Dinocephalia: Difference between revisions
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The infraorder Dinocephalia<ref name="Seeley 1894">Seeley, H.G. (1892) On ''Delphinognathus conocephalus'' (Seeley) from the middle Karroo beds, Cape Colony, preserved in the South African Museum, Cape Town. ''Quarterly Journal of the Geological Society, London.'' '''48''': 469-475.</ref> were a group of medium to large, heavily built therapsid mammal-like reptiles that lived during the Mid-Permian.<ref name="Rubidge 1991">Rubidge, B.S. (1991) A new primitive dinocephalian mammal-like reptile from the Permian of southern Africa. ''Palaeontology''. '''34''': 547-559.</ref><ref name="Kemp 2005">Kemp, T.S. (2005) ''The origin and evolution of mammals''. Oxford University Press, | The infraorder Dinocephalia<ref name="Seeley 1894">Seeley, H.G. (1892) On ''Delphinognathus conocephalus'' (Seeley) from the middle Karroo beds, Cape Colony, preserved in the South African Museum, Cape Town. ''Quarterly Journal of the Geological Society, London.'' '''48''': 469-475.</ref> were a group of medium to large, heavily built therapsid mammal-like reptiles that lived during the Mid-Permian.<ref name="Rubidge 1991">Rubidge, B.S. (1991) A new primitive dinocephalian mammal-like reptile from the Permian of southern Africa. ''Palaeontology''. '''34''': 547-559.</ref><ref name="Kemp 2005">Kemp, T.S. (2005) ''The origin and evolution of mammals''. Oxford University Press, | ||
Oxford.</ref> Dinocephalians are among the earliest group of [[Therapsida|therapsids]], and have been found in Brazil<ref name="Langer 2000">Langer, M.C. (2000) The first record of dinocephalians in South America, Late Permian | Oxford.</ref> Dinocephalians are among the earliest group of [[Therapsida|therapsids]], and have been found in Brazil<ref name="Langer 2000">Langer, M.C. (2000) The first record of dinocephalians in South America, Late Permian Rio do Rosta Formation) of the Paraná Basin, Brazil. ''Neues Jahrbuch für Geologie und Palaontologie, Abhandlungen''. '''215''': 69-95.</ref>, China (Li & Cheng 1995)<ref name="Cheng & Li 1996">Cheng, Z. & Li, J. (1996) First record of a primitive anteosaurid dinocephalian from the Upper Permian of Gansu, China. ''Vertebrate PalAsiatica.'' '''34''': 123-134.</ref><ref name="Cheng & Li 1997">Cheng, Z. & Li, J. (1997) A new genus of primitive dinocephalian: the third report on Late Permian Dashankou lower tetrapod fauna. ''Vertebrate PalAsiatica.'' '''35''': 35-43.</ref><ref name="Li et al. 1996">Li, J., Rubidge, B.S. & Cheng, Z. (1996) A primitive anteosaurid dinocephalian from China – implications for the distribution of earliest therapsid faunas. ''South African | ||
Rio do Rosta Formation) of the Paraná Basin, Brazil. ''Neues Jahrbuch für Geologie | |||
und Palaontologie, Abhandlungen''. '''215''': 69-95.</ref>, China (Li & Cheng 1995)<ref name="Cheng & Li 1996">Cheng, Z. & Li, J. (1996) First record of a primitive anteosaurid dinocephalian from the Upper Permian of Gansu, China. ''Vertebrate PalAsiatica.'' '''34''': 123-134.</ref><ref name="Cheng & Li 1997">Cheng, Z. & Li, J. (1997) A new genus of primitive dinocephalian: the third report on Late Permian Dashankou lower tetrapod fauna. ''Vertebrate PalAsiatica.'' '''35''': 35-43.</ref><ref name="Li et al. 1996">Li, J., Rubidge, B.S. & Cheng, Z. (1996) A primitive anteosaurid dinocephalian from China – implications for the distribution of earliest therapsid faunas. ''South African | |||
Journal of Science''. '''92''': 252-253.</ref>, Russia<ref name="Ivachnenko 1995">Ivachnenko, M.F. (1995) Primitive Late Permian dinocephalian-titanosuchids of Eastern Europe. ''Paleontological Journal''. '''29''': 120-129.</ref><ref name="Ivachnenko 2000">Ivachnenko, M.F. (2000) Estemmenosuchus and primitive theriodonts from the Late Permian. ''Paleontological Journal''. '''34''': 189-197.</ref> and southern Africa<ref name="Boonstra 1963">Boonstra, L.D. (1963) Diversity within the South African Dinocephalia. ''South African Journal of Science.'' '''59''': 196-206.</ref><ref name="Rubidge 1991"/><ref name="Rubidge 1994">Rubidge, B.S. (1994) ''Australosydon'', the first primitive anteosaurid dinocephalian from the Upper Permian of Gondwana. ''Palaeontology''. '''37''': 579-594.,/ref><ref name="Rubidge & van den Heever 1997">Rubidge, B.S. & van den Heever, J.A. (1997) Morphology and systematic position of the dinocephalian ''Styracocephalus platyrhynchus''. ''Lethaia.'' '''30''': 157-168.</ref><ref name="Lepper et al. 2000">Lepper, J., Raath, M.A. & Rubidge, B.S. (2000) A diverse dinocephalian fauna from | Journal of Science''. '''92''': 252-253.</ref>, Russia<ref name="Ivachnenko 1995">Ivachnenko, M.F. (1995) Primitive Late Permian dinocephalian-titanosuchids of Eastern Europe. ''Paleontological Journal''. '''29''': 120-129.</ref><ref name="Ivachnenko 2000">Ivachnenko, M.F. (2000) Estemmenosuchus and primitive theriodonts from the Late Permian. ''Paleontological Journal''. '''34''': 189-197.</ref> and southern Africa<ref name="Boonstra 1963">Boonstra, L.D. (1963) Diversity within the South African Dinocephalia. ''South African Journal of Science.'' '''59''': 196-206.</ref><ref name="Rubidge 1991"/><ref name="Rubidge 1994">Rubidge, B.S. (1994) ''Australosydon'', the first primitive anteosaurid dinocephalian from the Upper Permian of Gondwana. ''Palaeontology''. '''37''': 579-594.,/ref><ref name="Rubidge & van den Heever 1997">Rubidge, B.S. & van den Heever, J.A. (1997) Morphology and systematic position of the dinocephalian ''Styracocephalus platyrhynchus''. ''Lethaia.'' '''30''': 157-168.</ref><ref name="Lepper et al. 2000">Lepper, J., Raath, M.A. & Rubidge, B.S. (2000) A diverse dinocephalian fauna from | ||
Zimbabwe. ''South African Journal of Science''. '''96''': 403-405.</ref><ref name=Munyikwa 2001">Munyikwa, D. (2001) ''Cranial morphology of a primitive dinocephalian from the | Zimbabwe. ''South African Journal of Science''. '''96''': 403-405.</ref><ref name=Munyikwa 2001">Munyikwa, D. (2001) ''Cranial morphology of a primitive dinocephalian from the | ||
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Dinocephalia was initially considered to comprise six families, namely the | Dinocephalia was initially considered to comprise six families, namely the | ||
Estemmenosuchidae, Brithopodidae, Anteosauridae, Titanosuchidae, Tapinocephalidae and | Estemmenosuchidae, Brithopodidae, Anteosauridae, Titanosuchidae, Tapinocephalidae and | ||
the Styracocephalidae | the Styracocephalidae<ref name="Boonstra 1963"/>(Kemp 1982). In 1988 King reclassified these as | ||
subfamilies within the three families Estemmenosuchidae, Brithopodidae and Titanosuchidae. | subfamilies within the three families Estemmenosuchidae, Brithopodidae and Titanosuchidae. | ||
She considered ''Styracocephalus'', the only genus within the Styracocephalidae as ''incertia sedis''. Later Rubidge and van den Heever (1997), on the basis of the discovery of several | She considered ''Styracocephalus'', the only genus within the Styracocephalidae as ''incertia sedis''. Later Rubidge and van den Heever (1997), on the basis of the discovery of several | ||
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infraorder Dinocephalia (Boonstra 1969). The dentition of this taxon is thought to be more | infraorder Dinocephalia (Boonstra 1969). The dentition of this taxon is thought to be more | ||
specialised than that of other dinocephalians as their canines are reduced in size and resemble | specialised than that of other dinocephalians as their canines are reduced in size and resemble | ||
the incisors (Boonstra 1962, 1963, 1969). Both the incisors and canines are unusual as they have a “talon” and “heel” structure, with the talons of the upper and lower incisors | the incisors (Boonstra 1962, <ref name="Boonstra 1963"/>, 1969). Both the incisors and canines are unusual as they have a “talon” and “heel” structure, with the talons of the upper and lower incisors | ||
intermeshing to create a crushing surface between the heels (Boonstra 1962, 1963, 1969). | intermeshing to create a crushing surface between the heels (Boonstra 1962, <ref name="Boonstra 1963"/>, 1969). | ||
Boonstra (1962) also thought that the postcanine teeth of the maxilla may have intermeshed, | Boonstra (1962) also thought that the postcanine teeth of the maxilla may have intermeshed, | ||
but to a lesser extent. Tapinocephalians thus had crushing surfaces in the front of their | but to a lesser extent. Tapinocephalians thus had crushing surfaces in the front of their | ||
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Boonstra | Boonstra<ref name="Boonstra 1963"/> used the same classifications as Haughton and Brink (1954), but changed | ||
their taxonomic levels to that of subfamily. ''Moschosaurus'' was viewed as being | their taxonomic levels to that of subfamily. ''Moschosaurus'' was viewed as being | ||
morphologically the most primitive of the tapinocephalids yet discovered, and proposed that | morphologically the most primitive of the tapinocephalids yet discovered, and proposed that | ||
all other sub-families diversified from the Moschosaurinae | all other sub-families diversified from the Moschosaurinae<ref name="Boonstra 1963"/>. Boonstra later | ||
thought that the genera ''Moschosaurus'' and ''Struthiocephalellus'' were “growth stages” of | thought that the genera ''Moschosaurus'' and ''Struthiocephalellus'' were “growth stages” of | ||
''Struthiocephalus'' (1969, p39) and thus considered them as synonyms. As a result Boonstra | ''Struthiocephalus'' (1969, p39) and thus considered them as synonyms. As a result Boonstra | ||
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According to Boonstra’s (1969) classification the Tapinocephalinae consisted of the genera | According to Boonstra’s (1969) classification the Tapinocephalinae consisted of the genera | ||
''Keratocephalus'' (with synonym ''Pelosuchus'' (Broom 1905b), see Boonstra 1969), ''Mormosaurus'', ''Phocosaurus'' (Seeley 1888) and ''Tapinocephalus'' (with synonym ''Taurops'', see Boonstra 1969). King (1988) later reclassified Boontsra’s four subfamilies into three tribes, the Struthiocephalini | ''Keratocephalus'' (with synonym ''Pelosuchus'' (Broom 1905b), see Boonstra 1969), ''Mormosaurus'', ''Phocosaurus'' (Seeley 1888) and ''Tapinocephalus'' (with synonym ''Taurops'', see Boonstra 1969). King (1988) later reclassified Boontsra’s four subfamilies into three tribes, the Struthiocephalini<ref name="Boonstra 1963"/>, Tapinocephalini (Gregory 1926) and the Riebeeckosaurini<ref name="Boonstra 1963"/>. The Moschopinae, still considered as a valid taxon by Boonstra (1969), was sunk into the Tapinocephalini by King (1988). These groups were based predominantly on the length of the snout and the degree to which the skull had undergone pachyostosis. The genera contained within each tribe did not differ from Boonstra’s 1969 classification. The Struthiocephalini are characterised as having a reasonably long snout, and having a skull that has undergone moderate pachyostosis (King 1988). Tapinocephalini tend to have short, weak snouts, and great to moderate pachyostification of the skull. Some members of this tribe also have a well developed naso-frontal boss or swollen frons (King 1988). | ||
Revision as of 09:01, 3 February 2009
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Dinocephalia Fossil range: Permian | ||||||||
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Scientific classification | ||||||||
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The infraorder Dinocephalia[1] were a group of medium to large, heavily built therapsid mammal-like reptiles that lived during the Mid-Permian.[2][3] Dinocephalians are among the earliest group of therapsids, and have been found in Brazil[4], China (Li & Cheng 1995)[5][6][7], Russia[8][9] and southern Africa[10][2]Cite error: Closing </ref>
missing for <ref>
tag[11][12][3]. Despite
an early adaptive radiation (Ruben 1986), the dinocephalians became completely extinct by
the end of the Permian, leaving no descendent lineages (Kemp 1982).
Early Taxonomy
Dinocephalia was initially considered to comprise six families, namely the Estemmenosuchidae, Brithopodidae, Anteosauridae, Titanosuchidae, Tapinocephalidae and the Styracocephalidae[10](Kemp 1982). In 1988 King reclassified these as subfamilies within the three families Estemmenosuchidae, Brithopodidae and Titanosuchidae. She considered Styracocephalus, the only genus within the Styracocephalidae as incertia sedis. Later Rubidge and van den Heever (1997), on the basis of the discovery of several additional specimens, confirmed the validity of the genus and considered Styracocephalus to be the sister taxon to the Titanosuchidae. Of the three families of Dinocephalia recognised by King (1988), only the Estemmenosuchidae have yet to be found in Africa[2].
The subfamily Tapinocephalinae (Lydekker 1890) is considered to be the most derived of the
infraorder Dinocephalia (Boonstra 1969). The dentition of this taxon is thought to be more
specialised than that of other dinocephalians as their canines are reduced in size and resemble
the incisors (Boonstra 1962, [10], 1969). Both the incisors and canines are unusual as they have a “talon” and “heel” structure, with the talons of the upper and lower incisors
intermeshing to create a crushing surface between the heels (Boonstra 1962, [10], 1969).
Boonstra (1962) also thought that the postcanine teeth of the maxilla may have intermeshed,
but to a lesser extent. Tapinocephalians thus had crushing surfaces in the front of their
mouths, which was a novel adaptation.
Haughton and Brink (1954) compiled a list of the known genera of Tapinocephalia in South
Africa, recognising five families. These were; Mormosauridae: including the genera
Mormosaurus (Watson 1914), Struthiocephalus (Haughton 1915a), Struthiocephaloides
(Boonstra 1952e), Struthionops (Boonstra 1952b) and Taurocephalus (Broom 1928);
Moschopidae: including the genera Avenantia (Boonstra 1952f), 'Delphinognathus (Seeley
1892), Moschognathus (Broom 1914), Moschcoides (Byrne 1937), Moschops (Broom 1911)
and Pnigalion (Watson 1914); Moschosauridae: including the genera Agnosaurus (Boonstra
1952a) and Moschosaurus (Haughton 1915b); Tapinocephalidae: including the genera
Keratocephalus (von Huene 1931), Riebeeckosaurus (Boonstra 1952c), Tapinocephalus
(Owen 1876) and Taurops (Broom 1912); and Styracocephalidae; including the genus
Styracocephalus (Haughton 1929).
Boonstra[10] used the same classifications as Haughton and Brink (1954), but changed
their taxonomic levels to that of subfamily. Moschosaurus was viewed as being
morphologically the most primitive of the tapinocephalids yet discovered, and proposed that
all other sub-families diversified from the Moschosaurinae[10]. Boonstra later
thought that the genera Moschosaurus and Struthiocephalellus were “growth stages” of
Struthiocephalus (1969, p39) and thus considered them as synonyms. As a result Boonstra
(1969) synonymised the Moschosaurinae with the Struthiocephalinae. The Struthiocephalinae were described by Boonstra (1969) as having a long and fairly strong snout, as well as having
undergone moderate pachyostosis. Other genera of the Struthiocephalinae are
Struthiocephaloides, Struthionops and Taurocephalus.
The Moschopinae consisted of the genera Avenantia, Criocephalus (Broom 1928),
Delphinognathus and Moschops. Criocephalus was renamed Criocephalosaurus in 2002 by Kammerer and Sidor, as the name had been used previously to describe a species of
cerambycid beetle (Mulsant 1839). The genera Agnosaurus, Moschognathus, Moschoides and Pnigalion were synonymised with the genus Moschops (Boonstra 1969). Boonstra (1969)
suggested that Delphinognathus may be a skull of a young Moschops, but did not synonymise the two genera. These tapinocephalids were distinguished as having a short, gently curving snout (Boonstra 1969).
Riebeeckosaurus longirostris (Boonstra 1952c), the only species of the Riebeeckosaurinae, is
known from two skulls. These specimens have a long, slender snout and very narrow
intertemporal region, which forms a sigittal crest at the posterior of the skull (Boonstra 1969).
Boonstra (1969) described the Tapinocephalinae as large to massive tapinocephalids, with a
short, weak snout. Members of this taxon have skulls which are usually heavily pachyostosed,
and have either prominent naso-frontal boss or a greatly swollen frons (Boonstra 1969).
According to Boonstra’s (1969) classification the Tapinocephalinae consisted of the genera
Keratocephalus (with synonym Pelosuchus (Broom 1905b), see Boonstra 1969), Mormosaurus, Phocosaurus (Seeley 1888) and Tapinocephalus (with synonym Taurops, see Boonstra 1969). King (1988) later reclassified Boontsra’s four subfamilies into three tribes, the Struthiocephalini[10], Tapinocephalini (Gregory 1926) and the Riebeeckosaurini[10]. The Moschopinae, still considered as a valid taxon by Boonstra (1969), was sunk into the Tapinocephalini by King (1988). These groups were based predominantly on the length of the snout and the degree to which the skull had undergone pachyostosis. The genera contained within each tribe did not differ from Boonstra’s 1969 classification. The Struthiocephalini are characterised as having a reasonably long snout, and having a skull that has undergone moderate pachyostosis (King 1988). Tapinocephalini tend to have short, weak snouts, and great to moderate pachyostification of the skull. Some members of this tribe also have a well developed naso-frontal boss or swollen frons (King 1988).
References
in progress
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- ↑ 2.0 2.1 2.2 Rubidge, B.S. (1991) A new primitive dinocephalian mammal-like reptile from the Permian of southern Africa. Palaeontology. 34: 547-559.
- ↑ 3.0 3.1 Kemp, T.S. (2005) The origin and evolution of mammals. Oxford University Press, Oxford.
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- ↑ Cheng, Z. & Li, J. (1997) A new genus of primitive dinocephalian: the third report on Late Permian Dashankou lower tetrapod fauna. Vertebrate PalAsiatica. 35: 35-43.
- ↑ Li, J., Rubidge, B.S. & Cheng, Z. (1996) A primitive anteosaurid dinocephalian from China – implications for the distribution of earliest therapsid faunas. South African Journal of Science. 92: 252-253.
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- ↑ 10.0 10.1 10.2 10.3 10.4 10.5 10.6 10.7 Boonstra, L.D. (1963) Diversity within the South African Dinocephalia. South African Journal of Science. 59: 196-206.
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- ↑ Munyikwa, D. (2001) Cranial morphology of a primitive dinocephalian from the Madumabisa Mudstone Formation, Zimbabwe. Unpublished MSc dissertation, University of the Witwatersrand.