# 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.

## 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