Coptoclavidae

Coptoclavidae Temporal range: Late Triassic–Early Cretaceous PreꞒ Ꞓ O S D C P T J K Pg N
Holcoptera giebeli (specimen NMHUK PI II.3101)
Larva of Coptoclava
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Coleoptera
Suborder:	Adephaga
Family:	† Coptoclavidae Ponomarenko, 1961
Subfamilies
Charonoscaphinae Coptoclavinae Coptoclaviscinae Hispanoclavinae Timarchopsinae Coptoclavidae ( incertae sedis )

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 living aquatic beetles, including living predatory diving beetles (Dytiscidae). Coptoclavids are thought to have hunted on the water surface, similar to whirligig beetles (Gyrinidiae), with prey likely including small fish and larval amphibians. ^[7] 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. ^[8] 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. ^[8] Another study also suggested similarly for Coptoclavisca and possibly other coptoclaviscines. ^[9] A 2025 study suggested that Coptoclavidae as a whole, as well as most of the major subfamilies, were likely not monophyletic, but represent a number of separate lineages of dytiscoid beetles. ^[7]

Taxonomy

• † Agrascapha Lin, 1992 – Huangshanjie Formation, China, Carnian
• † Amblycephalonius Bode, 1953 – Posidonia Shale, Germany, Toarcian
• subfamily †Charonoscaphinae Ponomarenko, 1977
  • † Charonoscapha Ponomarenko, 1977 – Karabastau Formation, Kazakhstan, Callovian
  • † Charonoscaphidia Ponomarenko, 1977 – Karabastau Formation, Kazakhstan, Callovian
• † Coptoclavia Ping, 1928 – Cassange Group, Upper Triassic
• subfamily †Coptoclavinae Ponomarenko, 1961
  • † Bolbonectes Ponomarenko, 1987
    • †Bolbonectes intermediusPonomarenko, 1987 – Byankino Formation, Russia, Tithonian; Leskovskaya Formation, Russia, Barremian
    • †Bolbonectes occidentalisPonomarenko, 1993 – Ichetuy Formation, Russia, Oxfordian
    • †Bolbonectus lithographicusPonomarenko 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 elegansPonomarenko, 1980 – Mogotuin Formation, Mongolia, Aptian
    • †Coptoclavella inexpectaSoriano et al., 2007 – La Pedrera de Rúbies Formation, Spain, Barremian
    • †Coptoclavella jurassicaPonomarenko, 2014 – Sharteg, Mongolia, Tithonian
    • †Coptoclavella minorPonomarenko, 1980 – Takshin Formation, Kalgan Formation Russia, Callovian; Daya Formation, Russia, Hauterivian; Mogotuin Formation, Mongolia, Aptian; Turga Formation, Russia, Aptian
    • †Coptoclavella purbeckensisPonomarenko et al., 2005 – Durlston Formation, United Kingdom, Berriasian
    • †Coptoclavella vittataPonomarenko, 1986 – Gurvan-Eren Formation, Mongolia, Aptian
    • †Coptoclavella striataPonomarenko, 1986 – Gurvan-Eren Formation, Mongolia, Aptian
  • † Coptoclavisca Ponomarenko, 1987 – Laiyang Formation, China, Aptian; Tsagaantsav Formation, Mongolia, Early Cretaceous
  • † Holcoptera Handlirsch, 1906 (= StargelytronPonomarenko, 2015)
    • †Holcoptera alisonaeKelly et al., 2017 – Charmouth Mudstone Formation, United Kingdom, Sinemurian
    • †Holcoptera altus (Ponomarenko, 2015) (= Stargelytron altusPonomarenko, 2015) – Röt Formation, Germany, Anisian
    • †Holcoptera giebeliHandlirsch, 1906 – Lilstock Formation, United Kingdom, Rhaetian; Charmouth Mudstone Formation, United Kingdom, Sinemurian
    • †Holcoptera larissae (Ponomarenko, 2015) (= Stargelytron larissaePonomarenko, 2015) – Hassberge Formation, Germany, Carnian
    • †Holcoptera pigmentatusKelly et al., 2017 – Lilstock Formation, United Kingdom, Rhaetian; Portland Formation, Connecticut, Hettangian; Mount Toby Formation, Massachusetts, Hettangian
    • †Holcoptera rasnitsyniPonomarenko and Fedorenko, 2022 – Protopivka Formation, Ukraine, Norian
    • †Holcoptera schlotheimi(Giebel, 1856) (=Holcoptera confluensCockerell, 1915) – Argilliti di Riva di Solto Formation, Italy, Norian; Lilstock Formation, United Kingdom, Rhaetian; Blue Lias, United Kingdom, Hettangian; Portland Formation, Connecticut, Hettangian
    • †Holcoptera solitensisKelly et al., 2017 – Cow Branch Formation, North Carolina, Norian
• † Euroscapha Lin, 1992 – Huangshanjie Formation, China, Carnian
• subfamily †Hispanoclavinae Soriano et al., 2007
  • † Hispanoclavina Soriano et al., 2007 – Las Hoyas, Spain, Barremian
• 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 aquaticusPonomarenko, 1977 – Cheremkhovskaya Formation, Russia, Toarcian; Ichetuy Formation, Russia, Oxfordian
    • †Timarchopsis cyrenaicusPonomarenko, 1977 – Algeria, Hauterivian
    • †Timarchopsis czekanowskiiBrauer et al., 1889 – Cheremkhovskaya Formation, Russia, Toarcian
    • †Timarchopsis gigasPonomarenko, 1977 – Karabastau Formation, Kazakhstan, Callovian
    • †Timarchopsis gobiensisPonomarenko, 1987 – Ulaan-Ereg Formation, Mongolia, Tithonian
    • †Timarchopsis latusPonomarenko, 1977 – Sagul Formation, Kyrgyzstan, Toarcian
    • †Timarchopsis longusPonomarenko, 2014 – Sharteg, Mongolia, Tithonian
    • †Timarchopsis mongolicusPonomarenko, 1985 – Zhargalant Formation, Mongolia, Bathonian
    • †Timarchopsis sainshandensisPonomarenko, 1987 – Khamarkhoburinskaya Formation, Mongolia, Aalenian
  • † Tuhanectes Hong, 1995 – Sanjianfang Formation, China, Bathonian

References

1. Tree of Life Web Project. 1995. Liadytidae. Liadytes. Version 1 January 1995 (temporary). .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.
3. Grimaldi, David and Michael S. Engel (16 May 2005). Evolution of the Insects. Cambridge University Press. ISBN   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. (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. Bibcode:2009PalJ...43..652W. doi:10.1134/S0031030109060082. ISSN   1555-6174. S2CID   83756102.
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. Bibcode:2010AcGlS..84..680W. doi:10.1111/j.1755-6724.2010.00272.x. ISSN   1755-6724. S2CID   83806777.
7. Bao, Liang; Beutel, Rolf Georg; Niu, Kecheng; Bao, Tong (17 February 2025). "Redescription of the highly specialized aquatic †Tigrivia and the classification of the Mesozoic †Coptoclavidae (Coleoptera, Adephaga, Dytiscoidea)" . Zootaxa. 5588 (3): 401–425. doi:10.11646/zootaxa.5588.3.1. ISSN   1175-5334.
8. 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.
9. 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.