After Sansom, 2009 (fig. 5)
Alternative (older) phylogeny after Janvier, 1981, Carroll, 1988, Janvier, 1997 [“The Tree of Life”], Adrian & Wilson, 1994, and Mark-Kurik & Janvier, 1995
<==o Osteostraci Lankester, 1868 [Cephalaspidiformes] |--o †Atelaspididae Traquair, 1899 sensu Sansom, 2009 | |-- †Hirella gracilis [Micraspis] L. Dev. Eu. [paraphyletic Ateleaspidae] | `--o †Atelaspidinae Traquair, 1899 sensu Sansom, 2009 | |-- †Aceraspis robusta [Hemiteleaspis] [paraphyletic Ateleaspidae] | `-- †Atelaspis tessellata [paraphyletic Ateleaspidae] `--+-- †Hemicyclaspis murchisoni (Egerton, 1857) L. Dev. Eu. NNA. `--o †Cornuta Janvier, 1985b sensu Sansom, 2009 |--o †Cephalaspis [incl.? Alaspis] [Camptaspis; Escuminaspis; Eucephalaspis; Minestaspis; Pattenaspis; Scolenaspis] U. Sil. L. Dev. Eu. L.-M. Dev. Spits. L. Dev. EAs. L.-U. Dev. NA. [Cephalaspidida Stensiö, 1964] | |-- †C. lyelli Agassiz, 1835 | |-- †C. producta Wängsjö, 1952 | |-- †C. cradleyensis Stensiö, 1932 | |-- †C. sollasi Stensiö, 1932 | |?- †C. magnifica Obruchev, 1961 | |?- †C. microtuberculata [Alaspis] | `?- †C. powriei `--+--o †Zenaspidida Stensiö, 1958 sensu Sansom, 2009 | |?-o †Wladysagitta Voichyshyn, 2006; L.Dev. WEu. EEu. (sedis mutabilis) | | |-- †W. janvieri Voichyshyn, 2006 | | `-- †W. acutirostris (Stensiö, 1932) [†Cephalaspis acutirostris (Stensiö, 1932)] | |--o †Spangenhelmaspis Sansom, 2009 [Cephalaspis Agassiz, 1835; Stensiö, 1927 (in partim); Securiaspis Wängsjö, 1952 (in partim); Securiaspis janvier, 1985b (in partim)] | | |-- †S. staxrudi (Stensiö, 1927) [Cephalaspis staxrudi Stensiö, 1927] | | `-- †S. quadrata (Wängsjö, 1952) [Securiaspis quadrata Wängsjö, 1952] | |--o †Parametereoaspididae Sansom, 2009 | | |-- †Parametereoraspis gigas | | `--+?- †Trewinia (sedis mutabilis) | | |-- †Balticaspis latvica Lyarskaya, 1981 [incl.? Afanassievaspis Otto & Laurin, 2001] | | `--+--o †Escuminaspis [synon. of Cephalaspis] | | | `-- †E. laticeps (Traquair, 1880) [Cephalaspis rosamundae Roberts, 1937; Alaspis macrotuberculata Ørvik, 1957; †Alaspis rosamundae (juv.)] U.Dev. ENA. | | `-- †Levesquaspis patteni (Robertson, 1936) | `--o †Zenaspididae Stensiö, 1958 | |--+-- †Scolenaspis signata | | |-- †Ukrainaspis kozlowskii (Zych, 1937) (Sansom, 2009 [Cephalaspis Agassiz, 1835; Zych, 1937 (in partim); Cephalaspis Agassiz, 1835; Janvier, 1985b, 1985e (in partim); Heraspis Afanessieva, 1991, 2004 (preoccupied) non Megistaspis (Heraspis) Wandås, 1983, a trilobite subgenus] | | `--+--o †Machairaspis | | | |-- †M. ibex Wängsjö, 1952 | | | `-- †M. corystis Wängsjö, 1952 | | `-- †Superciliaspis gabrielsei (Dineley & Loeffler, 1976) Adrian & Wilson, 1994 [Cephalaspis? gabrielsei Dineley & Loeffler, 1976] | `--+--o †Tegaspis L.Dev. Spits. | | |-- †T. kolleri (Stensiö, 1927) | | `-- †T. pedata (Wängsjö, 1952) | `--+--o †Stensiopelta denison, 1951a; L.Dev. Eu. | | |-- †S. woodwardi (Stensiö, 1932) [Cephalaspis woodwardi Stensiö, 1932] | | `-- †S. pustulata Janvier, 1985b | `--+?- †“Diademaspis”? mackenziensis Adrian & Wilson, 1994 | |--o †Diademaspis Janvier, 1985b | | |-- †D. poplinae Janvier, 1985b | | |-- †D. jarviki (Wängsjö, 1952) Janvier, 1985b [Cephalaspis jarviki Wängsjö, 1952] | | `-- †D. stensioei Afanassieva, 1989 | `--o †Zenaspis Lankester, 1870 sensu Janvier, 1985b; L.Dev. NWEu. EEu. Spits. | |-- †Z. salweyi (Egerton, 1857) [†Cephalaspis salweyi Egerton, 1867] | |?- †Z. metopias (Wängsjö, 1952) | |-- †Z. podolica (Balabai, 1962) | `-- †Z. dzieduszyckii Voichyshyn, 2006 `--+--+?-+--o †Zychaspis Janvier, 1985; U.Sil. WEu., U.Dev. EEu. | | | |-- †Z. granulata Voichyshyn, 1998 | | | `-- †Z. siemiradzkii | | |--o †Pattenaspis [synon. of Cephalaspis?] | | | |-- †P. acuminata | | | |-- †P. deltoides | | | |-- †P. eurhynchus (Wängsjö, 1952) | | | `-- †P. rogalai (Balabai, 1962) | | `--+-- †Hildenaspis | | `-- †Mimetaspis hoeli | `--+--+--o †Waengsjoeaspis Janvier, 1985b | | | |-- †W. excellens (Wängsjö , 1952) Janvier, 1985b [†Cephalaspis excellens Wängsjö , 1952] | | | |-- †W. vogti (Stensiö, 1927) Janvier, 1985b [†Cephalaspis vogti Stensiö, 1927] | | | `-- †W. nahanniensis Adrian & Wilson, 1994 | | `--+-- †Camptaspis utahensis Branson & Mehl, 1931 | | `-- †Yvonaspis Otto & Laurin, 1999 | `--o †Benneviaspidida Denison, 1951a (non Halstead Tarlo, 1967) sensu Sansom, 2009 | |-- †Ectinaspis [Hoelaspis] L.Dev. Spits. | `--+--o †Securiaspis Stensiö, 1932; L.Dev. Eu. Spits. | | |-- †S. kitchini Stensiö, 1932 | | `-- †S. kingi Stensiö, 1932 | `--+-- †“Benneviaspis” longicornis | `--+--o †Benneviaspis Stebsiö, 1927 [incl. Citharaspis Afanassieva, 1989] L. Dev. Eu. Spits. [Benneviaspididae Denison, 1951] | | |-- †B. anglica Stensiö | | `--+-- †B. lankesteri Stensiö | | `--+-- †B. urvantsevi Mark-Kurik & Janvier, 1995; L. Dev. NAs. | | |-- †B. maxima | | |-- †B. holtedahli | | |-- †B. polonica Belles-Isles & Janvier, 1984 | | |-- †B. whitei Belles-Isles & Janvier, 1984 | | `-- †B. junia (Afanassieva, 1989) [Citharaspis junia Afanassieva, 1989] | `--o †Boreaspidoidei Stensiö, 1958 sensu Sansom, 2009 | |-- †Hoelaspis angulata Stensiö, 1927 | |-- †Severaspis rostralis Mark-Kurik & Janvier, 1995; L. Dev. NAs. | `--o †Boreaspididae Stensiö, 1958 | |-- †“Boreaspis” ceratops | |-- †“Boreaspis” intermedia | |--o †Boreaspis Stensiö, 1927 sensu stricto?; U.Dev. Spits. | | |-- †B. puella | | `== 12 other species? | |-- †Dicranaspis | `--+-- †Spatulaspis | `--+-- †Belonaspis | `--+-- †Hapilaspis aphales Mark-Kurik & Janvier, 1995; L. Dev. NAs. | `-- †Tauraspis rara Mark-Kurik & Janvier, 1995; L. Dev. NAs. `--o †Thyestiida Berg, 1940 sensu Sansom, 2009 [incl. Kiaeraspidida Stensiö, 1932; Tannuaspidida] |--o †Ilemoraspis L.Dev. EEu. | `-- †I. kirkinskayae Afanassieva, 1991 `--+-- †Procephalaspis U.Sil. Eu. `--+-- †Auchenaspis U.Sil. Eu. `--+--o †Thyestes [Auchenaspis] U.Sil.-L.Dev. Eu. [Thyestidae??] | `-- †T. verrucosus `--+-- †Witaaspis U. Sil. EEu. `--+--o †Kiaeraspidoidea (Stensiö, 1932) Afanassieva, 1991 | |-- †Didymaspis | `--o †Kiaeraspididae Stensiö, 1932 | |-- †Kieraspis L.Dev. Spits. | `--+?- †Norselaspis glacialis Janvier, 1981 (sedis mutabilis) | `--o †Axinaspidoidea Janvier, 1981a | |-- †Nectaspis L.-M.Dev. Spits. | `--+-- †Axinaspis L.Dev. Spits. | `--o †Acrotomaspidinae Janvier, 1981a | |-- †Acrotomaspis L.Dev. Spits. | `-- †Gustavaspis `--o †Tremataspidoidea (Woodward, 1891) Afanassieva, 1991 sensu Sansom, 2009 |--o †Tyriaspis U.Sil. Eu. | `-- †T. whitei |-- †Saaremaaspis [Dasylepis; ?Dictyolepis; Rotsikuellaspis obrutchevi Robertson, 1938] U. Sil. L. Dev. Eu. `--o †Tremataspididae Woodward, 1891 |-- †Aestiaspis |--o †Dartmuthia [Lophosteus] U.Sil. Eu. [Dartmuthiidae] | `-- †D. gemmifera `--+-- †Timanaspis U. Sil. EEu. L. Dev. Eu. `--+--o Tremataspidinae Woodward, 1891 | |-- †Oeselaspis [?Trachylepis] U.Sil. Eu. | `-- †Tremataspis [Odontododus, Stigmolepis] U. Sil. Eu. `--+-- †Dobraspis `--+-- †Tannuaspis levenkoi Obruchev, 1956 [incl. Tuvaspis margaritae Obruchev, 1956] L. Dev. NAs. `-- †Sclerodus [Eukeraspis] L. Dev. Eu. [Sclerodontidae]
†Osteostraci incertae sedis:
†Tuvaspis L. Dev. NAs. [paraphyletic Ateleaspidae]
†Hemiteleaspis heintzii Westoll, 1945 [=? Hemicyclaspis]
†Turinea [Cephalopterus] L.Dev. Eu.
Reference(s):
- Adrian, J. M. & Wilson, M. V. H., 1994: Early Devonian cephalaspids (Vertebrata: Osteostraci: Cornuata) from the southern Mackenzie Mountains, NorthWest Territories, Canada.
–Journal of Vertebrate Paleontology: Vol. 14, #3, pp. 301-319 - Carroll, R. L., 1988: Vertebrate paleontology and evolution.
–W. H. Freeman and company, New York, 1988, 698. - Carroll, R. L., 1988: Appendix. 594-648.
in Carroll, R. L., 1988: Vertebrate paleontology and evolution.
–W. H. Freeman and company, New York, 1988, 698. - Janvier, P., 1981: The phylogeny of the Craniata,
with particular referance to the significance of fossil "agnathans".
–Journal of Vertebrate Paleontology: vol. 1, #2, pp. 121-159. - Janvier , P., 1997: Vertebrata: Animals with backbones.
–iNet : Tree of Life; http://tolweb.org/tree/ - Janvier, P., Arsenault, M. & Desbiens, S., 2004: Calcified cartilage in the paired fins of the osteostracan Escuminaspis laticeps (Traquair, 1880), from the Late Devonian of Miguasha (Québec, Canada), with a consideration of the early evolution of the pectoral fin endoskeleton in vertebrates.
–Journal of Vertebrate Paleontology: Vol. 24, #4, pp. 773-779 - Lundberg, J. G., 1995, 1997: Chordata.
–iNet : Tree of Life; http://tolweb.org/tree/ - Long, J. A., 1995: The Rise of Fishes: 500 million years of evolution.
–Johns Hopkins University Press, Baltimore & London, pp. 1-223 - Mark-Kurik, E. & Janvier, P., 1995:
Early Devonian osteostracans from Severnaya Zemlya, Russia.
–Journal of Vertebrate Paleontology: Vol. 15, #3, pp. 449-462 - Nelson, J. S., 1994: Fishes of the world.
–John Wiley & Sons Inc., New York, 1994, xx-600 - Otto, M. & Laurin, M., 2001: Microanatomy of the dermal skeleton of Balticaspis latvica (Osteostraci, Middle Devonian).
–Journal of Vertebrate Paleontology: Vol. 21, #1, pp. 186-189 - Sansom, R. S., 2009: Phylogeny, classification and character polarity of the Osteostraci (Vertebrata).
–Journal of Systematic Palaeontology: Vol. 7, #1, pp. 95-115 - Voichyshyn, V., 2006: New osteostracans from the Lower Devonian terrigenous deposits of Podolia, Ukraine.
–Acta Palaeontologica Polonica: Vol. 51, #1, pp. 131-142 [http://app.pan.pl/acta51/app51-131.pdf]