Notocupes is an extinct genus of medium-sized archostematan beetles from the Mesozoic Era of Eurasia, including over 50 described species. Historically, the genus was classified as a member of the family Ommatidae, but the presence of characters such as the horizontal mandibular cutting edge, separated procoxae and overlapping abdominal sternites indicate that the genus may have a closer affinity with the family Cupedidae.[1] Notocupes is considered to be a junior synonym of Zygadenia by Kirejtshuk (2020),[2] but other researchers suggest to reserve the genus Zygadenia as a form taxon for isolated elytra that probably belong to the genus Notocupes, while retaining Notocupes as a valid genus for complete body fossils.[3][4][5] Most species of Notocupes were described from compression fossils. An additional three species were described from Cenomanian-aged Burmese amber, which were treated as a separate genus, Echinocups, by Kirejtshuk (2020), but Li et al. (2023) consider Echinocups to be a junior synonym of Notocupes.[6] Notocupes has a flattened body, which may suggest that it occupied narrow habitats, such as living under bark. Some species had serrated/spined margins of the carapace, which may have served as a defense against predators, or served as camouflage to resemble bark.[1]
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Strelnikova & Yan (2023) advise splitting up the genus Notocupes into four genera: Notocupes, Rhabdocupes, Conexicoxa and the new genus Brachilatus.[7]
Ladinian (242 Ma to ~237 Ma)
- Tongchuan Formation, China
- N. shiluoensis (Hong, 1984) = Chengdecupes shiluoensis
Carnian (237 Ma to 227 Ma)
- Koldzat Formation, Kazakhstan
- N. oxypygus Ponomarenko, 1969
- Madygen Formation, Kyrgyzstan
- N. laticella Ponomarenko, 1969
- N. rostratus Ponomarenko, 1969
- N. tenuis Ponomarenko, 1969
Hettangian (201.3 Ma to 199.3 Ma)
- Dzhil Formation, Kyrgyzstan
- N. cellulosus Ponomarenko, 1969
- N. issykkulensis Ponomarenko, 1969
- N. kirghizicus Ponomarenko, 1969
- N. latus Ponomarenko, 1969
- N. sogutensis Ponomarenko, 1969
- Guanyintan Formation, China
- N. homorus (Lin, 1986) = Conexicoxa homora
Toarcian (182.7 Ma to 174.1 Ma)
- Guanyintan Formation, China
- N. lini Ponomarenko et al., 2012
- Sulyukta Formation, Kyrgyzstan
- N. crassus Ponomarenko, 1969
Bajocian (170.3 Ma to 168.3 Ma)
- Bakhar Formation, Mongolia
- N. brachycephalus Ponomarenko, 1994
- N. exiguus Ponomarenko, 1994
- N. longicollis Ponomarenko, 1994
Callovian (166.1 Ma to 163.5 Ma)
- Haifanggou Formation, China
- N. daohugouensis Li and Cai in Lin et al., 2023
- N. dischides Zhang, 1986
- N. jurassicus (Hong, 1983) = Chengdecupes jurassicus
- N. pingi Ponomarenko and Ren, 2010
- N. robustus Li and Cai in Lin et al., 2023
- N. spinosus Li and Cai in Lin et al., 2023
- Tyumen Formation, Russia
- N. patulus Ponomarenko, 1985
Oxfordian (163.5 Ma to 157.3 Ma)
- Karabastau Formation, Kazakhstan
- N. foersteri Ponomarenko, 1971
- N. lapidarius Ponomarenko, 1968
- N. nigromonticola Ponomarenko, 1968
- N. picturatus Ponomarenko, 1964
- N. pulcher Ponomarenko, 1968
Tithonian (152.1 Ma To 145.0 Ma)
- Shar Teeg, Mongolia
- N. brachycephalus Ponomarenko, 1994
- N. exiguus Ponomarenko, 1994
- Solnhofen, Germany
- N. reticulatus (Oppenheim, 1888) = Procarabus reticulatus
- N. tripartitus (Oppenheim, 1888) = Procarabus tripartitus
Barremian (129.4 Ma to 125.0 Ma)
- Jianshangou Formation, China
- La Huérguina Formation, Las Hoyas, Spain
- N. diazromerali (Soriano & Delclòs, 2006) = Zygadenia diazromerali
- N. longicoxa (Soriano & Delclòs, 2006) = Zygadenia longicoxa
- N. siniestri (Soriano & Delclòs, 2006) = Zygadenia siniestri
- N. viridis (Soriano & Delclòs, 2006) = Zygadenia viridis
- La Pedrera de Rúbies Formation, Las Hoyas, Spain
- N. martinclosas (Soriano & Delclòs, 2006) = Zygadenia martinclosas
- N. oculatus (Soriano & Delclòs, 2006) = Zygadenia oculata
- Ulan-Argalant Formation, Mongolia
- N. mongolicus Ponomarenko, 1994
Aptian (125.0 to ~113.0 Ma)
- Argun Formation, Russia
- N. semen (Ponomarenko, 2000) = Zygadenia semen
- N. sibiricus (Ponomarenko, 2000) = Zygadenia sibirica
- Baojiatun Formation, China
- N. baojiatunensis (Hong, 1992) = Chengdecupes baojiatunensis
- Dzun-Bain Formation, Mongolia
- N. dundulaensis Ponomarenko, 1994
- N. elegans Ponomarenko, 1994
- Jehol Group, China
- N. validus (Lin, 1976) = Sinocupes validus
- Khasurty Formation, Russia
- N. khasurtyiensis Strelnikova, 2019
- Laiyang Formation, China
- N. laiyangensis (Hong, 1990) = Forticupes laiyangensis
- N. ludongensis Wang and Liu, 1996
- N. tuanwangensis (Hong, 1990) = Picticupes tuanwangensis
- Shahai Formation, China
- N. kezuoensis (Hong, 1987) = Chengdecupes kezuoensis
- Yixian Formation, China
- N. alienus (Tan and Ren, 2006) = Ovatocupes alienus
- N. cyclodontus (Tan et al., 2006) = Amblomma cyclodonta
- N. epicharis (Tan et al., 2005) = Amblomma epicharis
- N. eumeurus (Tan et al., 2006) = Amblomma eumeura
- N. minisculus (Tan et al., 2006) = Amblomma miniscula
- N. porrectus (Tan et al., 2006) = Amblomma porrecta
- N. protensus (Tan et al., 2006) = Amblomma protensa
- N. psilatus (Tan et al., 2005) = Amblomma psilata
- N. rudis (Tan et al., 2006) = Amblomma rudis
- N. stabilis (Tan et al., 2006) = Amblomma stabilis
- Zaza Formation, Russia
- N. caudatus Ponomarenko, 1966
- N. excellens Ponomarenko, 1966
- N. vitimensis Ponomarenko, 1966
- Burmese amber, Myanmar
- N. denticollis Jiang et al., 2020 (sometimes placed in separate genus Echinocups)
- N. neli Tihelka et al., 2019 (sometimes placed in separate genus Echinocups)
- N. ohmkuhnlei Jarzembowski et al., 2019 (sometimes placed in separate genus Echinocups)
Turonian (93.9 to 89.8 Ma)
- Emanra Formation, Russia
- N. khetanensis Ponomarenko in Cromov et al., 1993
- Kzyl-Zhar Locality, Kazakhstan
- N. caducus Ponomarenko, 1969