Coptoclavidae

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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, 1992Script error: No such module "Check for unknown parameters". – Huangshanjie Formation, China, Carnian
  • †Amblycephalonius Bode, 1953Script error: No such module "Check for unknown parameters". – Posidonia Shale, Germany, Toarcian
  • subfamily †Charonoscaphinae Ponomarenko, 1977Script error: No such module "Check for unknown parameters".
  • †Coptoclavia Ping, 1928Script error: No such module "Check for unknown parameters". – Cassange Group, Upper Triassic
  • subfamily †Coptoclavinae Ponomarenko, 1961Script error: No such module "Check for unknown parameters".
  • subfamily †Coptoclaviscinae Soriano et al., 2007Script error: No such module "Check for unknown parameters".
    • †Coptoclavella Ponomarenko, 1980Script error: No such module "Check for unknown parameters".
      • †Coptoclavella elegans Ponomarenko, 1980Script error: No such module "Check for unknown parameters". – Mogotuin Formation, Mongolia, Aptian
      • †Coptoclavella inexpecta Soriano et al., 2007Script error: No such module "Check for unknown parameters". – La Pedrera de Rúbies Formation, Spain, Barremian
      • †Coptoclavella jurassica Ponomarenko, 2014Script error: No such module "Check for unknown parameters". – Sharteg, Mongolia, Tithonian
      • †Coptoclavella minor Ponomarenko, 1980Script error: No such module "Check for unknown parameters". – Takshin Formation, Kalgan Formation Russia, Callovian; Daya Formation, Russia, Hauterivian; Mogotuin Formation, Mongolia, Aptian; Turga Formation, Russia, Aptian
      • †Coptoclavella purbeckensis Ponomarenko et al., 2005Script error: No such module "Check for unknown parameters". – Durlston Formation, United Kingdom, Berriasian
      • †Coptoclavella vittata Ponomarenko, 1986Script error: No such module "Check for unknown parameters". – Gurvan-Eren Formation, Mongolia, Aptian
      • †Coptoclavella striata Ponomarenko, 1986Script error: No such module "Check for unknown parameters". – Gurvan-Eren Formation, Mongolia, Aptian
    • †Coptoclavisca Ponomarenko, 1987Script error: No such module "Check for unknown parameters". – Laiyang Formation, China, Aptian; Tsagaantsav Formation, Mongolia, Early Cretaceous
    • †Holcoptera Handlirsch, 1906Script error: No such module "Check for unknown parameters". (= Stargelytron Ponomarenko, 2015Script error: No such module "Check for unknown parameters".)
      • †Holcoptera alisonae Kelly et al., 2017Script error: No such module "Check for unknown parameters". – Charmouth Mudstone Formation, United Kingdom, Sinemurian
      • †Holcoptera altus (Ponomarenko, 2015Script error: No such module "Check for unknown parameters".) (= Stargelytron altus Ponomarenko, 2015Script error: No such module "Check for unknown parameters".) – Röt Formation, Germany, Anisian
      • †Holcoptera giebeli Handlirsch, 1906Script error: No such module "Check for unknown parameters". – Lilstock Formation, United Kingdom, Rhaetian; Charmouth Mudstone Formation, United Kingdom, Sinemurian
      • †Holcoptera larissae (Ponomarenko, 2015Script error: No such module "Check for unknown parameters".) (= Stargelytron larissae Ponomarenko, 2015Script error: No such module "Check for unknown parameters".) – Hassberge Formation, Germany, Carnian
      • †Holcoptera pigmentatus Kelly et al., 2017Script error: No such module "Check for unknown parameters". – Lilstock Formation, United Kingdom, Rhaetian; Portland Formation, Connecticut, Hettangian; Mount Toby Formation, Massachusetts, Hettangian
      • †Holcoptera rasnitsyni Ponomarenko and Fedorenko, 2022Script error: No such module "Check for unknown parameters". – Protopivka Formation, Ukraine, Norian
      • †Holcoptera schlotheimi (Giebel, 1856)Script error: No such module "Check for unknown parameters". (=Holcoptera confluens Cockerell, 1915Script error: No such module "Check for unknown parameters".) – 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., 2017Script error: No such module "Check for unknown parameters". – Cow Branch Formation, North Carolina, Norian
  • †Euroscapha Lin, 1992Script error: No such module "Check for unknown parameters". – Huangshanjie Formation, China, Carnian
  • subfamily †Hispanoclavinae Soriano et al., 2007Script error: No such module "Check for unknown parameters".
    • †Hispanoclavina Soriano et al., 2007Script error: No such module "Check for unknown parameters". – Las Hoyas, Spain, Barremian
  • subfamily †Timarchopsinae Wang et al., 2010Script error: No such module "Check for unknown parameters".
    • †Actea Germar, 1842Script error: No such module "Check for unknown parameters". – Solnhofen, Germany, Tithonian
    • †Daohugounectes Wang et al., 2009Script error: No such module "Check for unknown parameters". – Daohugou, China, Callovian
    • †Ditomoptera Germar, 1839Script error: No such module "Check for unknown parameters". – Solnhofen, Germany, Tithonian
    • †Exedia Ponomarenko, 1977Script error: No such module "Check for unknown parameters". – Karabastau Formation, Kazakhstan, Callovian
    • †Ovonectes Soriano et al., 2007Script error: No such module "Check for unknown parameters". – Las Hoyas, Spain, Barremian
    • †Protonectes Prokin and Ponomarenko, 2013Script error: No such module "Check for unknown parameters". – Hassberge Formation, Germany, Carnian
    • †Pseudohydrophilus Deichmüller, 1886Script error: No such module "Check for unknown parameters". – Solnhofen, Germany, Tithonian
    • †Stygeonectes Ponomarenko, 1977Script error: No such module "Check for unknown parameters". – 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., 1889Script error: No such module "Check for unknown parameters".
      • †Timarchopsis aquaticus Ponomarenko, 1977Script error: No such module "Check for unknown parameters". – Cheremkhovskaya Formation, Russia, Toarcian; Ichetuy Formation, Russia, Oxfordian
      • †Timarchopsis cyrenaicus Ponomarenko, 1977Script error: No such module "Check for unknown parameters". – Algeria, Hauterivian
      • †Timarchopsis czekanowskii Brauer et al., 1889Script error: No such module "Check for unknown parameters". – Cheremkhovskaya Formation, Russia, Toarcian
      • †Timarchopsis gigas Ponomarenko, 1977Script error: No such module "Check for unknown parameters". – Karabastau Formation, Kazakhstan, Callovian
      • †Timarchopsis gobiensis Ponomarenko, 1987Script error: No such module "Check for unknown parameters". – Ulaan-Ereg Formation, Mongolia, Tithonian
      • †Timarchopsis latus Ponomarenko, 1977Script error: No such module "Check for unknown parameters". – Sagul Formation, Kyrgyzstan, Toarcian
      • †Timarchopsis longus Ponomarenko, 2014Script error: No such module "Check for unknown parameters". – Sharteg, Mongolia, Tithonian
      • †Timarchopsis mongolicus Ponomarenko, 1985Script error: No such module "Check for unknown parameters". – Zhargalant Formation, Mongolia, Bathonian
      • †Timarchopsis sainshandensis Ponomarenko, 1987Script error: No such module "Check for unknown parameters". – Khamarkhoburinskaya Formation, Mongolia, Aalenian
    • †Tuhanectes Hong, 1995Script error: No such module "Check for unknown parameters". – Sanjianfang Formation, China, Bathonian

References

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  1. ↑ Tree of Life Web Project. 1995. Liadytidae. Liadytes. Version 1 January 1995 (temporary). [1].01.01 in The Tree of Life Web Project, [2]
  2. ↑ Script error: No such module "citation/CS1".
  3. ↑ Script error: No such module "citation/CS1".
  4. ↑ Ponomarenko, A. G., Arnoldy, L., Jerikin, V. V., & Nikritin, L. M. (1992). Mesozoic Coleoptera. Suborder Adephaga, 1.
  5. ↑ Script error: No such module "Citation/CS1".
  6. ↑ Script error: No such module "Citation/CS1".
  7. ↑ a b Script error: No such module "Citation/CS1".
  8. ↑ a b Script error: No such module "Citation/CS1".
  9. ↑ Script error: No such module "Citation/CS1".

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