# 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. They are thought to have been xylophagous (wood eating), which is presumed to be the ancestral ecology of beetles.

## 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. However, other studies have recovered Talycupedidae as more closely related to crown-group beetles than to Permocupedidae.
* 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)
** "Archicupes" reichardti Rohdendorf, 1961 – considered indeterminate to genus by Kirejtshuk (2020)
* 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, 1961 – Russia: 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, 1961 – Russia: 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, 1961 – 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 Ponomarenko, 1963
** Kaltanicupes kargalensis Ponomarenko, 1963
** Kaltanicupes kitjakensis Ponomarenko, 1963
** Kaltanicupes richteri Rohdendorf, 1961 (Type species)
* Linicupes Ponomarenko, Yan & Huang, 2014 – 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, 1933 (syn Permocupoides Martynov, 1933) – 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 (Martynov, 1933)
** Permocupes grandis Ponomarenko, 1963
** Permocupes latus Ponomarenko, 2013
** Permocupes semenovi Martynov, 1933 (Type species)
** 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 sharovi (Rohdendorf, 1961)
** "Protocupoides" 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)