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Extinct family of beetles
Coptoclavidae is an extinct family of aquatic
beetles in the suborder
Adephaga .
[1]
[2] The Coptoclavidae lived from the
Late Triassic to the
Early Cretaceous .
[3] Coptoclavidae is a member of the adephagan clade
Dytiscoidea ,
[4]
[5]
[6] which contains other aquatic beetles. Suggested reasons for their extinction to include the rise of
teleost fish, or competition with
Gyrinidae and
Dytiscidae , which possess defensive secretions and sucking channels in the mandibles of larvae, which coptoclavids likely lacked. It has been suggested that the genus
Timarchopsis and the subfamily Timarchopsinae are only distantly related to other coptoclavids based on
cladistic analysis , with Timarchopsis being more closely related to geadephagans like
carabids and
trachypachids instead.
[7] Another study also suggested similarly for
Coptoclavisca and possibly other coptoclaviscines.
[8]
Taxonomy
†
Agrascapha Lin, 1992 –
Huangshanjie Formation , China,
Carnian
†
Amblycephalonius Bode, 1953 –
Posidonia Shale , Germany,
Toarcian
subfamily †Charonoscaphinae Ponomarenko, 1977
†
Coptoclavia Ping, 1928 –
Cassange Group ,
Upper Triassic
subfamily †Coptoclavinae Ponomarenko, 1961
†
Bolbonectes Ponomarenko, 1987
†Bolbonectes intermedius Ponomarenko, 1987 –
Byankino Formation , Russia,
Tithonian ;
Leskovskaya Formation , Russia,
Barremian
†Bolbonectes occidentalis Ponomarenko, 1993 –
Ichetuy Formation , Russia,
Oxfordian
†Bolbonectus lithographicus Ponomarenko and Martínez-Delclòs, 2000 –
La Pedrera de Rúbies Formation , Spain,
Barremian
†
Coptoclava Ping, 1928 –
Ustkarskaya Formation , Russia,
Hauterivian ;
Dabeigou Formation , China, Hauterivian;
Kutinskaya Formation , Russia, Barremian;
Dalazi Formation ,
Laiyang Formation ,
Jianchang Formation ,
Shouchang Formation ,
Jiufotang Formation ,
Chijinbao Formation ,
Aptian ;
Zaza Formation ,
Gidarinskaya Formation ,
Mangutskaya Formation ,
Turga Formation , Russia, Aptian
†
Hoyaclava Soriano et al. , 2007 –
Las Hoyas , Spain, Barremian
†
Megacoptoclava Ponomarenko and Martínez-Delclòs, 2000 – Las Hoyas, Spain, Barremian
subfamily †Coptoclaviscinae Soriano et al. , 2007
†
Coptoclavella Ponomarenko, 1980
†Coptoclavella elegans Ponomarenko, 1980 –
Mogotuin Formation , Mongolia, Aptian
†Coptoclavella inexpecta Soriano et al. , 2007 – La Pedrera de Rúbies Formation, Spain, Barremian
†Coptoclavella jurassica Ponomarenko, 2014 –
Sharteg , Mongolia, Tithonian
†Coptoclavella minor Ponomarenko, 1980 –
Takshin Formation ,
Kalgan Formation Russia, Callovian;
Daya Formation , Russia, Hauterivian; Mogotuin Formation, Mongolia, Aptian; Turga Formation, Russia, Aptian
†Coptoclavella purbeckensis Ponomarenko et al. , 2005 –
Durlston Formation , United Kingdom,
Berriasian
†Coptoclavella vittata Ponomarenko, 1986 –
Gurvan-Eren Formation , Mongolia, Aptian
†Coptoclavella striata Ponomarenko, 1986 – Gurvan-Eren Formation, Mongolia, Aptian
†
Coptoclavisca Ponomarenko, 1987 – Laiyang Formation, China, Aptian;
Tsagaantsav Formation , Mongolia,
Early Cretaceous
†
Stargelytron Ponomarenko, 2015 –
Röt Formation , Germany, Anisian;
Hassberge Formation , Germany, Carnian
†
Euroscapha Lin, 1992 – Huangshanjie Formation, China, Carnian
†
Holcoptera Handlirsch, 1906
†Holcoptera alisonae Kelly et al. , 2017 –
Charmouth Mudstone Formation , United Kingdom,
Sinemurian
†Holcoptera giebeli Handlirsch, 1906 –
Lilstock Formation , United Kingdom,
Rhaetian ; Charmouth Mudstone Formation, United Kingdom, Sinemurian
†Holcoptera pigmentatus Kelly et al. , 2017 – Lilstock Formation, United Kingdom, Rhaetian;
Portland Formation , Connecticut,
Hettangian ;
Mount Toby Formation , Massachusetts, Hettangian
†Holcoptera schlotheimi (Giebel, 1856) (=Holcoptera confluens Cockerell, 1915 ) –
Argilliti di Riva di Solto Formation , Italy,
Norian ; Lilstock Formation, United Kingdom, Rhaetian;
Blue Lias , United Kingdom, Hettangian; Portland Formation, Connecticut, Hettangian
†Holcoptera solitensis Kelly et al. , 2017 –
Cow Branch Formation , North Carolina, Norian
subfamily †Hispanoclavinae Soriano et al. , 2007
subfamily †Timarchopsinae Wang et al. , 2010
†
Actea Germar, 1842 –
Solnhofen , Germany, Tithonian
†
Daohugounectes Wang et al. , 2009 –
Daohugou , China, Callovian
†
Ditomoptera Germar, 1839 – Solnhofen, Germany, Tithonian
†
Exedia Ponomarenko, 1977 – Karabastau Formation, Kazakhstan, Callovian
†
Ovonectes Soriano et al. , 2007 – Las Hoyas, Spain, Barremian
†
Protonectes Prokin and Ponomarenko, 2013 – Hassberge Formation, Germany, Carnian
†
Pseudohydrophilus Deichmüller, 1886 – Solnhofen, Germany, Tithonian
†
Stygeonectes Ponomarenko, 1977 –
Cheremkhovskaya Formation , Russia, Toarcian;
Udinskaya Formation , Russia, Callovian;
Uda Formation , Ichetuy Formation, Russia, Oxfordian;
Glushkovo Formation , Russia, Tithonian;
Leskovskaya Formation ,
Bukachacha Formation , Russia, Barremian;
Godymboyskaya Formation , Turga Formation, Russia, Aptian
†
Timarchopsis Brauer et al. , 1889
†Timarchopsis aquaticus Ponomarenko, 1977 – Cheremkhovskaya Formation, Russia, Toarcian; Ichetuy Formation, Russia, Oxfordian
†Timarchopsis cyrenaicus Ponomarenko, 1977 – Algeria, Hauterivian
†Timarchopsis czekanowskii Brauer et al. , 1889 – Cheremkhovskaya Formation, Russia, Toarcian
†Timarchopsis gigas Ponomarenko, 1977 – Karabastau Formation, Kazakhstan, Callovian
†Timarchopsis gobiensis Ponomarenko, 1987 –
Ulaan-Ereg Formation, Mongolia, Tithonian
†Timarchopsis latus Ponomarenko, 1977 –
Sagul Formation , Kyrgyzstan, Toarcian
†Timarchopsis longus Ponomarenko, 2014 – Sharteg, Mongolia, Tithonian
†Timarchopsis mongolicus Ponomarenko, 1985 –
Zhargalant Formation , Mongolia,
Bathonian
†Timarchopsis sainshandensis Ponomarenko, 1987 –
Khamarkhoburinskaya Formation , Mongolia,
Aalenian
†
Tuhanectes Hong, 1995 –
Sanjianfang Formation , China, Bathonian
References
^ Tree of Life Web Project. 1995. Liadytidae. Liadytes. Version 1 January 1995 (temporary).
[1] .01.01 in The Tree of Life Web Project,
[2]
^ Ivan Löbl & Aleš Smetana (May 2003). "1". Catalogue of Palaearctic Coleoptera: Archostemata, Myxophaga, Adephaga . Vol. 1. Apollo Books. pp. 6–8.
ISBN
978-87-88757-73-6 .
^ Grimaldi, David and Michael S. Engel (16 May 2005). Evolution of the Insects .
Cambridge University Press .
ISBN
0-521-82149-5 .
^ Ponomarenko, A. G., Arnoldy, L., Jerikin, V. V., & Nikritin, L. M. (1992). Mesozoic Coleoptera. Suborder Adephaga , 1.
^ Wang, B.; Ponomarenko, A. G.; Zhang, H. -Ch. (15 December 2009). "A new coptoclavid larva (Coleoptera: Adephaga: Dytiscoidea) from the Middle Jurassic of China, and its phylogenetic implication". Paleontological Journal . 43 (6): 652.
doi :
10.1134/S0031030109060082 .
ISSN
1555-6174 .
S2CID
83756102 .
^ Bo, Wang; Ponomarenko, A. G.; Haichun, Zhang (2010). "Middle Jurassic Coptoclavidae (Insecta: Coleoptera: Dytiscoidea) from China: a Good Example of Mosaic Evolution". Acta Geologica Sinica - English Edition . 84 (4): 680–687.
doi :
10.1111/j.1755-6724.2010.00272.x .
ISSN
1755-6724 .
S2CID
83806777 .
^ Beutel, Rolf G.; Wang, Bo; Tan, Jing-Jing; Ge, Si-Qin; Ren, Dong; Yang, Xing-Ke (April 2013).
"On the phylogeny and evolution of Mesozoic and extant lineages of Adephaga (Coleoptera, Insecta)" . Cladistics . 29 (2): 147–165.
doi :
10.1111/j.1096-0031.2012.00420.x .
PMID
34814378 .
S2CID
85234655 .
^ Kirejtshuk, Alexander G.; Ansorge, Jörg (17 May 2022).
"An extraordinarily preserved new genus and species of Trachypachidae (Coleoptera, Adephaga) from the Early Jurassic of Germany and a review of fossil trachypachid genera" . Historical Biology . 35 (6): 958–975.
doi :
10.1080/08912963.2022.2071709 .
ISSN
0891-2963 .
S2CID
248869983 .