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Holocephalomorpha
Taxonomy
Holocephalomorpha was named by Lund (1986). It is not extant.
It was reranked as the infraclass Holocephalimorpha by Lund and Grogan (1997); it was misspelled as Holocephalimorpha by Lund and Grogan (2004), Goto and Okura (2004), Nelson (2006), Long (2011), Gouiric-Cavalli et al. (2015), Popov et al. (2020); it was corrected as Holocephalimorpha by Nelson et al. (2016), Ebersole et al. (2022), Popov et al. (2025).
It was assigned to Chondrichthyes by Lund and Grogan (1997), Lund and Grogan (2004); to Holocephali by Gouiric-Cavalli et al. (2015); and to Holocephali by Goto and Okura (2004), Nelson (2006), Long (2011), Nelson et al. (2016), Popov et al. (2020), Ebersole et al. (2022), Popov et al. (2025).
It was reranked as the infraclass Holocephalimorpha by Lund and Grogan (1997); it was misspelled as Holocephalimorpha by Lund and Grogan (2004), Goto and Okura (2004), Nelson (2006), Long (2011), Gouiric-Cavalli et al. (2015), Popov et al. (2020); it was corrected as Holocephalimorpha by Nelson et al. (2016), Ebersole et al. (2022), Popov et al. (2025).
It was assigned to Chondrichthyes by Lund and Grogan (1997), Lund and Grogan (2004); to Holocephali by Gouiric-Cavalli et al. (2015); and to Holocephali by Goto and Okura (2004), Nelson (2006), Long (2011), Nelson et al. (2016), Popov et al. (2020), Ebersole et al. (2022), Popov et al. (2025).
Synonymy list
| Year | Name and author |
|---|---|
| 1986 | Holocephalomorpha Lund |
| 1997 | Holocephalimorpha Lund and Grogan |
| 2004 | Holocephalomorpha Goto and Okura p. 221 |
| 2004 | Holocephalimorpha Lund and Grogan |
| 2006 | Holocephalimorpha Nelson p. 44 |
| 2011 | Holocephalomorpha Long p. 242 |
| 2015 | Holocephalomorpha Gouiric-Cavalli et al. p. 4 |
| 2016 | Holocephalimorpha Nelson et al. p. 50 |
| 2020 | Holocephalomorpha Popov et al. p. 5 |
| 2022 | Holocephalimorpha Ebersole et al. p. 373 |
| 2025 | Holocephalimorpha Popov et al. |
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If no rank is listed, the taxon is considered an unranked clade in modern classifications. Ranks may be repeated or presented in the wrong order because authors working on different parts of the classification may disagree about how to rank taxa.
Superor. †Holocephalomorpha Lund 1986
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Or. Chimaeriformes Obruchev 1953 [chimaera]
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G. †Arctacanthus Nielsen 1932
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†Arctacanthus exiguus Yamagishi 2004
†Arctacanthus multisectus Branson 1933
†Arctacanthus phosphoriensis Branson 1930
†Arctacanthus spitzbergensis Hulke 1873
†Arctacanthus uncinatus Nielsen 1932
†Arctacanthus wyomingensis Branson 1934
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Invalid names: Dolophonodus uncinatus Branson 1933 [replaced]
Invalid names: Ancistriodus Branson 1933 [synonym], Dolophonodus Branson 1933 [synonym], Hamatus Branson 1930 [synonym]
Fm. Callorhynchidae Garman 1901 [plownose chimaera]
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Subfm. Callorhynchinae Garman 1901
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G. †Ptyktoptychion Lees 1986
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†Ptyktoptychion tayyo Lees 1986
†Ptyktoptychion wadeae Bartholomai 2008
G. †Elasmodus Egerton
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†Elasmodus avirostrus Averianov 1999
†Elasmodus greenoughi Agassiz 1838
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Invalid names: Aptychus crassus Hebert 1855 [synonym], Elasmodus crassus Leriche 1906 [synonym], Elasmodus planus Leriche 1929 [synonym]
†Elasmodus kempi Ward 1977
†Elasmodus khosatzkyi Averianov 1999
†Elasmodus rossicus Averianov 1999
†Elasmodus sinzovi Averianov and Glickman 1994
†Elasmodus tsheganicus Averianov 2001
†Elasmodus zharyk Averianov 1999
Invalid names: Psaliodus Agassiz 1843 [synonym]
G. †Ischyodus Egerton 1843
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†Ischyodus avitus Meyer 1800
†Ischyodus beaumonti Egerton 1843
†Ischyodus bifurcatus Case 1978
†Ischyodus bohemicus Bayer 1905
†Ischyodus dufrenoyi Egerton 1843
†Ischyodus egertoni Buckland 1835
†Ischyodus emarginatus Egerton 1843
†Ischyodus incisus Newton 1878
†Ischyodus latus Newton 1878
†Ischyodus mortoni Chapman and Pritchard 1907
†Ischyodus panus Newton 1878
†Ischyodus quenstedti Wagner 1857
†Ischyodus rayhaasi Hoganson and Erickson 2005
†Ischyodus thurmanni Pictet and Campiche 1878
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Invalid names: Ischyodus brevirostris Newton 1876 [synonym]
†Ischyodus townsendi Buckland 1835
†Ischyodus yanschini Averianov 1991
†Ischyodus zinsmeisteri Applegate 1975
Invalid names: Edaphodontinae Stahl 1999 [empty]
Fm. Chimaeridae
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G. Chimaera Linnaeus 1758
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Subg. †Chimaera (Ischyodon) Agassiz 1843
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†Chimaera (Ischyodon) beaumontii Agassiz 1843
†Chimaera (Ischyodon) brevirostris Agassiz 1843
†Chimaera (Ischyodon) dufrenoyi Agassiz 1843
†Chimaera (Ischyodon) dutertrii Agassiz 1843
†Chimaera (Ischyodon) emarginata Agassiz 1843
†Chimaera (Ischyodon) helvetica Agassiz 1843
†Chimaera (Ischyodon) johnsonii Agassiz 1843
†Chimaera (Ischyodon) tessoni Buckland 1835
†Ischyodon bucklandi Agassiz 1843
Subg. †Chimaera (Psittacodon) Agassiz 1843
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†Chimaera (Psittacodon) falcata Agassiz 1843
†Chimaera (Psittacodon) mantellii Buckland 1835
†Chimaera (Psittacodon) psttacina Agassiz 1843
†Chimaera furcata Fritsch 1878
†Chimaera pliocenica Woodward 1891
†Chimaera venablesae Cassier 1966
G. †Hedronchus Cope 1876
G. †Hydrolagus Gill 1862
G. †Leptomylus Cope 1869
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†Leptomylus cookii Cope 1870
†Leptomylus densus Cope 1869
†Leptomylus forfex Cope 1875
Subor. Chimaeroidei
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G. †Anteliochimaera Tanaka 1909
Fm. Callorhinchidae Garman 1901 [plownose chimaera]
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G. Callorhinchus Lacépède 1798
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†Callorhinchus alfordi Cicimurri and Ebersole 2015
†Callorhinchus phillipsi Cicimurri and Ebersole 2015
Callorhinchus stahli Kriwet and Gazdizicki 2003
G. †Ganodus Agassiz 1843
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†Chimaera (Ganodus) colei Buckland 1835
†Chimaera (Ganodus) curvidens Agassiz 1843
†Chimaera (Ganodus) neglecta Agassiz 1843
†Chimaera (Ganodus) owenii Buckland 1835
†Chimaera (Ganodus) rugulosa Agassiz 1843
†Ganodus dentatus Egerton 1847
†Ganodus oweni Agassiz 1843
†Ganodus rugulosus Egerton 1847
G. †Pachymylus Woodward 1892
Superfm. Callorhinchoidea
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G. †Chalcodus Zittel 1887
Superfm. Chimaeroidea
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G. †Chimaerotheca Brown 1946
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†Chimaerotheca alaskana Brown 1946
†Chimaerotheca besselsi Brown 1946
†Chimaerotheca germanica Brown 1946
†Chimaerotheca montanana Brown 1946
†Chimaerotheca newmexicana Brown 1946
†Chimaerotheca reperere Gottfried and Fordyce 2015
†Chimaerotheca wyomingana Brown 1946
Fm. †Edaphodontidae Owen 1846
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G. †Edaphodon Buckland 1838
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Invalid names: Diphrissa Cope 1875 [synonym], Dipristis Marsh [synonym]
G. †Eumylodus Leidy 1873
G. †Crassidonta Branson 1916
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†Crassidonta stuckenbergi Branson 1916
†Crassidonta subcrenulata Teichert 1943
G. †Icthypriapus
Subor. †Myriacanthoidei Patterson 1965
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Fm. †Chimaeropsidae
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Fm. †Myriacanthidae Woodward 1889
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G. †Halonodon Duffin 1984
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†Halonodon luxembourgensis Duffin and Delsate 1993
†Halonodon warneri Duffin 1984
G. †Myriacanthus Agassiz 1836
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†Myriacanthus retrorsus Agassiz 1837
Invalid names: Prognathodus Egerton 1872 [synonym]
Invalid names: Acanthorhinidae Patterson 1965 [synonym]
Invalid names: Chimeroidei Patterson 1965 [empty], Myracanthiodei [empty], Squalorajoidei Patterson 1965 [empty]
Or. †Chondrenchelyiformes Moy-Thomas 1939
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Fm. †Chondrenchelyidae Berg 1940
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G. †Platyxystrodus Hay 1899
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†Platyxystrodus bellulus St John and Worthen 1875
†Platyxystrodus imitatus St John and Worthen 1875
†Platyxystrodus inconditus Saint John and Worthen 1883
†Platyxystrodus occidentalis St. John 1870
†Platyxystrodus simplex St John and Worthen 1875
†Platyxystrodus striatus M'Coy 1855
†Platyxystrodus verus John and Worthen 1883
Invalid names: Xystrodus Morris and Roberts 1862 [replaced]
Or. †Cochliodontiformes Obruchev 1953
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Fm. †Cochliodontidae Owen 1867
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G. †Aspidodus Newberry and Worthen 1866
G. †Chitonodus Stjohn and Worthen 1883
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†Chitonodus antiquus Saint John and Worthen 1883
†Chitonodus liratus Saint John and Worthen 1883
†Chitonodus springeri Saint John and Worthen 1883
G. †Cochliodus Agassiz 1838
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†Cochliodus acutus Agassiz 1843
†Cochliodus contortus Agassiz 1837
†Cochliodus latus Leidy 1857
†Cochliodus leidyi Saint John and Worthen 1883
†Cochliodus occidentalis Leidy 1857
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Invalid names: Deltodus stellatus Newberry and Worthen 1866 [synonym]
†Cochliodus vanhornii Saint John and Worthen 1883
G. †Cranodus Trautschold 1879
G. †Cyrtonodus Davis 1884
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†Cyrtonodus brevis Goto and Okura 2004
†Cyrtonodus fukujiensis Goto and Okura 2004
†Cyrtonodus hornei Woodward 1889
†Cyrtonodus nishioi Goto and Okura 2004
G. †Deltodopsis St John and Worthen 1883
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†Deltodopsis affinis Saint John and Worthen 1883
†Deltodopsis angustus Newberry and Worthen 1870
†Deltodopsis bialveatus Saint John and Worthen 1883
†Deltodopsis convolutus Saint John and Worthen 1883
†Deltodopsis exornatus Saint John and Worthen 1883
†Deltodopsis inflexus Saint John and Worthen 1883
†Deltodopsis sanctiludovici Saint John and Worthen 1883
G. †Deltodus Morris and Roberts 1862
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†Deltodus alatus Newberry and Worthen 1870
†Deltodus angularis Newberry and Worthen 1866
†Deltodus cinctulus Saint John and Worthen 1883
†Deltodus cinctus Newberry 1879
†Deltodus cingulatus Newberry and Worthen 1866
†Deltodus concha Trautschold 1874
†Deltodus contortus Stjohn and Worthen 1883
†Deltodus costatus Newberry and Worthen 1870
†Deltodus fasciatus Newberry and Worthen 1870
†Deltodus gibbus Woodward 1889
†Deltodus grandis Newberry 1879
†Deltodus inornatus Newberry 1897
†Deltodus intermedius Saint John and Worthen 1883
†Deltodus latior Saint John and Worthen 1883
†Deltodus littoni Newberry and Worthen 1870
†Deltodus mercurei Newberry 1876
†Deltodus obliquus Saint John and Worthen 1883
†Deltodus parvus Saint John and Worthen 1883
†Deltodus planidens Cope 1894
†Deltodus powellii Saint John and Worthen 1883
†Deltodus propinquus Saint John and Worthen 1883
†Deltodus rugosus Woodward 1889
†Deltodus spatulatus Newberry and Worthen 1866
†Deltodus sublaevis Agassiz 1843
†Deltodus trilobus Saint John and Worthen 1883
†Deltodus undulatus Newberry and Worthen 1866
G. †Dichelodus Giebel 1857
G. †Erismacanthus M'Coy 1848
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†Erismacanthus barbatus Eastman 1903
†Erismacanthus formosus Eastman 1902
†Erismacanthus jonesii M'Coy 1848
†Erismacanthus maccoyanus Stjohn and Worthen 1875
†Erismacanthus major Davis 1883
G. †Icanodus Miller 1892
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†Icanodus convexus Davis 1883
†Icanodus limitaris St John and Worthen 1875
Invalid names: Tomodus Davis 1883 [replaced]
G. †Orthopleurodus St John and Worthen 1880
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†Orthopleurodus convexus Saint John and Worthen 1883
†Orthopleurodus novomexicanus Saint John and Worthen 1883
G. †Oxytomodus Trautschold 1880
G. †Periplectrodus St John and Worthen 1875
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†Periplectrodus compressus Saint John and Worthen 1875
†Periplectrodus expansus Saint John and Worthen 1875
†Periplectrodus warreni Saint John and Worthen 1875
G. †Pleuroplax Woodward 1889
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†Pleuroplax attheyi Barkas 1874
†Pleuroplax rankinei Hancock and Atthey 1843
†Pleuroplax woodi Davis 1883
G. †Poecilodus Agassiz 1843
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†Poecilodus aliformis M'Coy 1848
†Poecilodus carbonarius Saint John and Worthen 1883
†Poecilodus cestriensis Saint John and Worthen 1883
†Poecilodus convolutus Newberry and Worthen 1870
†Poecilodus foveolatus M'Coy 1848
†Poecilodus jonesii Agassiz 1843
†Poecilodus oliquus Agassiz 1843
†Poecilodus paradoxus Waagen 1879
†Poecilodus parallelus Agassiz 1843
†Poecilodus rugosus Newberry and Worthen 1866
†Poecilodus sanctiludovici Saint John and Worthen 1883
†Poecilodus springeri Saint John and Worthen 1883
†Poecilodus tribulis Stjohn and Worthen 1883
†Poecilodus varsoviensis Saint John and Worthen 1883
†Poecilodus wortheni Saint John 1883
G. †Rhamphodus Davis 1883
G. †Sandalodus Newberry and Worthen 1866
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†Sandalodus angustus Newberry and Worthen 1866
†Sandalodus carbonarius Newberry and Worthen 1866
†Sandalodus complanatus Newberry and Worthen 1866
†Sandalodus crassus Newberry and Worthen 1866
†Sandalodus elllipticus Newberry 1897
†Sandalodus laevissimus Newberry and Worthen 1866
†Sandalodus minor Newberry and Worthen 1866
†Sandalodus morrisii Davis 1883
†Sandalodus parvulus Newberry and Worthen 1866
†Sandalodus rhomboideus Newberry and Worthen 1866
G. †Stenopterodus St John and Worthen 1883
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†Stenopterodus elongatus Saint John and Worthen 1883
†Stenopterodus planus Saint John and Worthen 1883
G. †Streblodus Agassiz 1843
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†Streblodus angustus Eastman 1903
†Streblodus colei Davis 1883
†Streblodus egertoni Davis 1883
†Streblodus oblongus Agassiz 1843
G. †Taeniodus St John and Worthen 1883
G. †Trigonodus Newberry and Worthen 1866
G. †Vaticinodus St John and Worthen 1883
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†Vaticinodus carbonarius Saint John and Worthen 1883
†Vaticinodus discrepans Saint John and Worthen 1883
†Vaticinodus lepis Saint John and Worthen 1883
†Vaticinodus similis Saint John and Worthen 1883
†Vaticinodus simplex Saint John and Worthen 1883
†Vaticinodus vetustus Saint John and Worthen 1883
G. †Venustodus St. John and Worthen 1875
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†Venustodus argutus St John and Worthen 1875
†Venustodus leidyi St. John and Worthen 1875
†Venustodus robustus Saint John and Worthen 1875
†Venustodus tenuicristatus Saint John and Worthen 1875
†Venustodus variabilis Saint John and Worthen 1875
†Venustodus venustus Leidy 1857
Fm. †Psephodontidae Zangerl 1981
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G. †Psephodus Morris and Roberts 1862
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†Psephodus crenulatus Newberry and Worthen 1866
†Psephodus dubius Woodward 1889
†Psephodus latus Saint John and Worthen 1883
†Psephodus levissimus Agassiz 1837
†Psephodus lunulatus Saint John and Worthen 1883
†Psephodus magnus Agassiz 1843
†Psephodus obliquus Saint John and Worthen 1883
†Psephodus placenta Newberry and Worthen 1866
†Psephodus politus Newberry 1897
†Psephodus regularis Saint John and Worthen 1883
†Psephodus reticulatus Saint John and Worthen 1875
†Psephodus salopiensis Woodward 1889
†Psephodus symmetricus Saint John and Worthen 1883
Invalid names: Psephodus depressus Waagen 1880 [nomen vanum], Psephodus indicus Waagen 1880 [nomen vanum]
Or. †Copodontiformes Obruchev 1953
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Fm. †Copodontidae Davis 1883
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G. †Copodus Morris and Roberts 1862
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†Copodus auriculatus Davis 1883
†Copodus cornutus Davis 1843
†Copodus furcatus Davis 1862
†Copodus lingua Davis 1883
†Copodus minimus Davis 1883
†Copodus oblongus Davis 1883
†Copodus pusillus Saint John and Worthen 1883
†Copodus spatulatus Davis 1862
†Copodus spatulatus Davis 1862
†Copodus vanhornii Saint John and Worthen 1883
Or. †Menaspiformes
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Fm. †Deltoptychiidae Zangerl 1981
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G. †Deltoptychius Morris and Roberts 1862
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†Deltoptychius acutus Agassiz 1843
†Deltoptychius expansus Saint John and Worthen 1883
†Deltoptychius gibberuius Davis 1883
†Deltoptychius nitidus Leidy 1857
†Deltoptychius primus Saint John and Worthen 1883
†Deltoptychius varsoviensis Saint John and Worthen 1883
†Deltoptychius wachsmuthi Saint John and Worthen 1883
Fm. †Traquairiidae Lund and Grogan 1997
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G. †Traquairius Lund and Grogan 1997
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†Traquairius agkistrocephalus Lund and Grogan 1997
†Traquairius nudus Lund and Grogan 1997
†Traquairius spinosus Lund and Grogan 1997
Or. †Psammodontiformes Obruchev 1953
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Fm. †Psammodontidae
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G. †Apodometes Gistel 1848
G. †Astrabodus Davis 1884
G. †Dimyleus Davis 1883
G. †Homalodus Davis 1883
G. †Mylacodus Davis 1883
G. †Mylax Morris and Roberts 1883
G. †Pinacodus Davis 1883
G. †Pleurogomphus Agassiz 1862
G. †Psammodus Agassiz 1834
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†Psammodus angularis Newberry and Worthen 1866
†Psammodus caelatus Saint John and Worthen 1883
†Psammodus canaliculatus M'Coy 1848
†Psammodus cornutus Agassiz 1843
†Psammodus crassidens Saint John and Worthen 1883
†Psammodus expansus Davis 1884
†Psammodus glyptus Saint John and Worthen 1883
†Psammodus grandis Saint John and Worthen 1883
†Psammodus longidens Agassiz 1837
†Psammodus lovianus Saint John and Worthen 1883
†Psammodus plenus Saint John and Worthen 1883
†Psammodus porosus Agassiz 1837
†Psammodus reticulatus Newberry and Worthen 1866
†Psammodus rugosus Agassiz 1836
†Psammodus salopiensis Woodward 1889
†Psammodus springeri Saint John and Worthen 1883
†Psammodus trapeziformis Davis 1883
†Psammodus tumidus Saint John and Worthen 1883
†Psammodus turgidus Saint John and Worthen 1883
†Psammodus turgidus Agassiz 1838
G. †Rhymodus Davis 1883
Or. †Squalorajiformes Patterson 1965
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Fm. †Squalorajidae Woodward 1886
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G. †Squaloraja Riley 1833
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†Squaloraja polyspondyla Agassiz 1836
†Squaloraja tenuispina Woodward 1886
Invalid names: Spinacorhinus Agassiz 1836 [synonym]
Diagnosis
No diagnoses are available
Measurements
No measurements are available
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| Source: o = order, subp = subphylum | |||||
| References: Carroll 1988, Hendy et al. 2009, Wagner 2023 | |||||
Age range
Maximum range based only on fossils: base of the Givetian to the top of the Early/Lower Pliocene or 387.95000 to 3.60000 Ma
Minimum age of oldest fossil (stem group age): 382.31 Ma
Minimum age of oldest fossil (stem group age): 382.31 Ma
Collections (353 total)
Oldest occurrences | |||
| Time interval | Ma | Country or state | Original ID and collection number |
|---|---|---|---|
| Devonian | Poland | Psephodus sp. (236675) | |
| Givetian | France (Nord-Pas-de-Calais) | Melanodus loonesi (90628) | |
| Late/Upper Devonian | USA (California) | Cochliodontiformes indet. (206454) | |
| Kinderhookian | USA (Montana) | Psammodus sp. (545) | |
| Early/Lower Mississippian | USA (Iowa) | Erismacanthus barbatus (133258) | |
| Mississippian | United Kingdom | Cochliodus magnus (132784) Erismacanthus jonesi, Copodus cornutus, Copodus spatulatus, Copodus furcatus, Mesogomphus lingua, Pleurogomphus auriculatus, Psammodus trapeziformis, Psephodus magnus, Xystrodus striatus, Paecilodus jonesii, Paecilodus gibbosus, Streblodus oblongus, Streblodus colei, Deltoptychius acutus, Erismacanthus major, Cochliodus contortus (137333) Poecilodus foveolatus (132729) Poecilodus sublaevis, Helodus turgidus, Psammodus rugosus, Cochliodus acutus, Cochliodus magnus, Cochliodus oblongus, Cochliodus striatus, Erismacanthus jonesii, Poecilodus jonesii, Poecilodus oliquus, Poecilodus parallelus, Psammodus canaliculatus, Psammodus cornutus, Cochliodus contortus (132781) Psammodus rugosus (132789) Psephodus salopiensis, Psephodus dubius, Sandalodus morrisii (137332) | |
| Mississippian | USA (Alabama) | Psammodus sp. (84103) | |
| Carboniferous | USA (Arizona) | Psephodus sp. (236494) | |
| Osagean | USA (Indiana) | Sandalodus sp. (6670) | |
| Osagean | USA (Iowa) | Deltodus spatulatus (133270) Deltodus spatulatus, Deltodus occidentalis (133094) | |
| Arundian | United Kingdom (Scotland) | Deltoptychius armigerus, Chondrenchelys problematica (90682) | |
| Chadian | United Kingdom (England) | Psephodus magnus (5970) | |
| Visean | Belgium | Taeniodus contortus (133095) | |
| Visean | United Kingdom | Cochliodus magnus, Cochliodus striatus, Poecilodus jonesii, Psammodus cornutus (132783) | |
| Visean | Russian Federation (Komi) | Venustodus sp. (133366) | |
| Visean | Egypt | Psammodus sp., Psephodus sp. (46983) | |
| Asbian | United Kingdom (Northern Ireland) | Psammodus sp. (6949) | |
| Asbian - Brigantian | United Kingdom (England) | Psephodus magnus (5986) | |
| Brigantian | Thailand | Psammodontidae indet. (130649) | |
| Late/Upper Mississippian | USA (Alabama) | Psammodus sp. (84104) | |