Reconstruction of Lasanius problematicus, copyright Nobu Tamura.

Belongs within: Deuterostomia.
Contains: Cyclostomata, Conodonta, Birkeniida, Thelodontomorphi, Pteraspidomorphi, Galeaspida, Osteostraci, Gnathostomata.

The Craniata are a major subgroup of the chordates in which the neural tube is enlarged to form a brain that is protected by an endoskeletal cranium. Anterior sensory structures such as eye lenses and an olfactory epithelium are also developed in the head region. In development, these head structures are formed from ectodermal placodes while neural crest cells give rise to the dermal skeleton, skeletal material of jaws and branchial arches, trabeculae, parts of sensory capsules, pigment cells, some cardiac and aortic-arch motor neurons and some sensory cranial nerves. Most craniates also possess an internal skeleton with vertebrae, characterising the clade Vertebrata.

Our understanding of higher-level phylogeny within living craniates is relatively stable except for the position of the Myxinidae (hagfish). Morphological studies place hagfish, which possess a cranium but lack vertebrae, outside the vertebrates, but molecular studies place hagfish as sister to Petromyzontidae (lampreys), implying that their lack of some vertebrate synapomorphies may represent a secondary loss.

Craniata [Agnatha, Craniota, Pisces, Tetraploblastica] SCM03
    |--Haikouichthys Luo, Hu & Shu in Shu, Luo et al. 1999 SCM03, SL99
    |    `--*H. ercaicunensis Luo, Hu & Shu in Shu, Luo et al. 1999 SL99
    `--Vertebrata (see below for synonymy) BJ06
         |  i. s.: Boothialepis thorsteinssoni Märss 1999 SM01
         |         Canonia grossi MG01
         |         Jianichthys longicephalus ZBH03
         |         Longdeichthys lujiaxiaensis ZBH03
         |         Yanosteus longidorsalis ZBH03
         |         Irenichthys S02
         |         Paralogania BMM02
         |           |--P. kummerowi (Gross 1967) BMM02
         |           `--P. ludlowiensis Gross 1967 SM01
         |         Fagea Buen 1940 C92
         |         Lophacanthus Stock 1880 C92
         |         Pezopallichthys [Furcacaudiformes] SM01
         |           `--P. ritchiei Wilson & Caldwell 1998 SM01
         |         Angaralepis moskalenkoae K-TS03
         |         Skiichthys halsteadi K-TS03
         |         Achanarella Newman 2002 [Achanarellidae] N02
         |           `--*A. trewini Newman 2002 N02
         |         Hanilepis wangi W97
         |         Gantarostrataspis geni W97
         |         Machaerasthus bohemicus W97
         |         Clarorbis apponomedianus JP97
         |         Lepidorhombus bosci PP64
         |         Gastrobranchus Bloch 1797 B97
         |           `--*G. coecus Bloch 1797 B97
         |         Pituriaspida J93
         |           |--Pituriaspis doylei J93
         |           `--Neeyambaspis J93
         |         Pircanchaspis rinconensis Schultze in Erdtmann et al. 2000 TBN04
         |         Areyongalepis oervigi TBN04
         |         Proceros Rafinesque 1820 G00
         |         Myllokunmingia Shu, Zhang & Han in Shu, Luo et al. 1999 SL99
         |           `--*M. fengjiaoa Shu, Zhang & Han in Shu, Luo et al. 1999 SL99
         |--Cyclostomata MW07
         `--+--Conodonta GCR06
            `--+--Jamoytius [Jamoytiidae] GCR06
               |    `--J. kerwoodi White 1946 BMM02
               |--Euphaneropidae N02
               |    |--Legendrelepis N02
               |    |--Euphanerops Woodward 1900 MS16, W00
               |    |    `--*E. longaevus Woodward 1900 W00
               |    `--Cornovichthys Newman & Trewin 2001 N02
               |         `--C. blaauweni Newman & Trewin 2001 N02
               `--+--Anaspida [Anaspidiformes] GCR06
                  |    |  i. s.: Ctenopleuron nerepisense Matthew 1907 BMM02
                  |    |--Birkeniida BMM02
                  |    `--Lasanius Traquair 1898 [Lasaniidae] BMM02
                  |         |--L. armatus Traquair 1898 BMM02
                  |         `--L. problematicus Traquair 1898 BMM02
                  `--+--Thelodontomorphi MS16
                     `--+--Pteraspidomorphi MS16
                        `--+--Galeaspida GCR06
                           `--+--Osteostraci GCR06
                              `--Gnathostomata MW07

Craniata incertae sedis:
  Acus Müller 1774 BR05
  ‘Acus’ Swainson 1839 non Müller 1774 BR05
  ‘Gonostoma’ van Hasselt 1823 non Rafinesque 1810 BR05
  Aetheolepis mirabilis Woodward 1895 F71
  Apateolepis australis Woodward 1890 F71
  Aphelepis australis Woodward 1895 F71
  Madariscus robustus (Woodward 1895) [=Archaeomaene robustus] F71
  Belichthys minimus F71
  Corunegenys bowralensis Wade 1941 F71
  Enigmatichthys attenuatus Wade 1935 F71
  Gosfordia truncata Woodward 1890 F71
  Myriolepis F71
    |--M. clarkei F71
    `--M. lata Woodward 1890 [=M. latus] F71
  Proctotretus stantoni Girard 1854 DS89
  Homalopomanus trowbridgii Girard 1856 DS89
  Holconotus trowbridgii Girard 1858 DS89
  Metaspriggina Simonetta & Insom 1993 MS16, W96
  Chilodus Muller & Troschel 1844 BR17
  Zalophancylus morani Hannibal 1912 [=Lanx (Zalophancylus) morani] TTE93
  Taraxes Day 1881 K18
  Pseudogastromyzon myersi B93
  Morulius chrysophekadion B93
  Tanichthys chrysophekadion B93

Vertebrata [Birkeniiformes, Cephalaspidomorphi, Diplorhina, Hyperoartii, Jamoytiiformes, Monorhina, Myopterygii, Nantes] BJ06

*Type species of generic name indicated


[B93] Beresford, W. A. 1993. Cranial skeletal tissues: diversity and evolutionary trends. In: Hanken, J., & B. K. Hall (eds) 1993. The Skull vol. 2. Patterns of Structural and Systematic Diversity pp. 69–130. The University of Chicago Press.

[B97] Bloch, M. E. 1797. Sur le Gastrobranchus, nouveau genre de poisson. Bulletin des Sciences, par la Societé Philomathique de Paris 1 (4): 26.

[BMM02] Blom, H., T. Märss & C. G. Miller. 2002. Silurian and earliest Devonian birkeniid anaspids from the Northern Hemisphere. Transactions of the Royal Society of Edinburgh: Earth Sciences 92: 263–323.

[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.

[BR17] Bouchet, P., J.-P. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A. Nützel, P. Parkhaev, M. Schrödl & E. E. Strong. 2017. Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia 61 (1–2): 1–526.

[BJ06] Bourlat, S. J., T. Juliusdottir, C. J. Lowe, R. Freeman, J. Aronowicz, M. Kirschner, E. S. Lander, M. Thorndyke, H. Nakano, A. B. Kohn, A. Heyland, L. L. Moroz, R. R. Copley & M. J. Telford. 2006. Deuterostome phylogeny reveals monophyletic chordates and the new phylum Xenoturbellida. Nature 444 (7115): 85–89.

[C92] Crawford, R. L. 1992. Catalogue of the genera and type species of the harvestman superfamily Phalangioidea (Arachnida). Burke Museum Contributions in Anthropology and Natural History 8: 1–60.

[DS89] Dundee, H. A., & H. M. Smith. 1989. Nomenclatural eponym endings revisited. Systematic Zoology 38 (3): 279–283.

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

[GCR06] Gess, R. W., M. I. Coates & B. S. Rubidge. 2006. A lamprey from the Devonian period of South Africa. Nature 443: 981–984.

[G00] Grubb, P. 2000. Valid and invalid nomenclature of living and fossil deer, Cervidae. Acta Theriologica 45 (3): 289–307.

[J93] Janvier, P. 1993. Patterns of diversity in the skull of jawless fishes. In: Hanken, J., & B. K. Hall (eds) 1993. The Skull vol. 2. Patterns of Structural and Systematic Diversity pp. 131–188. The University of Chicago Press.

[JP97] Ji S. & Pan J. 1997. The macropetalichthyids (Placodermi) from Guangxi and Hunan, China. Vertebrata PalAsiatica 35 (1): 18–34.

[K-TS03] Karatajute-Talimaa, V., & M. M. Smith. 2003. Early acanthodians from the Lower Silurian of Asia. Transactions of the Royal Society of Edinburgh: Earth Sciences 93: 277–299.

[K18] Kury, A. B. 2018. Familial nomina in harvestmen (Arachnida, Opiliones). Bionomina 13: 1–27.

[MW07] Mallatt, J., & C. J. Winchell. 2007. Ribosomal RNA genes and deuterostome phylogeny revisited: more cyclostomes, elasmobranchs, reptiles, and a brittle star. Molecular Phylogenetics and Evolution 43: 1005–1022.

[MG01] Märss, T., & P.-Y. Gagnier. 2001. A new chondrichthyan from the Wenlock, Lower Silurian, of Baillie-Hamilton Island, the Canadian Arctic. Journal of Vertebrate Paleontology 21 (4): 693–701.

[MS16] McCoy, V. E., E. E. Saupe, J. C. Lamsdell, L. G. Tarhan, S. McMahon, S. Lidgard, P. Mayer, C. D. Whalen, C. Soriano, L. Finney, S. Vogt, E. G. Clark, R. P. Anderson, H. Petermann, E. R. Locatelli & D. E. G. Briggs. 2016. The 'Tully monster' is a vertebrate. Nature 532: 496–499.

[N02] Newman, M. J. 2002. A new naked jawless vertebrate from the Middle Devonian of Scotland. Palaeontology 45 (5): 933–941.

[PP64] Peres, J. M., & J. Picard. 1964. Nouveau manuel de bionomie benthique de la mer Mediterranee. Recueil des Travaux de la Station Marine d'Endoume, Bulletin 31 (27): 5–137.

[SCM03] Shu, D.-G., S. Conway Morris, J. Han, Z.-F. Zhang, K. Yasul, P. Janvier, L. Chen, X.-L. Zhang, J.-N. Liu, Y. Li & H.-Q. Liu. 2003. Head and backbone of the Early Cambrian vertebrate Haikouichthys. Nature 421: 526–529.

[SL99] Shu, D.-G., H.-L. Luo, S. C. Morris, X.-L. Zhang, S.-X. Hu, L. Chen, J. Han, M. Zhu, Y. Li & L.-Z. Chen. 1999. Lower Cambrian vertebrates from south China. Nature 402: 42–46.

[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.

[SM01] Soehn, K. L., T. Märss, M. W. Caldwell & M. V. H. Wilson. 2001. New and biostratigraphically useful thelodonts from the Silurian of the Mackenzie Mountains, Northwest Territories, Canada. Journal of Vertebrate Paleontology 21: 651–659.

[TTE93] Tracey, S., J. A. Todd & D. H. Erwin. 1993. Mollusca: Gastropoda. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 131–167. Chapman & Hall: London.

[TBN04] Turner, S., A. Blieck & G. S. Nowlan. 2004. Vertebrates (agnathans and gnathostomes). In: Webby, B. D., F. Paris, M. L. Droser & I. G. Percival (eds) The Great Ordovician Biodiversification Event pp. 327–335. Columbia University Press.

[W97] Wang N. 1997. Restudy of thelodont microfossils from the lower part of the Cuifengshan Group of Qujing, eastern Yunnan, China. Vertebrata PalAsiatica 35 (1): 1–17.

[W00] Woodward, A. S. 1900. On a new ostracoderm (Euphanerops longaevus) from the Upper Devonian of Scaumenac Bay, Province of Quebec, Canada. Annals and Magazine of Natural History, series 7, 5: 416–419, pl. 10.

[ZBH03] Zhou, Z., P. M. Barrett & J. Hilton. 2003. An exceptionally preserved Lower Cretaceous ecosystem. Nature 421: 807–814.

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