Notocupes

Notocupes; Temporal range: Ladinian–Turonian PreꞒ Ꞓ O S D C P T J K Pg N
	Various specimens of Notocupes from the Middle Jurassic of China and mid-Cretaceous Burmese amber
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Coleoptera
Suborder:	Archostemata
Family:	incertae sedis
Genus:	†Notocupes; Ponomarenko, 1964
Synonyms
	Sinocupes Lin, 1976; Chengdecupes Hong, 1983; ?Conexicoxa Lin, 1986; Echinocups Kirejtshuk and Jarzembowski, 2020; Amblomma Tan et al., 2005; Ovatocupes Tan and Ren, 2006;

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]

Strelnikova & Yan (2023) advise splitting up the genus Notocupes into four genera: Notocupes, Rhabdocupes, Conexicoxa and the new genus Brachilatus.^[7]

Triassic species

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

Jurassic species

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

Cretaceous species

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

References

^ ^a ^b Li, Yan-Da; Tihelka, Erik; Yamamoto, Shûhei; Newton, Alfred F.; Xia, Fang-Yuan; Liu, Ye; Huang, Di-Ying; Cai, Chen-Yang (2023-08-22). "Mesozoic Notocupes revealed as the sister group of Cupedidae (Coleoptera: Archostemata)". Frontiers in Ecology and Evolution. 11. doi:10.3389/fevo.2023.1015627. ISSN 2296-701X.
^ Kirejtshuk, Alexander G. (2020). "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.
^ Strelnikova, O.D.; Yan, E.V. (2021). "Redescriptions of beetles of the Notocupes generic complex (Coleoptera: Archostemata: Ommatidae) from the Lower Cretaceous of Buryatia". Palaeoentomology. 4 (5): 499–514. doi:10.11646/palaeoentomology.4.5.15. S2CID 240527664.
^ Strelnikova, O.D.; Yan, E.V. (2023). "Redescriptions of the Triassic Notocupes beetles (Archostemata: Ommatidae) from Kyrgyzstan and South Kazakhstan". Palaeoentomology. 6 (2): 174–190. doi:10.11646/palaeoentomology.6.2.9. S2CID 258403180.
^ Soo Bin Lee; Gi Soo Nam; Yan-Da Li (2022). "A new species of Notocupes (Coleoptera: Archostemata) from the Lower Cretaceous (Albian) Jinju Formation in South Korea". Cretaceous Research. 140: 105357. Bibcode:2022CrRes.14005357L. doi:10.1016/j.cretres.2022.105357. S2CID 252147645.
^ Li, Y.-D.; Tihelka, E.; Newton, A. F.; Huang, D.-Y.; Cai, C.-Y. (2023). "New species of Notocupes (Coleoptera: Archostemata) from the Middle Jurassic Daohugou beds, with discussion on the generic circumscription". Palaeoentomology. 6 (4): 398–415. doi:10.11646/palaeoentomology.6.4.11.
^ Strelnikova, O.D.; Yan, E.V. (2023). "On splitting of the genus Notocupes (Coleoptera: Archostemata): new data on morphology and taxonomy". Far Eastern Entomologist. 488: 1–28. doi:10.25221/fee.488.1.

[:0-1] Li, Yan-Da; Tihelka, Erik; Yamamoto, Shûhei; Newton, Alfred F.; Xia, Fang-Yuan; Liu, Ye; Huang, Di-Ying; Cai, Chen-Yang (2023-08-22). "Mesozoic Notocupes revealed as the sister group of Cupedidae (Coleoptera: Archostemata)". Frontiers in Ecology and Evolution. 11. doi:10.3389/fevo.2023.1015627. ISSN 2296-701X.

[2] Kirejtshuk, Alexander G. (2020). "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.

[3] Strelnikova, O.D.; Yan, E.V. (2021). "Redescriptions of beetles of the Notocupes generic complex (Coleoptera: Archostemata: Ommatidae) from the Lower Cretaceous of Buryatia". Palaeoentomology. 4 (5): 499–514. doi:10.11646/palaeoentomology.4.5.15. S2CID 240527664.

[4] Strelnikova, O.D.; Yan, E.V. (2023). "Redescriptions of the Triassic Notocupes beetles (Archostemata: Ommatidae) from Kyrgyzstan and South Kazakhstan". Palaeoentomology. 6 (2): 174–190. doi:10.11646/palaeoentomology.6.2.9. S2CID 258403180.

[5] Soo Bin Lee; Gi Soo Nam; Yan-Da Li (2022). "A new species of Notocupes (Coleoptera: Archostemata) from the Lower Cretaceous (Albian) Jinju Formation in South Korea". Cretaceous Research. 140: 105357. Bibcode:2022CrRes.14005357L. doi:10.1016/j.cretres.2022.105357. S2CID 252147645.

[6] Li, Y.-D.; Tihelka, E.; Newton, A. F.; Huang, D.-Y.; Cai, C.-Y. (2023). "New species of Notocupes (Coleoptera: Archostemata) from the Middle Jurassic Daohugou beds, with discussion on the generic circumscription". Palaeoentomology. 6 (4): 398–415. doi:10.11646/palaeoentomology.6.4.11.

[7] Strelnikova, O.D.; Yan, E.V. (2023). "On splitting of the genus Notocupes (Coleoptera: Archostemata): new data on morphology and taxonomy". Far Eastern Entomologist. 488: 1–28. doi:10.25221/fee.488.1.

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