, Late Permian (Changhsingian)
    • Proterocupes permiana (Ponomarenko, 2015) (Type species) Poldarsa Formation, Vologda, Russia, Late Permian (Wuchiapingian)
  • Simmondsia Dunstan, 1923 Blackstone Formation, Australia Late Triassic (Norian)
    • Simmondsia cylindrica Dunstan, 1923
    • Simmondsia subpiriformis Dunstan, 1923 (Type species)
  • Taldycupes Rohdendorf, 1961 (syn Cryptocupes Rohdendorf, 1961, Taldycupidium Rohdendorf, 1961)
    • Taldycupes bergi (Rohdendorf, 1961) Gramoteinskaya Formation, Kemerovo Oblast, Russia, Late Permian
    • Taldycupes cellulosus Ponomarenko, 2013 Poldarsa Formation, Vologda, Russia, Late Permian (Wuchiapingian)
    • Taldycupes khalfini Rohdendorf, 1961 Gramoteinskaya Formation, Kemerovo Oblast, Russia, Late Permian
    • Taldycupes moltshanovi Rohdendorf, 1961 Gramoteinskaya Formation, Kemerovo Oblast, Russia, Late Permian
    • Taldycupes pingi Ponomarenko et al. 2014 Yinping Formation, Anhui, China, Middle Permian (Capitanian)
    • Taldycupes reticulatus Ponomarenko, 1969 Akkolka Formation, Kazakhstan, Late Permian (Changhsingian)
    • Taldycupes rosanovi Ponomarenko, 2006 Limptekon Formation, Krasnoyarsk, Russia, Late Permian (Changhsingian)
    • Taldycupes vasjuchitshevi Rohdendorf, 1961 Gramoteinskaya Formation, Kemerovo Oblast, Russia, Late Permian
  • Tecticupes Rohdendorf, 1961 (syn Stegocupes Rohdendorf, 1961
    • Tecticupes heckeri Rohdendorf, 1961 (Type species) Kazankovo-Markinskaya Formation, Kemerovo, Russia, Middle Permian (Wordian) Mitina Formation, Kemerovo, Russia, Middle Permian (Roadian)
    • Tecticupes indistinctus Rohdendorf, 1961 Kazankovo-Markinskaya Formation, Kemerovo, Russia, Middle Permian (Wordian)
    • Tecticupes martynovi Ponomarenko, 2013 Amanak Formation. Orenburg, Middle Permian (Capitanian)
  • Tychticupes Rohdendorf, 1961 Gramoteinskaya Formation, Kemerovo Oblast, Russia, Late Permian
    • Tychticupes beljanini Rohdendorf, 1961
    • Tychticupes radtshenkoi Rohdendorf, 1961
    • Tychticupes ragozini Rohdendorf 1961
    • "Tychticupes ussovi" Rohdendorf 1961 (nomen dubium)
  • Tychticupoides Rohdendorf, 1961 Gramoteinskaya Formation, Kemerovo Oblast, Russia, Late Permian
    • Tychticupoides grjazevi Rohdendorf, 1961
  • Uskaticupes Rohdendorf, 1961 (nomen dubium) Gramoteinskaya Formation, Kemerovo Oblast, Russia, Late Permian

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. 92 (4): 587–591. doi:10.1007/s12542-018-0426-4. ISSN 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 (eds.). "Early evolution of beetles regulated by the end-Permian deforestation". eLife. 10: e72692. doi:10.7554/eLife.72692. ISSN 2050-084X.
  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. Bibcode:2020Geosc..10...73K. doi:10.3390/geosciences10020073. ISSN 2076-3263.
  4. A. G. Ponomarenko, E. V. Yan and Di-ying Huang (2014). "New beetles (Coleoptera) from the terminal Middle Permian of China". Paleontological Journal 48 (2): 191–200. doi:10.1134/S0031030114010109.
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