After Lloyd, Wang & Brusatte, 2012
Note: “Ceratodiformes” = The least inclusive clade containing †Ceratodus latissimus Agassiz, Neoceratodus forsteri (Krefft), Lepidosiren paradoxa Fitzinger and Protopterus annectens Owen.
<==o Ceratodiformes [incl. Lepidosireniformes; Ceratodontoidei Vorobyeva & Obruchev, 1964; Ceratodontidae Gill, 1872 sensu Carroll, 1988] (modern lungfishes; nykykeuhkokalat l. australiankeuhkokalat & siimaeväkeuhkokalat) |--+-- †Parasagenodus sibiricus Vorobyeva, 1972 | `--+--o †Archaeoceratodus Kemp, 1997b | | |-- †A. avus (Woodward, 1906) | | |-- †A. djelleh (Kemp, 1982) [Neoceratodus djelleh Kemp, 1982] | | |-- †A. theganus Kemp, 1997b | | `-- †A. rowleyi Kemp, 1997b | `--o †Ceratodontidae Gill, 1872 sensu Kemp, 1997 | |--o †Metaceratodus (Chapman, 1914) Schultze, 1992 [Ceratodus (Metaceratodus) Chapman, 1914] Aust. | | |-- †M. wollastoni Kemp, 1997a [type] | | |-- †M. ellioti Kemp, 1997 | | |-- †M. palmeri (Krefft, 1874) Kemp, 1997 [Ceratodus palmeri Krefft, 1874; Neoceratodus forsteri (Krefft, 1870) de Vis, 1884; Epiceratodus palmeri (Krefft, 1874) Hills, 1934] | | |-- †M. bonei Kemp, 1997 | | |-- †M. kaopen (Apesteguía, Agnolin & Claeson, 2007) Cione & Gouiric-Cavalli, 2012 [Ceratodus kaopen Apesteguía, Agnolin & Claeson, 2007; Ceratodus kaopen Agnolin, 2010; M. kaopen Cione & Gouiric-Cavalli, 2012] | | `-- †M. wichmanni (Apesteguía, Agnolin & Claeson, 2007) Cione & Gouiric-Cavalli, 2012 [Ceratodus iheringi Ameghino, 1898 (in partim); Ceratodus sp. Wichmann, 1927; Ceratodus iheringi Pascual & Bondesio, 1976; ‘Ceratodus’ iheringi Cione, 1987; ‘Ptychoceratodus’ iheringi Arratia & Cione, 1996; Ceratodus sp. González-Riga, 1999; Ptychoceratodus wichmanni Apesteguía, Agnolin & Claeson, 2007; Ptychoceratodus cionei Apesteguía, Agnolin & Claeson, 2007; Ferganoceratodus wichmanni Agnolin, 2010; Metaceratodus wichmanni Cione & Gouiric-Cavalli, 2012] | `--+-- †Grossipterus crassus (Gross, 1933) Vorobyeva & Obruchev, 1964 [Dipterus crassus Gross, 1933] | `--o †Ceratodus Agassiz, 1838 [Hemictenodus; Metaceratodus; Tellerodus Lehman, 1975; Scropha] L. Trias. U. Jur. Eu. L. Trias. L. Cret. As. L. Trias. U. Cret. Mad. NA. Aus. L. Trias. Paleoc. Af. L. Trias. Spits. | `-- †C. latissimus Agassiz, 1838 [C. bicornis Agassiz, 1838-44] [type] `--+--o †Gnathorhiza Cope, 1883; Penn. L. Perm. NA. U. Perm. Eu. [Gnathorhizidae Miles, 1977] | |?- †G. flavosus (Cope, 1884) [Proceratodus flavosus] | |-- †G. serrata Cope, 1883 | |-- †G. dikeloda Olson, 1951 | `-- †G. noblensis Olson & Daly, 1972 `--+--+-- †Microceratodus angolensis (Texeira, 1947) Teixeira, 1954 [Ceratodus angolensis] Trias. NAf. L.Trias. Mad. | `--+-- †Aphelodus anapes Kemp, 1993 | `--+-- †Palaeophichthys parvulus (a terrasiiform?) | `--+--o †Ptychoceratodus Jaekel, 1926 [Ceratodus (in partim)]; U.Trias. Eu. [Ptychoceratodontidae Martin, 1982b] | | |?- †P. philippsi | | |?- †P. rectangulus | | `-- †P. serratus (Agassiz, 1838) [Ceratodus serratus Agassiz, 1838] | `--+-- †Paraceratodus germaini Lehman, Chateau, Laurain & Nauche, 1959; L. Trias. Mad. | `--+--o †Ferganoceratodus Kaznyshkin & Nessov, 1985; U.Jur. SEAs. | | |-- †F. szechuanensis | | `-- †F. martini Cavin, Suteethorn, Buffetaut & Tong, 2007 | `--o †Arganodus Martin, 1979; U.Trias. NAf. | |-- †A. atlanticus Martin, 1979 [Asiatoceratodus atlantis Kemp, 1998] | `-- †A. tiguidensis [Ceratodus tiguidiensis] `--+-- †Beltanodus ambilobensis (Beltan, 1968) Schultze, 1981 `--+-- †Namatozodia pitikanta Kemp, 1993; Trias. Aust. `--+-- †Ariguna formosa (Wade, 1935) Kemp, 1994 [= Ceratodus formosus Wade, 1935; Tellerodus formosus Martin, 1982b; Ceratodus formosus Kemp, 1991; Ceratodus formosa Lloyd, Wang & Brusatte, 2012] `--+-- †Tellerodus sturii (Teller, 1891) Lehman, 1975 [Ceratodus sturii Teller, 1891] `--+--+-- †Gosfordia truncata Woodward, 1890; L. Trias. Aus. | `-- †Asiatoceratodus sharovi Vorobyeva, 1967 [Asiatoceratodontidae Vorobyeva, 1967] `==o Neoceratodontidae Miles, 1977 [Ceratodidae] |?- †Equinoxiodus alcantarensis Toledo, de Sousa, Medeiros & Bertini, 2011; U.Cret. NESAm. |--o Neoceratodus de Castelnau, 1876 [Epiceratodus; ?Ompax] U.Cret.-R. Aust. (australiankeuhkokalat) | |-- †N. nargun (Kemp, 1983) Kemp, 1997b [Ceratodus nargun Kemp, 1983] U.Cret. SEAust. | |-- †N. denticulatus (Agassiz) ?Eoc. NEAust. | |-- †N. eyrensis (White, 1925) Kemp, 1997b [Epiceratodus eyrensis White, 1925] | |-- †N. djelleh Kemp, 1982 | `-- N. forsteri (Krefft, 1870) [Ceratodus forsteri Krefft, 1870] (Australian lungfish; australiankeuhkokala) `--+--o †Mioceratodus Kemp, 1992; U. Olig. L. Mioc. Aus. | |-- †M. anemosyrus Kemp, 1992 | |-- †M. gregoryi (White, 1925) ["Epiceratodus" gregoryi White, 1925; Neoceratodus gregoryi (White, 1925)] | |-- †M. diaphorus Kemp, 1997b | `-- †M. poastrus Kemp, 1997b `--o Lepidosireniformes Berg, 1940 sensu Longrich, 2017 |?- †Retodus tuberculatus (Tabaste, 1963) Churcher, De Iuliis & Kleindienst, 2006 [Ceratodus tuberculatus Tabaste, 1963, Cappetta 1972, Kemp 1997, Lehman 1966, Martin 1984b, Schultze 1992; Neoceratodus tuberculatus (Tabaste) Martin 1982a, 1982b, Schaal 1984, Churcher & De Iuliis, 2001; “Neoceratodus tuberculatus” (Tabaste) Churcher 1999] U. Cret. NAf. |?- †“Neoceratodus” africanus |?-o †Lavocatodidae Longrich, 2017 | |-- †?Ceratodus diutinus Kemp, 1993 | |-- †Lavocatodus? sp. [Kem Kem] | |-- †Lavocatodus protopteroides (Tabaste, 1963) Claeson, Sallam, O'Connor & Sertich, 2014 [“Ceratodus” protopteroides Tabaste, 1963; “Protopterus” protopteroides (Tabaste); Lavocatodus? protopteroides Longrich, 2017] | |-- †Lavocatodus humei (Priem, 1914) Claeson, Sallam, O'Connor & Sertich, 2014 [Ceratodus? humei Priem, 1914; “Protopterus” humei (Priem, 1914); Lavocatodus? humei Longrich, 2017] | `--+-- †Lavocatodus giganteus Martin, 1995 | `-- †Xenoceratodus labyrinthus Longrich, 2017; Eoc. NAf. `--o Lepidosirenidae Bonaparte, 1841 [non Woodward, 1891][incl. Protopteridae] (siimaeväkeuhkokalat; Aestivating lungfishes) |-- †Lepidosiren? sp. [Fajum] |-- †Protopterus? sp. [Adrar Mgom] |-- †Protopterus? elongus |-- †Protopterus? sp. [Dur at Talah] |-- †Protopterus? nigeriensis Martin, 1997 |-- Lepidosiren paradoxa Fitzinger, 1837 [Amphibichthys] Mioc. R. SA. [Lepidosirenidae] (eteläamerikankeuhkokalat; South American lungfish) `--o Protopterus Owen, 1839 [Protomalus, Rhinocryptis] U.Cret. Eoc. R. Af. [Protopteridae] (afrikankeuhkokalat) |-- †P. sp. [Fayum] |-- †P. regulatus |-- †P. crassidens Churcher & de Iuliis, 2001 |--o P. aethiopicus Heckel, 1851 (ethiopiankeuhkokala) | |-- P. a. aethiopicus Heckel, 1851 (Marbled lungfish) | |-- P. a. congicus Poll, 1961 | `-- P. a. mesmaekersi Poll, 1961 |-- P. amphibius (Peters, 1844) (sambesinkeuhkokala; Gilled lungfish) |--o P. annectens Owen, 1839 (afrikankeuhkokala) | |-- P. a. annectens (Owen, 1839) (West African lungfish) | `-- P. a. brieni Poll, 1961 (Southern lungfish) `-- P. dolloi Boulenger, 1900 (kongonkeuhkokala; Slender lungfish)
Ceratodiformes incertae sedis:
†Atlantoceratodus patagonicus Agnolin, 2010
†Chaoceratodus portezuelensis Apesteguía et al. 2007
†Potamoceratodus guentheri (Marsh, 1878) Pardo, Huttenlocker, Small & Gorman, 2010 [Ceratodus guentheri Marsh, 1878]
†Ceratodus Agassiz, 1838 [Hemictenodus; Metaceratodus; Tellerodus Lehman, 1975; Scropha] L. Trias. U. Jur. Eu. L. Trias. L. Cret. As. L. Trias. U. Cret. Mad. NA. Aus. L. Trias. Paleoc. Af. L. Trias. Spits.
†C. acutus
†C. africanus [Neoceratodus africanus sensu Churcher & de Iuliis, 2001, incl. C. pectinatus Tabaste, 1963; C. brasiliensis de Sousa Cunha & Simoes Ferreira, 1980]
†C. altus
†C. americanus
†C. arganensis
†C. argentinus Apesteguía et al. 2007
†C. bovisrivi
†C. capensis
†C. concinnus
†C. crosbiensis
†C. cruciferus
†C. curvus
†C. daedaleus
†C. disauris
†C. donensis
†C. emarginatus
†C. eruciferus
†C. felchi
†C. frazieri
†C. gibbus
†C. gustasoni
†C. heshanggouensis
†C. heteromorphus
†C. hieroglyphus
†C. hislopianus
†C. hunterianus
?†C. iheringi
†C. kaupii [Ceratodus kaupi]
†C. nageshwarai
†C. obtusus
†C. ornatus
†C. parvus
†C. planasper
†C. planus
†C. rectangularis
†C. robustus
†C. rucinatus
†C. szechuanensis
†C. vinslovii
†C. virapa
After Long, 1995, Cloutier & Ahlberg, 1996, and Kemp, 1998<==o ?Ceratodiformes |?- †Arganodus |?- †Gosfordia |?- †Ferganoceratodus |?- †Microceratodus |?- †Paraceratodus |--o Ceratodiformes sensu stricto?? | |?-o †Asiatoceratodontidae | | `--o †Asiatoceratodus | | |-- †A. sharovi | | `-- †A. atlantis | `--+?-o †Ptychoceratodontidae | | `--o †Ptychoceratodus | | |?- †P. philippsi | | |?- †P. rectangulus | | `-- †P. serratus | `--+?-o Ceratodontidae Gill, 1872 | | `--o Ceratodontinae Gill, 1872 | | |?- †“Ceratodus” africanus | | |-- †Retodus tuberculatus | | |--o †Ceratodus | | | |-- †C. latissimus | | | |-- †C. nargun | | | |-- †C. humei | | | |-- †C. sturii | | | |?- †D. diutinus | | | `?- †C. iheringi | | `--o †Metaceratodus | | |-- †M. wollastoni | | |-- †M. ellioti | | |-- †M. palmeri | | `-- †M. bonei | `--o Neoceratodontidae | |--o †Mioceratodus | | |-- †M. anemosyrus | | `-- †M. gregoryi | `--o Neoceratodus | |-- †N. denticulatus | |-- †N. eyrensis | `-- N. forsteri `--o Lepidosireniformes |?-o †Gnathorhizidae | |?- †Palaeophichthys (a terrasiiform?) | `--o †Gnathorhiza | |-- †G. flavosus | `-- †G. dikeloda `--o Lepidosirenidae |-- Lepidosiren paradoxa `--o Protopterus |-- †P. regulatus |-- †P. protopteroides |-- †P. crassidens |-- P. aethiopicus |-- P. amphibius |-- P. annectens `-- P. dolloi |
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