Coptoclavidae Explained

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 – Huangshanjie Formation, China, Carnian
• †Amblycephalonius – Posidonia Shale, Germany, Toarcian
• subfamily †Charonoscaphinae
  • †Charonoscapha – Karabastau Formation, Kazakhstan, Callovian
  • †Charonoscaphidia – Karabastau Formation, Kazakhstan, Callovian
• †Coptoclavia – Cassange Group, Upper Triassic
• subfamily †Coptoclavinae
  • †Bolbonectes
    • †Bolbonectes intermedius – Byankino Formation, Russia, Tithonian; Leskovskaya Formation, Russia, Barremian
    • †Bolbonectes occidentalis – Ichetuy Formation, Russia, Oxfordian
    • †Bolbonectus lithographicus – La Pedrera de Rúbies Formation, Spain, Barremian
  • †Coptoclava – 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 – Las Hoyas, Spain, Barremian
  • †Megacoptoclava – Las Hoyas, Spain, Barremian
• subfamily †Coptoclaviscinae
  • †Coptoclavella
    • †Coptoclavella elegans – Mogotuin Formation, Mongolia, Aptian
    • †Coptoclavella inexpecta – La Pedrera de Rúbies Formation, Spain, Barremian
    • †Coptoclavella jurassica – Sharteg, Mongolia, Tithonian
    • †Coptoclavella minor – Takshin Formation, Kalgan Formation Russia, Callovian; Daya Formation, Russia, Hauterivian; Mogotuin Formation, Mongolia, Aptian; Turga Formation, Russia, Aptian
    • †Coptoclavella purbeckensis – Durlston Formation, United Kingdom, Berriasian
    • †Coptoclavella vittata – Gurvan-Eren Formation, Mongolia, Aptian
    • †Coptoclavella striata – Gurvan-Eren Formation, Mongolia, Aptian
  • †Coptoclavisca – Laiyang Formation, China, Aptian; Tsagaantsav Formation, Mongolia, Early Cretaceous
  • †Stargelytron – Röt Formation, Germany, Anisian; Hassberge Formation, Germany, Carnian
• †Euroscapha – Huangshanjie Formation, China, Carnian
• †Holcoptera
  • †Holcoptera alisonae – Charmouth Mudstone Formation, United Kingdom, Sinemurian
  • †Holcoptera giebeli – Lilstock Formation, United Kingdom, Rhaetian; Charmouth Mudstone Formation, United Kingdom, Sinemurian
  • †Holcoptera pigmentatus – Lilstock Formation, United Kingdom, Rhaetian; Portland Formation, Connecticut, Hettangian; Mount Toby Formation, Massachusetts, Hettangian
  • †Holcoptera schlotheimi (=Holcoptera confluens) – Argilliti di Riva di Solto Formation, Italy, Norian; Lilstock Formation, United Kingdom, Rhaetian; Blue Lias, United Kingdom, Hettangian; Portland Formation, Connecticut, Hettangian
  • †Holcoptera solitensis – Cow Branch Formation, North Carolina, Norian
• subfamily †Hispanoclavinae
  • †Hispanoclavina – Las Hoyas, Spain, Barremian
• subfamily †Timarchopsinae
  • †Actea – Solnhofen, Germany, Tithonian
  • †Daohugounectes – Daohugou, China, Callovian
  • †Ditomoptera – Solnhofen, Germany, Tithonian
  • †Exedia – Karabastau Formation, Kazakhstan, Callovian
  • †Ovonectes – Las Hoyas, Spain, Barremian
  • †Protonectes – Hassberge Formation, Germany, Carnian
  • †Pseudohydrophilus – Solnhofen, Germany, Tithonian
  • †Stygeonectes – 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
    • †Timarchopsis aquaticus – Cheremkhovskaya Formation, Russia, Toarcian; Ichetuy Formation, Russia, Oxfordian
    • †Timarchopsis cyrenaicus – Algeria, Hauterivian
    • †Timarchopsis czekanowskii – Cheremkhovskaya Formation, Russia, Toarcian
    • †Timarchopsis gigas – Karabastau Formation, Kazakhstan, Callovian
    • †Timarchopsis gobiensis – Ulaan-Ereg Formation, Mongolia, Tithonian
    • †Timarchopsis latus – Sagul Formation, Kyrgyzstan, Toarcian
    • †Timarchopsis longus – Sharteg, Mongolia, Tithonian
    • †Timarchopsis mongolicus – Zhargalant Formation, Mongolia, Bathonian
    • †Timarchopsis sainshandensis – Khamarkhoburinskaya Formation, Mongolia, Aalenian
  • †Tuhanectes – Sanjianfang Formation, China, Bathonian

Notes and References

1. Tree of Life Web Project. 1995. Liadytidae. Liadytes. Version 1 January 1995 (temporary). http://tolweb.org/Liadytes/8898/1995.01.01 in The Tree of Life Web Project, http://tolweb.org/
2. Book: Catalogue of Palaearctic Coleoptera: Archostemata, Myxophaga, Adephaga. Ivan Löbl & Aleš Smetana. May 2003. Apollo Books. 978-87-88757-73-6. 1. 6–8. 1.
3. Book: Grimaldi, David and Michael S. Engel. Evolution of the Insects. 2005-05-16. Cambridge University Press. 0-521-82149-5.
4. Ponomarenko, A. G., Arnoldy, L., Jerikin, V. V., & Nikritin, L. M. (1992). Mesozoic Coleoptera. Suborder Adephaga, 1.
5. Wang. B.. Ponomarenko. A. G.. Zhang. H. -Ch.. 2009-12-15. A new coptoclavid larva (Coleoptera: Adephaga: Dytiscoidea) from the Middle Jurassic of China, and its phylogenetic implication. Paleontological Journal. 43. 6. 652. 10.1134/S0031030109060082. 2009PalJ...43..652W . 83756102 . 1555-6174.
6. 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. 10.1111/j.1755-6724.2010.00272.x. 2010AcGlS..84..680W . 83806777 . 1755-6724.
7. 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 . 10.1111/j.1096-0031.2012.00420.x. 34814378 . 85234655 .
8. Kirejtshuk . Alexander G. . Ansorge . Jörg . 2022-05-17 . 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 . 10.1080/08912963.2022.2071709 . 248869983 . 0891-2963.