Actinopteri

Reconstruction of Birgeria, copyright DiBgd.

Belongs within: Actinopterygii.
Contains: Neopterygii, Palaeonisciformes, Acipenseriformes.

The Actinopteri are a clade of fishes including all modern ray-finned fishes with the exception of the bichirs of the Polypteridae. Members of this clade have the premaxilla confined to a region anterior to the orbit, parasphenoid teeth absent, and proximal radials of the dorsal fin substantially enlarged relative to the middle radials (Giles et al. 2017).

<==Actinopteri XG11
    |--+--Neopterygii XG11
    |  `--+--Palaeonisciformes GX17
    |     `--Acipenserifomes GX17
    `--+--Brachydegma [Brachydegmidae] GX17
       |    `--B. caelatum Dunkle 1939 G93
       `--Birgeriidae G93
            |--Psilichthys selwyni Hall 1900 G93
            `--Birgeria Stensiö 1919 GX17, G93
                 `--B. groenlandica GX17

Actinopteri incertae sedis:
  Willomorichthyidae [Rhabdolepididae] G93
    |--Willomorichthys striatulus Gardiner 1969 G93
    |--Rhabdolepis macropterus (Bronn 1829) G93
    |--Tienshaniscus longipterus Lui & Wang 1978 G93
    |--Sinoniscus macrolepis Lui & Wang 1978 G93
    `--Bendenius Traquair 1878 G93
         `--B. deneensis (Traquair 1878) G93
  Phanerorhynchus [Phanerorhynchidae, Phanerorhynchiformes] G93
    `--P. armatus Gill 1923 G93
  Holuridae G93
    |--Holurus parki Traquair 1881 G93
    |--Australichthys longidorsalis Gardiner 1969 G93
    |--Holuropsis yavorskyi Berg 1947 G93
    `--Palaeonisconotus G93
  Atherstoniidae G93
    |--Aestuarichthys fulcratus Gardiner 1969 G93
    `--Atherstonia Woodward 1889 G93
         `--A. australis Woodward 1902 F71
  Dwykia [Dwykiidae] G93
    `--D. analensis Gardiner 1969 G93
  Phanerosteon [Carbovelidae] G93
    `--P. mirabile Traquair 1881 G93
  Strepheoschemidae G93
    |--Strepheoschema fouldenensis White 1927 G93
    `--Aetheretmon valentiacum White 1927 G93
  Cornuboniscus [Cornuboniscidae] G93
    `--C. budensis White 1939 G93
  Amblypteridae G93
    |--Paramblypterus decorus (Egerton 1850) G93
    |--Amblypterus Agassiz 1833 G93
    |--Amblypterina Berg 1940 G93
    |--Gardinerichthys tewarii Gupta et al. 1978 G93
    `--Korutichthys korutensis Kazantseva 1980 G93
  Haplolepididae G93
    |--Haplolepis Miller 1892 G93
    `--Pyritocephalus sculptus Fric 1894 G93
  Tarrasius [Tarrasiidae, Tarrasiiformes] G93
    `--T. problematicus Traquair 1881 G93
  Aeduelliidae G93
    |--Aeduella blainvillei (Agassiz 1833) G93
    |--Westollia crassus (Pholig 1892) G93
    |--Igornella Heyler 1969 G93
    |--Decazella Heyler 1969 G93
    |--Burbonella Heyler 1969 G93
    `--Palaeothrissum Blainville 1818 G93
  Cryphiolepis [Cryphiolepididae] G93
    `--C. striatus (Traquair 1881) G93
  Urosthenes [Urosthenidae] G93
    |--U. australis Dana 1848 G93
    `--U. latus Woodward 1931 G93
  Lawnia [Lawniidae] G93
    `--L. taylorensis Wilson 1953 G93
  Trissolepididae G93
    |--Sceletophorus biserialis Frič 1894 G93
    `--Sphaerolepis kounoviensis Frič 1877 G93
  Commentrya [Commentryidae] G93
    `--C. traquairi Sauvage 1888 G93
  Dorypterus [Dorypteridae, Dorypteriformes, Dorypteroidei] G93
    |--D. althausi (Münster 1842) G93
    `--D. hoffmanni Germar 1842 G93

*Type species of generic name indicated

REFERENCES

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[G93] Gardiner, B. G. 1993. Osteichthyes: basal actinopterygians. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 611–619. Chapman & Hall: London.

[GX17] Giles, S., G.-H. Xu, T. J. Near & M. Friedman. 2017. Early members of 'living fossil' lineage imply later origin of modern ray-finned fishes. Nature 549: 265–268.

[XG11] Xu, G.-H. & K.-Q. Gao. 2011. A new scanilepiform from the Lower Triassic of northern Gansu Province, China, and phylogenetic relationships of non-teleostean Actinopterygii. Zoological Journal of the Linnean Society 161: 595–612.

Semionotiformes

Lepidotes elvensis, copyright Ghedoghedo.

Belongs within: Holostei.

The Semionotiformes are a group of fishes including the modern gars and their fossil relatives, united by the absence of the intercalar and gular bones, the presence of a series of antorbitals anterior to the circum-orbital ring, and enlarged basal and fringing fulcra (Cavin 2010).

Semionotiformes [Ginglymodi, Semionotoidei]
    |  i. s.: Clastes Cope 1872 C77
    |           |--C. aganus Cope 1877 C77
    |           |--C. anax C77
    |           |--C. atrox [=Lepidosteus atrox] C77
    |           |--C. cycliferus C77
    |           |--C. integer Cope 1877 C77
    |           |--C. notabilis [=Lepidosteus notabilis] C77
    |           `--‘Lepidosteus’ simplex C77
    |         Pneumatosteus C77
    |--+--Notagogus ferreri C10, S02
    |  |--Propterus GX17
    |  |    |--P. elongatus GX17
    |  |    `--P. vidali S02
    |  `--Macrosemiidae G93
    |       |--Macrosemius rostratus GX17, FS10
    |       |--Legonotus Egerton 1854 G93
    |       `--Enchelyolepis Woodward 1918 G93
    `--+--Macrosemimimus lennieri GX17
       |--Sangiorgichthys aldae C10
       `--Semionotidae
            |  i. s.: Lepidotyle Meunier & Gayet 1992 GM01
            |           `--L. enigmatica Meunier & Gayet 1992 GM01
            |         Alleiolepis Heller 1953 G93
            |         Aphelolepis Heller 1953 G93
            |         Asialepidotus Su 1959 G93
            |         Sinosemionotus Yuan & Koh 1936 G93
            |         Allelepidotus Frič 1906 G93
            |         Pericentrophorus Jörg 1969 G93
            |--+--Semionotus Agassiz 1832 C10, G93
            |  |    |--S. bergeri C10
            |  |    `--+--S. elegans C10
            |  |       `--S. kanabensis C10
            |  `--Lepidotes Agassiz 1832 C10, G93
            |       |--+--L. lennieri C10
            |       |  `--+--L. gloriae C10
            |       |     `--L. tendaguruensis C10
            |       `--+----L. minor C10
            |          `--+--L. mantelli C10
            |             `--+--L. semiserratus C10
            |                |--+--L. elvensis C10
            |                |  `--+--L. latifrons C10
            |                |     `--Isanichthys palustris C10
            |                `--+--L. piauhyensis C10
            |                   `--Paralepidotus Stolley 1920 C10, G93
            |                        `--P. ornatus C10
            `--+--Neosemionotus puntanus C10
               `--+--‘Lepidotes’ microrhis C10
                  `--+--‘Lepidotes’ buddhabutrensis C10
                     `--+--+--Araripelepidotes temnurus C10
                        |  `--Pliodetes nigeriensis C10
                        `--+--Masillosteus kelleri C10
                           `--Lepisosteidae [Lepisosteiformes, Lepisostoidei] B-RB13
                                |  i. s.: Paralepidosteus Arambourg & Joleaud 1943 G93
                                |--Obaichthys decoratus GX17
                                `--+--Oniichthys falipoui C10
                                   |--Lepisosteus Lacépède 1903 GX17, G93
                                   |    |--L. osseus GM01
                                   |    `--L. platostomus FS10
                                   `--Atractosteus GX17
                                        |--A. ferox C77
                                        |--A. spatula B96
                                        |--A. tristoechus R88 [=Lepisosteus tristoechus G88]
                                        `--A. tropicus R88

*Type species of generic name indicated

REFERENCES

[B-RB13] Betancur-R., R., R. E. Broughton, E. O. Wiley, K. Carpenter, A. López, C. Li, N. I. Holcroft, D. Arcila, M. Sanciangco, J. C. Cureton, II, F. Zhang, T. Buser, M. A. Campbell, J. A. Ballesteros, A. Roa-Varon, S. Willis, W. C. Borden, T. Rowley, P. C. Reneau, D. J. Hough, G. Lu, T. Grande, G. Arratia & G. Ortí. 2013. The tree of life and a new classification of bony fishes. PLoS Currents Tree of Life April 18 2013. doi: 10.1371/currents.tol.53ba26640df0ccaee75bb165c8c26288.

[B96] Bond, C. E. 1996. Biology of Fishes 2^nd ed. Saunders College Publishing: Fort Worth.

[C10] Cavin, L. 2010. Diversity of Mesozoic semionotiform fishes and the origin of gars (Lepisosteidae). Naturwissenschaften 97: 1035–1040.

[C77] Cope, E. D. 1877. Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874. Geographical Surveys West of the One Hundredth Meridian 4 (2): i–iv, 1–370.

[FS10] Friedman, M., K. Shimada, L. D. Martin, M. J. Everhart, J. Liston, A. Maltese & M. Triebold. 2010. 100-million-year dynasty of giant planktivorous bony fishes in the Mesozoic seas. Science 327: 990–993.

[G93] Gardiner, B. G. 1993. Osteichthyes: basal actinopterygians. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 611–619. Chapman & Hall: London.

[GM01] Gayet, M., L. G. Marshall, T. Sempere, F. J. Meunier, H. Cappetta & J-C. Rage. 2001. Middle Maastrichtian vertebrates (fishes, amphibians, dinosaurs and other reptiles, mammals) from Pajcha Pata (Bolivia). Biostratigraphic, palaeoecologic and palaeobiostratigraphic implications. Palaeogeography, Palaeoclimatology, Palaeoecology 169: 39–68.

[GX17] Giles, S., G.-H. Xu, T. J. Near & M. Friedman. 2017. Early members of 'living fossil' lineage imply later origin of modern ray-finned fishes. Nature 549: 265–268.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[R88] Rauchenberger, M. 1988. Historical biogeography of poeciliid fishes in the Caribbean. Systematic Zoology 37 (4): 356–365.

[S02] Sinitshenkova, N. D. 2002. Ecological history of the aquatic insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 388–426. Kluwer Academic Publishers: Dordrecht.

Anguilloidei

European eels Anguilla anguilla, copyright Dmitriy Konstantinov.

Belongs within: Anguilliformes.

The Anguilloidei are a group of eels in which the premaxillaries are fused with the mesethmoid to form a tooth-bearing bone at the front of the upper jaw (Bond 1996). They include the typical eels of the genus Anguilla, species of which spend the greater part of their post-larval life cycle in fresh water before returning to the oceans to breed.

Anguilloidei [Anguilloidea]
    |--Milananguilla [Milananguillidae] P93
    |    `--M. lehmani Blot 1978 P93
    |--Anguilloididae P93
    |    |--Anguilloides branchiostegalis (Eastman 1905) P93
    |    `--Veronanguilla ruffoi Blot 1978 P93
    |--Heterenchelyidae B96
    |    |--Heterenchelys [incl. Pythonichthys] B96
    |    |--Panturichthys B96
    |    `--Heterenchelyidarum richardsi Nolf 1988 P93
    |--Moringuidae [Ratabouridae] IM01
    |    |  i. s.: ‘Leptocephalus’ tuberculatus Castle 1965 C65
    |    |--Neoconger tuberculatus (Castle 1965) S99b
    |    |--Stilbiscus Jordan & Bollman 1889 [incl. Anguillichthys Mowbray 1927] C65
    |    |    `--S. bahamensis (Mowbray 1927) C65
    |    |--Aphthalmichthys M58
    |    |    |--A. bicolor (Kaup 1856) [=Moringua bicolor] M58
    |    |    |--A. floresiana (Weber & de Beaufort 1916) [=Moringua floresiana] M58
    |    |    |--A. macrocephalus Bleeker 1863 [=Moringua macrocephalus] M58
    |    |    `--A. macrochir (Bleeker 1855) [=Moringua macrochir] M58
    |    `--Moringua Gray 1831 TP86
    |         |--*M. raitaborua Ham-Buch. 1822 S62
    |         |--M. abbreviata (Bleeker 1863) TP86 [=Aphthalmichthys abbreviatus M58]
    |         |--‘Leptocephalus’ diptychus Eigenmann & Kennedy 1900 C65
    |         |--M. edwardsi IM01
    |         |--M. javanica (Kaup 1856) TP86 [=Aphthalmichthys javanicus M58]
    |         |--M. lumbricoidea JR10
    |         |--M. microchir Bleeker 1853 S62 [=Aphthalmichthys microchir M58; incl. M. ferruginea Bliss 1883 S62]
    |         `--M. raitaborua Hamilton-Buchanan 1822 C65
    `--Anguillidae IM01
         |--Eoanguilla leptoptera (Agassiz 1835) P93
         `--Anguilla IM01
              |--A. amboinensis JR10
              |--A. anguilla (Linnaeus 1758) SE08
              |--A. annosa Stinton 1975 P93
              |--A. australis Richardson 1841 S99a
              |--A. bengalensis CS77
              |--A. bicolor McClelland 1844 S99a
              |    |--A. b. bicolor AC90
              |    `--A. b. pacifica Schmidt 1932 AC90
              |--A. celebesensis Kaup 1856 (see below for synonymy) M58
              |--A. chrysypa D56
              |--A. dieffenbachii WH02
              |--A. ignota G88
              |--A. interioris Whitley 1938 S99a
              |--A. japonica Temminck & Schlegel 1846 S99a
              |--A. malgumora Schlegel in Kaup 1846 [incl. A. borneensis] S99a
              |--A. manillensis JR10
              |--A. marmorata Quoy & Gaimard 1824 AC90
              |--A. mauritiania Bennett 1831 [=Muraena mauritiania; incl. A. labiata Weber 1908] M58
              |--A. megastoma Kaup 1856 S99a
              |--A. obscura Günther 1871 M58
              |--A. pachyura G88
              |--A. reinhardtii Steindachner 1867 M58
              |--A. rostrata (LeSueur 1817) ST02 [incl. A. texana Günther 1870 F16]
              |--A. spengeli Weber 1912 M58
              `--A. vulgaris B96

Inorganic: Anguilla japonica minilorientalis Okamura 1987 O87

Anguilla celebesensis Kaup 1856 [=Muraena celebesensis; incl. A. aneitensis Macleay 1880, A. otaheitensis Günther 1910] M58

*Type species of generic name indicated

REFERENCES

[AC90] Allen, G. R., & D. Coates. 1990. An ichthyological survey of the Sepik River, Papua New Guinea. Records of the Western Australian Museum Supplement 34: 31–116.

[B96] Bond, C. E. 1996. Biology of Fishes 2^nd ed. Saunders College Publishing: Fort Worth.

[C65] Castle, P. H. J. 1965. Moringuid leptocephali in Australasian waters. Transactions of the Royal Society of New Zealand, Zoology 7 (7): 125–133.

[CS77] Cramp, S., & K. E. L. Simmons (eds) 1977. Handbook of the Birds of Europe, the Middle East and North Africa: The Birds of the Western Palaearctic vol. 1. Ostrich to Ducks. Oxford University Press: Oxford.

[D56] Dawes, B. 1956. The Trematoda with special reference to British and other European forms. University Press: Cambridge.

[F16] Fowler, H. W. 1916. The fishes of Trinidad, Grenada, and St. Lucia, British West Indies. Proceedings of the Academy of Natural Sciences of Philadelphia 67 (3): 520–548.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[IM01] Inoue, J. G., M. Miya, K. Tsukamoto & M. Nishida. 2001. Complete mitochondrial DNA sequence of Conger myriaster (Teleostei: Anguilliformes): novel gene order for vertebrate mitochondrial genomes and the phylogenetic implications for anguilliform families. Journal of Molecular Evolution 52: 311–320.

[JR10] Jordan, D. S., & R. E. Richardson. 1910. Check-list of the species of fishes known from the Philippine archipelago. Bureau of Printing: Manila.

[M58] Munro, I. S. R. 1958. The fishes of the New Guinea region: a check-list of the fishes of New Guinea incorporating records of species collected by the Fisheries Survey Vessel “Fairwind” during the years 1948 to 1950. Papua and New Guinea Agricultural Journal 10 (4): 97–369 (reprinted 1958. Territory of Papua and New Guinea Fisheries Bulletin no. 1).

[O87] Okamura, C. 1987. New facts: Homo and all Vertebrata were born simultaneously in the former Paleozoic in Japan. Original Report of the Okamura Fossil Laboratory 15: 347–573.

[P93] Patterson, C. 1993. Osteichthyes: Teleostei. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 621–656. Chapman & Hall: London.

[SE08] Sevcsik, A., & T. Erös. 2008. A revised catalogue of freshwater fishes of Hungary and the neighbouring countries in the Hungarian Natural History Museum (Pisces). Annales Historico-Naturales Musei Nationalis Hungarici 100: 331–383.

[S99a] Smith, D. G. 1999a. Anguillidae. Freshwater eels. In: Carpenter, K. E., & V. H. Niem (eds) The Living Marine Resources of the Western Central Pacific vol. 3. Batoid fishes, chimaeras and bony fishes part 1 (Elopidae to Linophrynidae) pp. 1630–1636. FAO: Rome.

[S99b] Smith, D. G. 1999b. Moringuidae. Spaghetti eels. In: Carpenter, K. E., & V. H. Niem (eds) The Living Marine Resources of the Western Central Pacific vol. 3. Batoid fishes, chimaeras and bony fishes part 1 (Elopidae to Linophrynidae) pp. 1637–1638. FAO: Rome.

[ST02] Smith, D. G., & K. A. Tighe. 2002. Feshwater eels. Family Anguillidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3^rd ed. pp. 92–95. Smithsonian Institute Press: Washington.

[S62] Smith, J. L. B. 1962. Sand-dwelling eels of the western Indian Ocean and the Red Sea. Ichthyological Bulletin 24: 447–466.

[TP86] Thinès, G., & G. Proudlove. 1986. Pisces. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 709–733. E. J. Brill/Dr W. Backhuys: Leiden.

[WH02] Worthy, T. H., & R. N. Holdaway. 2002. The Lost World of the Moa: Prehistoric life of New Zealand. Indiana University Press: Bloomington (Indiana).

Last updated: 9 February 2022.

Pycnodontoidei

Apomesodon gibbosus, copyright Ghedoghedo.

Belongs within: Pycnodontiformes.
Contains: Pycnodontidae.

The Pycnodontoidei are a clade of pycnodontiform fishes characterised by the loss of scales over much of the body (Poyato-Ariza & Wenz 2002).

Characters (from Poyato-Ariza & Wenz 2002): Opercular process of hyomandibular absent; ossifications in gular region absent; two branchiostegal rays, thin and separated; crenulations on vomerine and prearticular teeth occasional and weak if present; scales not covering the whole body; scales rows between bases of lepidotrichia of unpaired fins absent.

Pycnodontoidei
    |--Eomesodon Woodward 1918 P-AW02
    |    |--*E. liassicus (Egerton 1855) [=Pycnodus liassicus] P-AW02
    |    |--E. barnesi (Woodward 1906) P-AW02
    |    |--E. depressus Woodward 1918 P-AW02
    |    `--E. hoeferi (Gorjanovic-Kramberger 1905) P-AW02
    `--+--Apomesodon Poyato-Ariza & Wenz 2002 P-AW02
       |    |--*A. gibbosus (Wagner 1851) [=Mesodon gibbosus, Gyrodus gibbosus Agassiz 1843 (n. n.)] P-AW02
       |    |--A. comosus (Thiollière 1858) [=Mesodon comosus, Macromesodon comosus] P-AW02
       |    `--A. surgens Poyato-Ariza & Wenz 2002 P-AW02
       `--Pycnodontoidea P-AW02
            |--Pycnodontidae P-AW02
            `--Coccodontidae P-AW02
                 |--Trewavasia White & Moy-Thomas 1941 P-AW02
                 |    `--*T. carinatus (Davis 1887) [=Xenopholis carinatus] P-AW02
                 `--+--Ichthyoceros Gayet 1984 P-AW02
                    |    `--*I. spinosus Gayet 1984 P-AW02
                    `--Coccodus Pictet 1850 P-AW02
                         |--*C. armatus Pictet 1850 P-AW02
                         `--C. lindstroemi Davis 1890 (n. d.) P-AW02

*Type species of generic name indicated

REFERENCES

[P-AW02] Poyato-Ariza, F. J., & S. Wenz. 2002. A new insight into pycnodontiform fishes. Geodiversitas 24 (1): 139–248.

Proscinetinae

Proscinetes cf. elegans, copyright Ghedoghedo.

Belongs within: Pycnodontidae.

The Proscinetidae are a group of pycnodontid fishes known from the Late Jurassic and Early Cretaceous, characterised by the alternation of one large and two small teeth in the main vomerine tooth row (Poyato-Ariza & Wenz 2002).

Characters (from Poyato-Ariza & Wenz 2002): Three vomerine tooth rows; main vomerine tooth row alternating between one large and two small teeth, with eight or nine teeth in total (counting each couple of alternating small teeth as one tooth); about 30 to 34 vertebrae; dorsal ridge scales separated from each other; 18 to 21 ventral keel scales; spines on ventral keel scales placed in the posterior region of the midline.

Proscinetinae
    |--Neoproscinetes Figueiredo & Silva Santos 1987 PA-W02
    |    `--*N. penalvai (Silva Santos 1970) [=Microdon penalvai] PA-W02
    `--Proscinetes Gistl 1848 PA-W02
         |--*P. elegans (Agassiz 1833) [=Microdon elegans] PA-W02
         |--P. bassanii (D’Erasmo 1914) [=Coelodus bassanii, Palaeobalistium bassanii] PA-W02
         |--P. bernardi (Thollière 1852) PA-W02
         |--P. egertoni (Thollière 1852) PA-W02
         |--P. itieri (Thollière 1850) PA-W02
         |--P. radiatus (Agassiz 1839) PA-W02
         |--P. sauvanasi (Thollière 1852) PA-W02
         |--P. thollieri (Saint-Seine 1949) PA-W02
         `--P. wagneri (Thollière 1852) [=Gyrodus wagneri] PA-W02

*Type species of generic name indicated

REFERENCES

[P-AW02] Poyato-Ariza, F. J., & S. Wenz. 2002. A new insight into pycnodontiform fishes. Geodiversitas 24 (1): 139–248.

Palaeonisciformes

Platysomus, from here.

Belongs within: Actinopteri.

The Palaeonisciformes are an early group of heavily scaled ray-finned fishes known from the Devonian to the Cretaceous (Bond 1996).

Characters (from Bond 1996): Body form varying from fusiform to compressiform; usually covered by ganoid scales. Head large, bony; mouth generally long, teeth conical; maxillary large, probably not movable.

<==Palaeonisciformes [Platysomiformes]
    |  i. s.: Pygopteridae P99
    |           |--Watsonichthys pectinatus (Traquair 1877) P99
    |           |--Nematoptychius Traquair 1875 P99
    |           |    `--*N. greenocki Traquair 1875 P99
    |           `--Progyrolepis Fritsch 1895 P99
    |                |--*P. speciosus (Fritsch 1875) P99
    |                |--P. heyleri Poplin 1999 P99
    |                `--P. tricessimalaris Dunkle 1946 P99
    |--Redfieldioidei B96
    |--Dorypteroidei B96
    |--Palaeoniscus M79 [Palaeoniscidae, Palaeoniscoidei B96]
    |    `--P. macropomus M79
    `--Platysomoidei B96
         |--Chirondontidae B96
         `--Platysomidae PL00
              |--Kargalichthys TT05
              |    |--K. efremovi Minikh 1986 TT05
              |    `--K. pritokensis Minikh 1992 TT05
              `--Platysomus PL00
                   |--P. bashkirus Minikh 1992 TT05
                   |--P. biarmicus Von Eichwald 1861 TT05
                   |--P. soloduchi Minikh 1992 TT05
                   `--P. superbus PL00

*Type species of generic name indicated

REFERENCES

[B96] Bond, C. E. 1996. Biology of Fishes 2^nd ed. Saunders College Publishing: Fort Worth.

[M79] Müller, A. H. 1979. Fossilization (taphonomy). In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A2–A78. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[P99] Poplin, C. 1999. Un paléoniscoïde (Pisces, Actinopterygii) de Buxières-les-Mines, témoin des affinités fauniques entre Massif central et Bohême au passage Carbonifère-Permien. Geodiversitas 21 (2): 147–155.

[PL00] Poplin, C., & R. Lund. 2000. Two new deep-bodied palaeoniscoid actinopterygians from Bear Gulch (Montana, USA, Lower Cretaceous). Journal of Vertebrate Paleontology 20 (3): 428–449.

[TT05] Tverdokhlebov, V. P., G. I. Tverdokhlebova, A. V. Minikh, M. V. Surkov & M. J. Benton. 2005. Upper Permian vertebrates and their sedimentological context in the South Urals, Russia. Earth-Science Reviews 69: 27–77.

Actinopterygii

Cheirolepis canadensis, from Parc national de Miguasha.

Belongs within: Gnathostomata.
Contains: Elonichthyiformes, Amphicentridae, Scanilepiformes, Actinopteri.

The Actinopterygii contain the ray-finned fishes in which the external fin is mostly composed of a ray-supported membrane. However, some basal members of this group (such as the modern Polypteridae) retain lobate pectoral fins. Basal ray-finned fishes (including the living Polypteriformes, Acipenseriformes and Holostei) have a covering of bony ganoid scales, often diamond-shaped and non-overlapping. Many fossil examples of these heavily armoured fishes have previously been classified in the Palaeoniscimorpha, now recognised as paraphyletic to more derived actinopteygians. The basalmost actinopterygians include the Devonian Cheirolepiformes which retained an appearance similar to related basal fishes in the Sarcopterygii and Acanthodii. The Saurichthyidae were long-bodied, long-jawed predatory fishes known from the late Permian to the mid-Jurassic.

Synapomorphies (from Xu & Gao 2011): Extrascapular paired; single pair only of nasal bones; squamosal absent; dentary with enclosed mandibular canal; presupracleithrum present; caudal fin heterocercal, upper caudal fin rays short.

<==Actinopterygii [Palaeoniscimorpha]
    |--Meemannia Zhu, Yu et al. 2006 GX17, ZY06
    |    `--*M. eos Zhu, Yu et al. 2006 ZY06
    `--+--Cheirolepiformes TT05
       |    |--Samarichthys [Karaunguriidae] TT05
       |    |    `--S. luxus Minikh 1990 TT05
       |    |--Boreolepis [Boreolepedidae] TT05
       |    |    |--B. jenseni Aldinger 1937 G93
       |    |    `--B. tataricus Esin 1996 TT05
       |    `--Cheirolepis Agassiz 1835 XG11, P99 [Cheirolepidae]
       |         |--C. trailli Agassiz 1835 GX17, CDZ93
       |         `--+--C. canadensis Whiteaves 1881 GX17, CDZ93
       |            `--C. schultzei GX17
       `--+--+--Osorioichthys [Osorioichthyidae] GX17
          |  |    `--O. marginis (Casier 1952) G93
          |  `--Tegeolepis [Tegeolepididae] GX17
          |       `--T. clarki (Newberry 1888) G93
          `--+--+--Mimipiscis GX17
             |  |    |--M. toombsi GX17
             |  |    `--+--M. bartrami GX17
             |  |       `--Gogosardina coatesi GX17
             |  |--+--Donnrosenia schaefferi GX17
             |  |  `--Mimiidae G93
             |  |       |--Mimia toombsi Gardiner & Bartram 1977 G93
             |  |       `--Howqualepis rostridens Long 1988 GX17, G93
             |  `--+--Raynerius splendens GX17
             |     `--Stegotrachelidae G93
             |          |--Orvikuina Gross 1953 G93
             |          |--Stegotrachelus Woodward & White 1926 G93
             |          |    `--S. finlayi GX17
             |          `--Moythomasia Gross 1950 GX17, G93
             |               |--M. durgaringa GX17
             |               |--M. lineata GX17
             |               |--M. nitida Gross 1953 P99
             |               `--M. perforata GX17
             `--+--+--Woodichthys bearsdeni GX17
                |  |--Melanecta anneae GX17
                |  `--Elonichthyiformes GX17
                `--+--Pholidopleuridae [Pholidopleuriformes] G93
                   |    |--Arctosomus sibiricus Berg 1941 G93
                   |    |--Pholidopleurus typus Bronn 1858 G93
                   |    |--Australosomus Piveteau 1930 GX17, G93
                   |    |    `--A. kochi GX17
                   |    `--Macroaethes Wade 1932 G93
                   |         `--M. brookvalei Wade 1932 F71
                   `--+--+--Amphicentridae GX17
                      |  `--+--Fouldenia ischiptera GX17
                      |     `--Styracopteridae G93
                      |          |--Styracopterus fulcatus Traquair 1890 GX17, G93
                      |          |--Whiteichthys Moy-Thomas 1942 G93
                      |          `--Chichia gracilis Liu & Wang 1978 G93
                      `--+--+--Scanilepiformes GX17
                         |  `--Actinopteri GX17
                         `--Saurichthyidae [Belonorhynchidae, Saurichthyiformes] G93
                              |--Belonorhynchus gigas Woodward 1890 F71
                              |--Eosaurichthys Liu & Wel 1988 G93
                              |--Saurorhynchus brevirostris (Woodward 1895) G93
                              `--Saurichthys Agassiz 1834 GX17, G93
                                   |--S. madagascarensis GX17
                                   `--S. parvidens F71

Actinopterygii incertae sedis:
  Amphistichus B02
    |--A. argenteus B02
    |--A. heermanni Girard 1854 DS89
    `--A. rhodoterus B78
  Eutrigla gurnardus OC03
  Pseudocaranx dentex OC03
  Spectrunculus grandis OC03
  Lepidotus notidanus BO02
  Niwaella delicata T90
  Squalidus T90
    |--S. chankaensis T90
    |    |--S. c. chankaensis T90
    |    `--S. c. biwae T90
    |--S. gracilis T90
    `--S. japonicus T90
  Pseudogobio T90
    |--P. esocinus T90
    `--P. rivularis T90
  Leucopsarion petersi T90
  Hemibarbus T90
    |--H. barbus T90
    `--H. longirostris T90
  Biwa zezera T90
  Odonthestes bonariensis T90
  Pungtungia herzi T90
  Ischikauia steenackeri T90
  Chirolophus nugator B96
  Haplopagrus guentheri B96
  Hoplerythrinus unitaeniatus B96
  Sicyases sanguinus B96
  Pyrrhulina filamentosa B96
  Xenocharax spilurus B96
  Genyonemus lineatus B96
  Tautagolabrus adspersus B96
  Xererpes fucorum B96
  Mylopharyngodon piceus B96
  Salminus maxillosus B96
  Steindachnerella Eiglemann 1897 B78
  Neoclinus blanchardi B78
  Damalichthys vacca B78
  Pahaerodon furcatus B78
  Apomotis cyanellus D51
  Clevelandia ios CH97
  Lionurus carapinus CH97
  Usclasichthys Heyler 1977 P99
    `--*U. macrodens Heyler 1977 P99
  Gyrolepis Agassiz 1833 P99
  Crinodus lophodon CC98
  Zalembius rosaceus AFL84
  Leptatherina wallacei MG06
  Pseudogobius olorum MG06
  Eleginus navaga W81
  Horalabiosa joshuai AJ02
  Sillaginodes punctatus S45
  Limichthys polyactis HS01
  Parika scaber HS01
  Obliquichthys maryannae HS01
  Grahamina capito HS01
  Notolabrus celidotus HS01
  Notoclinops segmentatus HS01
  Ruanoho whero HS01
  Trachylochismus melobesia HS01
  Hymenophysa curta USDI77
  Acanthorutilus handlirschi USDI77
  Moapa coriacea USDI77
  Probarbus jullieni USDI77
  Plagopterus argentissimus USDI77
  Brazilichthys macrognathus Cox & Hutchinson 1991 WO06
  Sargus PP89
    |--S. annularis D56
    |--S. rondeletii PP89
    |--S. salviani D56
    `--S. vulgaris D56
  Corvina PP89
    |--C. chrysoleuca G66
    |--C. nigra PP89
    |--C. ronchus G66
    `--C. vermicularis G66
  Gymnocharacinus bergi Steindachner 1903 RGG06
  Gymnetrus banksii T66
  Meletta libertatis G66
  Plagiogeneion rubiginosus P49
  Oxygaster PP07
    |--O. baccaila PP07 [=Chela bacaila PH72]
    `--O. clupeoides RM72
  Catla catla PP07
  Mystus CS77
    |--M. aor DR72
    |--M. cavasius PP07
    |--M. gulio CS77
    |--M. seenghala DR72
    `--M. vittatus PP07
  Gadusia chaptra PP07
  Poeciliops M68
    |--P. formosa M68
    |--P. latipinna M68
    `--P. sphenops M68
  Discordichthyidae [Discordichthyiformes] TT05
    |--Discordichthys spinifer (Minikh 1998) TT05
    `--Geryonichthys TT05
         |--G. burchardi Minikh 1998 TT05
         `--G. longus Minikh 1998 TT05
  Auchenoceros punctatus H01
  Pseudophycis bachus H01
  Austrophycis marginata H01
  Lepidorhynchus denticulatus H01
  Amphichthys cryptocentrus Valenciennes in Cuvier & Valenciennes 1837 BC01
  Joturus pichardi R86
  Polysteganus caeruleopunctatus Klunzinger 1871 S63
  Pneumatophorus grex P61
  Mollienisia F16b
    |--M. latipinna F15
    `--M. sphenops F16b
  Eupomotis CS77
    |--E. gibbosus CS77
    `--E. holbrooki F15
  Hepatus Gronow 1765 [incl. Teuthis Linnaeus 1766, Theutis] JR10
    |--H. bahianus F15
    |--H. celebicus JR10
    |--H. coeruleus F16b
    |--H. elongatus JR10
    |--H. hepatus F15
    |--H. matoides JR10
    `--H. olivaceus JR10
  Glaridichthys uninotatus F15
  Vomer F16b
    |--V. setapinnis F16b
    `--V. spixii F15
  Bathystoma F16b
    |--B. rimator F15
    `--B. striatum F16b
  Priapichthys annectens F15
  Coptostomobarbus vittei CS77
  Lophopsetta maculata F16a
  Pygosteus pungitius F16a
  Lasiancistrus guacharote [=Ancistrus guacharote; incl. A. trinitatis Regan 1906] F16b
  Odontostilbe pulcher (Gill 1858) [=Poecilurichthys pulcher, Chirodon pulcher] F16b
  Stevardia altipinnis Gill 1858 (see below for synonymy) F16b
  Hoploerythrinus unitaeniatus [=Erythrinus unitaeniatus; incl. E. cinereus Gill 1858] F16b
  Seserinus paru F16b
  Petrometopon cruentatus F16b
    |--P. c. cruentatus F16b
    `--P. c. coronatus [=Serranus coronatus; incl. Epinephelus guttatus Boulenger 1895] F16b
  Eudulus auriga [=Serranus auriga] F16b
  Brachygenys F16b
    |--B. chrysargyreus (Günther 1859) [=Haemulon chrysargyreum] F16b
    `--B. rimator F16b
  Corvula subaequalis [incl. C. sanctaeluciae Jordan 1889] F16b
  Philypnus dormitor F16b
  Alphestes chloropterus [incl. Epinephelus afer Boulenger 1895] F16b
  Myriapristis jacobus F16b
  Angelichthys isabelita F16b
  Colocopus lambdurus Gill 1884 JR10
  Prontocanthus sahelalmae Davis 1887 [=Holosteus sahelalmae] P05
  Porocottus allisi SSE10
  Opisthocentrus ocellatus SSE10
  Davidijordania lacertina SSE10
  Cuneoganthus gerdineri GX17
  Krasnoyarichthys jesseni GX17
  Limnomis delayni GX17
  Novagonatodus kasantsevae GX17
  Paedocypris progenetica SBM11
  Vallecillichthys multivertebratum VF21
  Robertichthys riograndensis VF21
  Esomus RS72
    |--E. altus [=E. danrica var. alta] RS72
    `--E. danricus RS72 [=E. danrica PH72]
         |--E. d. danricus RS72
         |--E. d. jabalpurensis Rao & Sharma 1972 RS72
         `--E. d. malabaricus RS72
  Amblypharyngodon mola PH72
  Eutroplus suratensis RM72
  Potamalosa novahollandiae O97
  Amphacanthus hexagonatus O97
  Batrachus trispinosus O97
  Crepidogaster tasmaniensis O97

Stevardia altipinnis Gill 1858 [incl. Corynopoma riisei Gill 1858, C. riisii, Nematopoma searlesii Gill 1858, C. veedoni Gill 1858] F16b

*Type species of generic name indicated

REFERENCES

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[BC01] Boyko, C. B., & J. R. Cordeiro. 2001. Catalog of Recent type specimens in the Division of Invertebrate Zoology, American Museum of Natural History. V. Mollusca, part 2 (class Gastropoda [exclusive of Opisthobranchia and Pulmonata], with supplements to Gastropoda [Opisthobranchia], and Bivalvia). Bulletin of the American Museum of Natural History 262: 1–170.

[B78] Brusca, R. C. 1978. Studies on the cymothoid fish symbionts of the eastern Pacific (Crustacea: Isopoda: Cymothoidae) II. Systematics and biology of Lironeca vulgaris Stimpson 1857. Occasional Papers of the Allan Hancock Foundation New Series 2: 1–19.

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[JR10] Jordan, D. S., & R. E. Richardson. 1910. Check-list of the species of fishes known from the Philippine archipelago. Bureau of Printing: Manila.

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[P99] Poplin, C. 1999. Un paléoniscoïde (Pisces, Actinopterygii) de Buxières-les-Mines, témoin des affinités fauniques entre Massif central et Bohême au passage Carbonifère-Permien. Geodiversitas 21 (2): 147–155.

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[T66] Tristram, H. B. 1866. Notice of the capture of a ribbonfish (Gymnetrus banksii) off the coast of Durham. Proceedings of the Zoological Society of London 1866: 147.

[TT05] Tverdokhlebov, V. P., G. I. Tverdokhlebova, A. V. Minikh, M. V. Surkov & M. J. Benton. 2005. Upper Permian vertebrates and their sedimentological context in the South Urals, Russia. Earth-Science Reviews 69: 27–77.

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[VF21] Vullo, R., E. Frey, C. Ifrim, M. A. González González, E. S. Stinnesbeck & W. Stinnesbeck. 2021. Manta-like planktivorous sharks in Late Cretaceous oceans. Science 371: 1253–1256.

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[XG11] Xu, G.-H., & K.-Q. Gao. 2011. A new scanilepiform from the Lower Triassic of northern Gansu Province, China, and phylogenetic relationships of non-teleostean Actinopterygii. Zoological Journal of the Linnean Society 161: 595–612.

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Last updated: 18 December 2021.

Teleostei

Fossil of Pholidophorus bechei in the Urwelt-Museum Hauff Holzmaden, photographed by Ghedoghedo.

Belongs within: Neopterygii.
Contains: Pachycormidae, Elopomorpha, Osteoglossocephalai.

The Teleostei are a clade including the majority of living ray-finned fishes. Teleosts are characterised by the presence of one or more uroneurals, modifications of the vertebral neural arches that support the caudal fin. Early members include the Mesozoic Pholidophoriformes, small fishes up to forty centimetres long that retain plesiomorphic features such as ganoid scales and enamel on the skull bones. The Triassic Leptolepidae possess more ossified vertebrae than Pholidophoriformes and bear cycloid scales (Bond 1996).

Synapomorphies (from Friedman et al. 2010): Internal carotid foramen present through parasphenoid; symplectic articulation on inner medial surface of quadrate; uroneurals present; pectoral propterygium fused to first pectoral fin-ray.

Teleostei [Archaeophylaces, Isospondyli, Malacopterygii, Teleosteomorpha]
    |--+--Pachycormidae FS10
    |  |--Tetragonolepis semicincta GX17
    |  `--Dapediidae G93
    |       |--Dapedium pholidotum A01, GX17
    |       |--Dandya White & Moy-Thomas 1940 G93
    |       |--Prohalecites Deeke 1889 G93
    |       `--Heterostrophus Wagner 1860 G93
    `--+--Ichthyokentemidae [Catervariolidae, Galkiniidae] P93
       |    |--Ichthyokentema purbeckensis (Davis 1887) GX17, P93
       |    |--Catervariolus G88
       |    |--Elpistoichthys pectinatus Griffith 1977 P93
       |    `--Ceramurus macrocephalus Egerton 1854 P93
       |--Pholidophoriformes B96
       |    |  i. s.: Seefeldia watsoni D01
       |    |         Pholidolepis africana D01
       |    |--Pleuropholis P93 [Ligulellidae, Majokiidae, Pleuropholidae G88]
       |    |    |--P. decastroi Bravi 1991 P93
       |    |    `--P. thiollieri Sauvage 1883 P93
       |    `--Pholidophoridae [Archaeomenidae] AS09
       |         |--Pholidophoretes salvus Griffith 1977 P93
       |         |--Gurvanichthys mongoliensis V-RR93
       |         |--Archaeomaene tenuis Woodward 1895 F71
       |         |--Neopholidophoropsis serrata Taverne 1981 P93
       |         |--Wadeichthys oxyops Waldman 1971 P93
       |         |--Pholidophoristion P93
       |         |    |--P. ornatus (Agassiz 1844) P93
       |         |    `--P. spaethi Taverne 1981 P93
       |         `--Pholidophorus FS10
       |              |--P. bechei FS10
       |              |--P. germanicus FS10
       |              |--P. latisculus A01
       |              `--P. macrocephalus FS10
       `--+--+--Dorsetichthys bechei GX17
          |  `--Leptolepidae [Koonwarriidae, Leptolepididae, Leptolepidiformes, Leptolepimorpha] D01
          |       |--Koonwarria manifrons Waldman 1971 P93
          |       |--Proleptolepis Nybelin 1974 D01
          |       |    |--P. elongata D01
          |       |    |--P. furcata D01
          |       |    `--P. megalops D01
          |       `--Leptolepis A01
          |            |--L. coryphaenoides FS10
          |            |--L. koonwarri Waldman 1971 P93
          |            |--L. normandica D01
          |            `--L. talbragarensis Woodward 1895 [incl. L. gregarius Woodward 1895, L. lowei Woodward 1895] F71
          `--+--Varasichthyidae A01
             `--+--Teleocephala [Elopocephala] A01
                |    |  i. s.: Protostomiatidae P93
                |    |           |--Protostomias maroccanus Arambourg 1954 P93
                |    |           `--Pronotacanthus Woodward 1900 P93, W00
                |    |                `--*P. sahelalmae (Davis 1887) [=Anguilla sahelalmae] W00
                |    |--Elopomorpha FS10
                |    `--Osteoglossocephalai FS10
                `--Ichthyodectidae [Icthyodectiformes, Ichthyodectoidei] A01
                     |--Xiphactinus Leidy 1870 P93, FS10
                     |--Gillicus P93
                     |--Cladocyclus G88
                     |    |--C. gardneri BWW93
                     |    `--C. sweeti F71
                     `--Thrissops P93
                          |--T. formosus Agassiz 1833 P93
                          `--T. subovatus Münster 1833 P93

Teleostei incertae sedis:
  Ascalabos viothii FS10
  Telestes pleurobipunctatus KKV03
  Membras martinicus K-M02
  Tselfatoidiformes B96
  Platypoecilus maculatus D37
  Palaeolabrus [Palaeolabridae] G88
  Siyuichthyidae P93
  Allothrissops [Allothrissopidae] P93
    |--A. mesogaster (Agassiz 1833) P93
    |--A. regleyi (Thollière 1854) P93
    `--A. salmoneus (Blainville 1818) P93
  Saurodontidae P93
    |--Saurodon intermedius (Newton 1878) P93
    `--Saurocephalus lanciformis Harlan 1824 P93
  Molinichthys inopinatus Gayet 1982 P93
  Protaulopsis bolcensis Woodward 1901 P93
  Aidachar paludalis Nessov 1981 B93
  Neocybium rostratum Leriche 1908 F91

*Type species of generic name indicated

REFERENCES

[AS09] Alfaro, M. E., F. Santini, C. Brock, H. Alamillo, A. Dornburg, D. L. Rabosky, G. Carnevale & L. J. Harmon. 2009. Nine exceptional radiations plus high turnover explain species diversity in jawed vertebrates. Proceedings of the National Academy of Sciences of the USA 106 (32): 13410–13414.

[A01] Arratia, G. 2001. The sister-group of Teleostei: consensus and disagreements. Journal of Vertebrate Paleontology 21 (4): 767–773.

[B93] Benton, M. J. 1993. Reptilia. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 681–715. Chapman & Hall: London.

[B96] Bond, C. E. 1996. Biology of Fishes 2^nd ed. Saunders College Publishing: Fort Worth.

[BWW93] Briggs, D. E. G., M. J. Weedon & M. A. Whyte. 1993. Arthropoda (Crustacea excluding Ostracoda). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 321–342. Chapman & Hall: London.

[D01] Delsate, D. 2001. L’ichthyofaune du Pliensbachian (Jurassique inferieur) de Lorraine et des Ardennes (France): premiers resultats. Bulletin de l’Académie Lorraine des Sciences 40 (1–2): 47–69.

[D37] Dobzhansky, T. 1937. Genetics and the Origin of Species. Columbia University Press: New York.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[F91] Freess, W. B. 1991. Beiträge zur Kenntnis von Fauna und Flora des marinen Mitteloligozäns bei Leipzig. Altenburger Naturwissenschaftliche Forschungen 6: 3–74.

[FS10] Friedman, M., K. Shimada, L. D. Martin, M. J. Everhart, J. Liston, A. Maltese & M. Triebold. 2010. 100-million-year dynasty of giant planktivorous bony fishes in the Mesozoic seas. Science 327: 990–993.

[G93] Gardiner, B. G. 1993. Osteichthyes: basal actinopterygians. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 611–619. Chapman & Hall: London.

[GX17] Giles, S., G.-H. Xu, T. J. Near & M. Friedman. 2017. Early members of 'living fossil' lineage imply later origin of modern ray-finned fishes. Nature 549: 265–268.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[K-M02] Klein-MacPhee, G. 2002. Jacks. Family Carangidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3^rd ed. pp. 411–427. Smithsonian Institute Press: Washington.

[KKV03] Kontula, T., S. V. Kirilchik & R. Väinölä. 2003. Endemic diversification of the monophyletic cottoid fish species flock in Lake Baikal explored with mtDNA sequencing. Molecular Phylogenetics and Evolution 27: 143–155.

[P93] Patterson, C. 1993. Osteichthyes: Teleostei. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 621–656. Chapman & Hall: London.

[V-RR93] Vickers-Rich, P., & T. H. Rich. 1993. The ICI Australia Catalogue of the Great Russian Dinosaurs Exhibition 1993–1995. Monash University.

[W00] Woodward, A. S. 1900. Evidence of an extinct eel (Urenchelys anglicus, sp. n.) from the English Chalk. Annals and Magazine of Natural History, series 7, 5: 321–323, pl. 9.

Last updated: 13 February 2022.