Notocupes

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. Notocupes is considered to be a junior synonym of Zygadenia by Kirejtshuk (2020), 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. 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. 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.

Strelnikova & Yan (2023) advise splitting up the genus Notocupes into four genera: Notocupes, Rhabdocupes, Conexicoxa and the new genus Brachilatus.

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

Hauterivian (132.9 Ma to 129.4 Ma)

 * Dabeigou Formation, China
 * N. lentus (Ren, 1995) = Tetraphalerus lentus

Barremian (129.4 Ma to 125.0 Ma)

 * Jianshangou Formation, China
 * N. laetus (Lin, 1976)
 * 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

Albian (~113.0 to 100.5 Ma)

 * Jinju Formation, South Korea
 * N. premeris Lee et al., 2022

Cenomanian (100.5 to 93.9 Ma)

 * 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