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<mods:title id="EDDBB7DFAB35A9C1EB126813A84AF964">A proposed higher taxonomy of anomodont therapsids</mods:title>
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<subSubSection id="C35D6535961A0F14DBAFF9D9FB57F948" pageId="1" pageNumber="2" type="nomenclature">
<paragraph id="8BF836BE961A0F14DBAFF9D9F8E3F9D7" blockId="1.[151,492,1578,1604]" box="[151,492,1578,1604]" pageId="1" pageNumber="2">
<heading id="D0B081D2961A0F14DBAFF9D9F8E3F9D7" bold="true" box="[151,492,1578,1604]" fontSize="11" level="1" pageId="1" pageNumber="2" reason="1">
<emphasis id="B933EAAC961A0F14DBAFF9D9F8E3F9D7" bold="true" box="[151,492,1578,1604]" pageId="1" pageNumber="2">
<taxonomicName id="4C474D3D961A0F14DBAFF9D9F8D3F9D7" authority="Owen, 1860" authorityName="Owen" authorityYear="1860" box="[151,476,1578,1604]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="1" pageNumber="2" phylum="Chordata" rank="order">
Anomodontia
<bibRefCitation id="EFD64B4F961A0F14DA71F9D9F8D3F9D7" author="Owen" box="[329,476,1578,1604]" pageId="1" pageNumber="19" refString="Owen, R. (1860 b) On the orders of fossil and recent Reptilia, and their distribution in time. Report of the Twenty-Ninth Meeting of the British Association for the Advancement of Science, 1859, 153 - 166." type="journal article" year="1860" yearSuffix="b">Owen, 1860</bibRefCitation>
</taxonomicName>
b
</emphasis>
</heading>
</paragraph>
<paragraph id="8BF836BE961A0F14DBAFF980FB57F948" blockId="1.[151,1438,1651,2026]" pageId="1" pageNumber="2">
<emphasis id="B933EAAC961A0F14DBAFF980F828F91E" bold="true" box="[151,295,1651,1677]" pageId="1" pageNumber="2">Definition:</emphasis>
All taxa more closely related to
<taxonomicName id="4C474D3D961A0F14D9F2F987FD84F91D" authority="Owen, 1845" authorityName="Owen" authorityYear="1845" box="[714,1163,1652,1678]" class="Reptilia" family="Dicynodontidae" genus="Dicynodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="lacerticeps">
<emphasis id="B933EAAC961A0F14D9F2F987FAE9F91E" box="[714,998,1652,1677]" italics="true" pageId="1" pageNumber="2">Dicynodon lacerticeps</emphasis>
<bibRefCitation id="EFD64B4F961A0F14D8D7F987FD84F91D" author="Owen" box="[1007,1163,1652,1678]" pageId="1" pageNumber="19" refString="Owen, R. (1845) Report on the reptilian fossils of South Africa. Part I. - Description of certain fossil crania, discovered by A. G. Bain, Esq., in sandstone rocks at the south-eastern extremity of Africa, referable to different species of an extinct genus of Reptilia (Dicynodon), and indicative of a new tribe or sub-order of Sauria. Transactions of the Geological Society of London, Second Series, 7, 59 - 84." type="journal article" year="1845">Owen, 1845</bibRefCitation>
</taxonomicName>
, than
<taxonomicName id="4C474D3D961A0F14DFE0F987F8D7F927" authority="Owen, 1876" authorityName="Owen" authorityYear="1876" class="Insecta" family="Buprestidae" genus="Tapinocephalus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="2" phylum="Arthropoda" rank="species" species="atherstonei">
<emphasis id="B933EAAC961A0F14DFE0F987F823F927" italics="true" pageId="1" pageNumber="2">Tapinocephalus atherstonei</emphasis>
<bibRefCitation id="EFD64B4F961A0F14DA0DF969F8D7F927" author="Owen" box="[309,472,1690,1716]" pageId="1" pageNumber="19" refString="Owen, R. (1876) Descriptive and Illustrated Catalogue of the Fossil Reptilia of South Africa in the Collection of the British Museum. Taylor and Francis, London, 88 pp." type="book" year="1876">Owen, 1876</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C474D3D961A0F14DADFF968FAB6F927" authority="Broom, 1913" authorityName="Broom" authorityYear="1913" box="[487,953,1690,1716]" class="Reptilia" family="Ictidorhinidae" genus="Ictidorhinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="martinsi">
<emphasis id="B933EAAC961A0F14DADFF968FA0FF927" box="[487,768,1691,1716]" italics="true" pageId="1" pageNumber="2">Ictidorhinus martinsi</emphasis>
<bibRefCitation id="EFD64B4F961A0F14D832F969FAB6F927" author="Broom" box="[778,953,1690,1716]" pageId="1" pageNumber="17" refString="Broom, R. (1913 b) On some new carnivorous therapsids. Bulletin of the American Museum of Natural History, 32, 557 - 561." type="journal article" year="1913" yearSuffix="b">Broom, 1913</bibRefCitation>
</taxonomicName>
b,
<taxonomicName id="4C474D3D961A0F14D8E1F968FC7CF927" authority="Owen, 1876" authorityName="Owen" authorityYear="1876" box="[985,1395,1690,1716]" class="Synapsida" family="Gorgonopsidae" genus="Gorgonops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="torvus">
<emphasis id="B933EAAC961A0F14D8E1F968FDC8F927" box="[985,1223,1691,1716]" italics="true" pageId="1" pageNumber="2">Gorgonops torvus</emphasis>
<bibRefCitation id="EFD64B4F961A0F14DFE8F969FC7CF927" author="Owen" box="[1232,1395,1690,1716]" pageId="1" pageNumber="19" refString="Owen, R. (1876) Descriptive and Illustrated Catalogue of the Fossil Reptilia of South Africa in the Collection of the British Museum. Taylor and Francis, London, 88 pp." type="book" year="1876">Owen, 1876</bibRefCitation>
</taxonomicName>
, or
<taxonomicName id="4C474D3D961A0F14DBAFF932FB4BF948" authority="Broom, 1903" authorityName="Broom" authorityYear="1903" box="[151,580,1729,1755]" class="Reptilia" family="Euchambersiidae" genus="Scylacosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="sclateri">
<emphasis id="B933EAAC961A0F14DBAFF932F890F949" box="[151,415,1729,1754]" italics="true" pageId="1" pageNumber="2">Scylacosaurus sclateri</emphasis>
<bibRefCitation id="EFD64B4F961A0F14DA9FF932FB4BF948" author="Broom" box="[423,580,1729,1755]" pageId="1" pageNumber="16" refString="Broom, R. (1903 b) On some new primitive theriodonts in the South African Museum. Annals of the South African Museum, 4, 147 - 158." type="journal article" year="1903" yearSuffix="b">Broom, 1903</bibRefCitation>
</taxonomicName>
b.
</paragraph>
</subSubSection>
<subSubSection id="C35D6535961A0F17DBFEF914FBCDFA46" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2" type="description">
<paragraph id="8BF836BE961A0F17DBFEF914FDE0FD43" blockId="1.[151,1438,1651,2026]" lastBlockId="2.[151,1437,152,1493]" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2">
<emphasis id="B933EAAC961A0F14DBFEF914F8EFF892" bold="true" box="[198,480,1767,1793]" pageId="1" pageNumber="2">Taxonomic comments:</emphasis>
<taxonomicName id="4C474D3D961A0F14DADFF91BFB87F891" box="[487,648,1768,1794]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="1" pageNumber="2" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
has had a convoluted taxonomic history, and usage of the term has gone through several major phases.
<collectingCountry id="F350762E961A0F14D972F8FDFB64F8BB" box="[586,619,1806,1832]" name="American Samoa" pageId="1" pageNumber="2">As</collectingCountry>
initially conceived by
<bibRefCitation id="EFD64B4F961A0F14D8BAF8FDFD3CF8BB" author="Owen" box="[898,1075,1806,1832]" pageId="1" pageNumber="19" refString="Owen, R. (1860 b) On the orders of fossil and recent Reptilia, and their distribution in time. Report of the Twenty-Ninth Meeting of the British Association for the Advancement of Science, 1859, 153 - 166." type="journal article" year="1860" yearSuffix="b">Owen (1860b)</bibRefCitation>
,
<taxonomicName id="4C474D3D961A0F14DF78F8FDFDEAF8BB" box="[1088,1253,1806,1832]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="1" pageNumber="2" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
contained taxa currently considered to be dicynodonts (Owens “families” Dicynodontia, for
<taxonomicName id="4C474D3D961A0F14DF14F8C6FDA2F8DD" box="[1068,1197,1845,1870]" class="Reptilia" family="Dicynodontidae" genus="Dicynodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis id="B933EAAC961A0F14DF14F8C6FDA2F8DD" box="[1068,1197,1845,1870]" italics="true" pageId="1" pageNumber="2">Dicynodon</emphasis>
</taxonomicName>
and
<taxonomicName id="4C474D3D961A0F14DFDEF8C6FC9BF8DD" box="[1254,1428,1845,1870]" class="Malacostraca" family="Varunidae" genus="Ptychognathus" kingdom="Animalia" order="Decapoda" pageId="1" pageNumber="2" phylum="Arthropoda" rank="genus">
<emphasis id="B933EAAC961A0F14DFDEF8C6FC9BF8DD" box="[1254,1428,1845,1870]" italics="true" pageId="1" pageNumber="2">Ptychognathus</emphasis>
</taxonomicName>
, and Cryptodontia, for
<taxonomicName id="4C474D3D961A0F14DA81F8AFFB42F8E6" box="[441,589,1884,1909]" class="Reptilia" family="Cryptodontidae" genus="Oudenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis id="B933EAAC961A0F14DA81F8AFFB42F8E6" box="[441,589,1884,1909]" italics="true" pageId="1" pageNumber="2">Oudenodon</emphasis>
</taxonomicName>
) as well as non-synapsid reptiles (“family” Gnathodontia for the archosauromorph
<taxonomicName id="4C474D3D961A0F14DA4BF870FB22F80F" box="[371,557,1923,1948]" class="Reptilia" family="Rhynchosauridae" genus="Rhynchosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rhynchosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis id="B933EAAC961A0F14DA4BF870FB22F80F" box="[371,557,1923,1948]" italics="true" pageId="1" pageNumber="2">Rhynchosaurus</emphasis>
</taxonomicName>
).
<bibRefCitation id="EFD64B4F961A0F14D97FF871FBE7F80F" author="Owen" box="[583,744,1922,1948]" pageId="1" pageNumber="19" refString="Owen, R. (1861) Palaeontology, or a Systematic Summary of Extinct Animals and Their Geological Relations, 2 nd Edition. Adam and Charles Black, Edinburgh, 463 pp." type="book" year="1861">Owen (1861)</bibRefCitation>
removed Gnathodontia from
<taxonomicName id="4C474D3D961A0F14DF6AF871FDF7F80F" box="[1106,1272,1922,1948]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="1" pageNumber="2" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
and included Cynodontia instead. In Owens (1876) monographic treatment of South African fossil reptiles, he removed cynodonts from
<taxonomicName id="4C474D3D961A0F14DA4FF823FB22F879" box="[375,557,2000,2026]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="1" pageNumber="2" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
, restricting it to dicynodonts (Dicynodontia, Cryptodontia, and Endothiodontia). In the late 19th and early 20th century, various workers (
<emphasis id="B933EAAC96190F17DF2EFF6AFD2CFF21" box="[1046,1059,153,178]" italics="true" pageId="2" pageNumber="3">e</emphasis>
.
<emphasis id="B933EAAC96190F17DF13FF6AFD35FF21" box="[1067,1082,153,178]" italics="true" pageId="2" pageNumber="3">g</emphasis>
.,
<bibRefCitation id="EFD64B4F96190F17DF6AFF6AFDE7FF20" author="Seeley" box="[1106,1256,153,179]" pageId="2" pageNumber="20" refString="Seeley, H. G. (1888) Researches on the structure, organisation, and classification of the fossil Reptilia. VI. On the anomodont Reptilia and their allies [Abstract]. Proceedings of the Royal Society of London, 44, 381 - 383." type="journal article" year="1888">Seeley 1888</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17DFCEFF6AFC9BFF20" author="Watson" box="[1270,1428,153,179]" pageId="2" pageNumber="20" refString="Watson, D. M. S. (1917) A sketch classification of the pre-Jurassic tetrapod vertebrates. Proceedings of the Zoological Society of London, 1917, 167 - 186." type="proceedings paper" year="1917">Watson 1917</bibRefCitation>
) expanded the composition of
<taxonomicName id="4C474D3D96190F17DAC2FF4CFB91FF4A" box="[506,670,191,217]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="2" pageNumber="3" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
to include all non-mammalian synapsids.
<bibRefCitation id="EFD64B4F96190F17DFA1FF4CFC4BFF4A" author="Broom" box="[1177,1348,191,217]" pageId="2" pageNumber="16" refString="Broom, R. (1905) On the use of the term Anomodontia. Records of the Albany Museum, 1, 266 - 269." type="journal article" year="1905">Broom (1905)</bibRefCitation>
argued that
<taxonomicName id="4C474D3D96190F17DBF2FF15F864FE93" box="[202,363,230,256]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="2" pageNumber="3" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
should be restricted to dicynodonts, but this usage did not regain wide acceptance until the 1920s and 1930s (
<emphasis id="B933EAAC96190F17DA43FEFEF887FEB5" box="[379,392,269,294]" italics="true" pageId="2" pageNumber="3">e</emphasis>
.
<emphasis id="B933EAAC96190F17DAA9FEFEF8AFFEB5" box="[401,416,269,294]" italics="true" pageId="2" pageNumber="3">g</emphasis>
.,
<bibRefCitation id="EFD64B4F96190F17DA81FEFFFB6FFEB5" author="Nopcsa" box="[441,608,268,294]" pageId="2" pageNumber="19" refString="Nopcsa, F. (1923) Die Familien der Reptilien. Verlag von Gebruder Borntraeger, Berlin, 210 pp." type="book" year="1923">Nopcsa 1923</bibRefCitation>
,
<bibRefCitation id="EFD64B4F96190F17D955FEFFFBA1FEB5" author="Nopcsa" box="[621,686,268,294]" pageId="2" pageNumber="19" refString="Nopcsa, F. (1928) The genera of reptiles. Palaeobiologica, 1, 163 - 188." type="journal article" year="1928">1928</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17D984FEFFFA52FEB5" author="Broom" box="[700,861,268,294]" pageId="2" pageNumber="17" refString="Broom, R. (1932) The Mammal-Like Reptiles of South Africa and the Origin of Mammals. H. F. &amp; G. Witherby, London, 376 pp." type="book" year="1932">Broom 1932</bibRefCitation>
). Watson &amp; Romers (1956) highly influential classification of therapsids united dicynodonts, the Russian “venyukoviamorphs”, and herbivorous dinocephalians (tapinocephalians) within
<taxonomicName id="4C474D3D96190F17D9B5FEA9FA39FEE7" box="[653,822,346,372]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="2" pageNumber="3" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
. Later,
<bibRefCitation id="EFD64B4F96190F17D8B7FEA9FD36FEE7" author="Romer" box="[911,1081,346,372]" pageId="2" pageNumber="20" refString="Romer, A. S. (1966) Vertebrate Paleontology, 3 rd Edition. The University of Chicago Press, Chicago, 468 pp." type="book" year="1966">Romer (1966)</bibRefCitation>
also included the supposedly carnivorous dinocephalians (
<taxonomicName id="4C474D3D96190F17DAD2FE72FB99FE08" box="[490,662,385,411]" class="Reptilia" family="Anteosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="2" pageNumber="3" phylum="Chordata" rank="family">Anteosauridae</taxonomicName>
and
<taxonomicName id="4C474D3D96190F17D9E8FE72FA89FE08" box="[720,902,385,411]" class="Reptilia" family="Titanosuchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="2" pageNumber="3" phylum="Chordata" rank="family">Titanosuchidae</taxonomicName>
, the latter of which are now believed to have been herbivores [
<emphasis id="B933EAAC96190F17DA59FE5BF861FE52" box="[353,366,424,449]" italics="true" pageId="2" pageNumber="3">e</emphasis>
.
<emphasis id="B933EAAC96190F17DA4EFE5BF88AFE52" box="[374,389,424,449]" italics="true" pageId="2" pageNumber="3">g</emphasis>
.,
<bibRefCitation id="EFD64B4F96190F17DAA2FE54FB46FE52" author="Boonstra" box="[410,585,423,449]" pageId="2" pageNumber="16" refString="Boonstra, L. D. (1963) Diversity within the South African Dinocephalia. South African Journal of Science, 59, 176 - 195." type="journal article" year="1963">Boonstra 1963</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17D96DFE54FBEEFE52" author="Kemp" box="[597,737,423,449]" pageId="2" pageNumber="18" refString="Kemp, T. S. (1982) Mammal-Like Reptiles and the Origin of Mammals. Academic Press, New York, 363 pp." type="book" year="1982">Kemp 1982</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17D9D5FE54FD20FE52" author="Hopson" box="[749,1071,423,449]" pageId="2" pageNumber="17" refString="Hopson, J. A. &amp; Barghusen, H. R. (1986) An analysis of therapsid relationships. In: Hotton, N., III, MacLean, P., Roth, J. &amp; Roth, E. (Eds.), The Ecology and Biology of the Mammal-Like Reptiles. Smithsonian Institution Press, Washington, DC, pp. 83 - 106." type="book chapter" year="1986">Hopson &amp; Barghusen 1986</bibRefCitation>
]) within
<taxonomicName id="4C474D3D96190F17DFA0FE54FC33FE52" box="[1176,1340,423,449]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="2" pageNumber="3" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
, and the use of
<taxonomicName id="4C474D3D96190F17DBDDFE3DF889FE7B" box="[229,390,462,488]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="2" pageNumber="3" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
to refer to
<taxonomicName id="4C474D3D96190F17D931FE3DFBAAFE7B" authority="Dicynodontia" authorityName="Dicynodontia" box="[521,677,462,488]" class="Reptilia" family="Anteosauridae" genus="Dinocephalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">Dinocephalia</taxonomicName>
+ Dicynodontia was dominant until the 1990s (
<emphasis id="B933EAAC96190F17DFEDFE3DFDEDFE74" box="[1237,1250,462,487]" italics="true" pageId="2" pageNumber="3">e</emphasis>
.
<emphasis id="B933EAAC96190F17DFD2FE3DFDF7FE74" box="[1258,1272,462,487]" italics="true" pageId="2" pageNumber="3">g</emphasis>
.,
<bibRefCitation id="EFD64B4F96190F17DE36FE3DFC80FE7B" author="King" box="[1294,1423,462,488]" pageId="2" pageNumber="18" refString="King, G. M. (1988) Anomodontia. Handbuch der Palaoherpetologie 17 C. Gustav Fischer Verlag, Stuttgart, 174 pp." type="book" year="1988">King 1988</bibRefCitation>
). With the application of cladistic methodology to studying therapsid relationships, the monophyly of
<taxonomicName id="4C474D3D96190F17DBAFFDE8F836FDA6" authority="Dicynodontia" authorityName="Dicynodontia" box="[151,313,539,565]" class="Reptilia" family="Anteosauridae" genus="Dinocephalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">Dinocephalia</taxonomicName>
+ Dicynodontia was called into question (
<bibRefCitation id="EFD64B4F96190F17D807FDE8FD9EFDA6" author="Hopson" box="[831,1169,539,565]" pageId="2" pageNumber="17" refString="Hopson, J. A. &amp; Barghusen, H. R. (1986) An analysis of therapsid relationships. In: Hotton, N., III, MacLean, P., Roth, J. &amp; Roth, E. (Eds.), The Ecology and Biology of the Mammal-Like Reptiles. Smithsonian Institution Press, Washington, DC, pp. 83 - 106." type="book chapter" year="1986">Hopson &amp; Barghusen 1986</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17DFA6FDE8FC98FDA6" author="Gauthier" box="[1182,1431,539,565]" pageId="2" pageNumber="17" refString="Gauthier, J., Kluge, A. G. &amp; Rowe, T. (1988) Amniote phylogeny and the importance of fossils. Cladistics, 4, 105 - 209." type="journal article" year="1988">
Gauthier
<emphasis id="B933EAAC96190F17DE29FDE8FC40FDA7" box="[1297,1359,539,564]" italics="true" pageId="2" pageNumber="3">et al.</emphasis>
1988
</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17DBAFFDB1F833FDCF" author="Hopson" box="[151,316,578,604]" pageId="2" pageNumber="17" refString="Hopson, J. A. (1991) Systematics of the nonmammalian Synapsida and implication for patterns of evolution in synapsids. In: Schultze, H. - P. &amp; Trueb, L. (Eds.), Origin of the Higher Groups of Tetrapods: Controversy and Consensus. Cornell University Press, Ithaca, pp. 635 - 693." type="book chapter" year="1991">Hopson 1991</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17DA71FDB1F8D9FDCF" author="Grine" box="[329,470,578,604]" pageId="2" pageNumber="17" refString="Grine, F. E. (1997) Dinocephalians are not anomodonts. Journal of Vertebrate Paleontology, 17, 177 - 183." type="journal article" year="1997">Grine 1997</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17DADBFDB1FBFDFDCF" author="Sidor" box="[483,754,578,604]" pageId="2" pageNumber="20" refString="Sidor, C. A. &amp; Hopson, J. A. (1998) Ghost lineages and &quot; mammalness &quot;: assessing the temporal pattern of character acquisition in the Synapsida. Paleobiology, 24, 254 - 273." type="journal article" year="1998">Sidor &amp; Hopson 1998</bibRefCitation>
), and dinocephalians were removed from
<taxonomicName id="4C474D3D96190F17DFD7FDB1FC97FDCF" box="[1263,1432,578,604]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="2" pageNumber="3" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
. Current usage of
<taxonomicName id="4C474D3D96190F17DA5CFD9AFB0AFD10" box="[356,517,617,643]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="2" pageNumber="3" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
universally refers to the clade containing both dicynodonts and the basal taxa formerly called “dromasaurs” or “venyukoviamorphs” (
<emphasis id="B933EAAC96190F17D81DFD63FA3DFD3A" box="[805,818,656,681]" italics="true" pageId="2" pageNumber="3">e</emphasis>
.
<emphasis id="B933EAAC96190F17D801FD63FA48FD3A" box="[825,839,656,681]" italics="true" pageId="2" pageNumber="3">g</emphasis>
.,
<bibRefCitation id="EFD64B4F96190F17D865FD7CFD45FD3A" author="Modesto" box="[861,1098,655,681]" pageId="2" pageNumber="19" refString="Modesto, S. P., Rubidge, B. S. &amp; Welman, J. (1999) The most basal anomodont therapsid and the primacy of Gondwana in the evolution of the anomodonts. Proceedings of the Royal Society of London. Series B: Biological Sciences, 266, 331 - 337." type="journal article" year="1999">
Modesto
<emphasis id="B933EAAC96190F17D8F3FD63FD0AFD3A" box="[971,1029,656,681]" italics="true" pageId="2" pageNumber="3">et al.</emphasis>
1999
</bibRefCitation>
,
<bibRefCitation id="EFD64B4F96190F17DF6EFD7CFDAEFD3A" author="Modesto" box="[1110,1185,655,681]" pageId="2" pageNumber="19" refString="Modesto, S. P., Rubidge, B. S., Visser, I. &amp; Welman, J. (2003 a) A new basal dicynodont from the Upper Permian of South Africa. Palaeontology, 46, 211 - 223." type="journal article" year="2003" yearSuffix="a">2003a</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17DF95FD7CF9DAFD43" author="Modesto" pageId="2" pageNumber="19" refString="Modesto, S. P. &amp; Rubidge, B. S. (2000) A basal anomodont therapsid from the lower Beaufort Group, Upper Permian of South Africa. Journal of Vertebrate Paleontology, 20, 515 - 521." type="journal article" year="2000">Modesto &amp; Rubidge 2000</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17DBD9FD45FB3CFD43" author="Angielczyk" box="[225,563,694,720]" pageId="2" pageNumber="16" refString="Angielczyk, K. D. &amp; Kurkin, A. A. (2003 a) Phylogenetic analysis of Russian Permian dicynodonts (Therapsida: Anomodontia): implications for Permian biostratigraphy and Pangaean biogeography. Zoological Journal of the Linnean Society, 139, 157 - 212." type="journal article" year="2003" yearSuffix="a">Angielczyk &amp; Kurkin 2003a</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17D907FD45FA04FD43" author="Angielczyk" box="[575,779,694,720]" pageId="2" pageNumber="16" refString="Angielczyk, K. D. (2004) Phylogenetic evidence for and implications of a dual origin of propaliny in anomodont therapsids (Synapsida). Paleobiology, 30, 268 - 296." type="journal article" year="2004">Angielczyk 2004</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17D82FFD45FACCFD43" author="Frobisch" box="[791,963,694,720]" pageId="2" pageNumber="17" refString="Frobisch, J. (2007) The cranial anatomy of Kombuisia frerensis Hotton (Synapsida, Dicynodontia) and a new phylogeny of anomodont therapsids. Zoological Journal of the Linnean Society, 150, 117 - 144." type="journal article" year="2007">Fröbisch 2007</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17D8F7FD45FDECFD43" author="Frobisch" box="[975,1251,694,720]" pageId="2" pageNumber="17" refString="Frobisch, J. &amp; Reisz, R. R. (2008) A new species of Emydops (Synapsida, Anomodontia) and a discussion of dental variability and pathology in dicynodonts. Journal of Vertebrate Paleontology, 28, 770 - 787." type="journal article" year="2008">Fröbisch &amp; Reisz 2008</bibRefCitation>
).
</paragraph>
<paragraph id="8BF836BE96190F17DBFEFD2FFD2FFB7E" blockId="2.[151,1437,152,1493]" pageId="2" pageNumber="3">
<bibRefCitation id="EFD64B4F96190F17DBFEFD2FFB60FD65" author="Modesto" box="[198,623,732,758]" pageId="2" pageNumber="19" refString="Modesto, S. P. &amp; Rubidge, B. S. (2000) A basal anomodont therapsid from the lower Beaufort Group, Upper Permian of South Africa. Journal of Vertebrate Paleontology, 20, 515 - 521." type="journal article" year="2000" yearSuffix="p">Modesto &amp; Rubidge (2000, p. 516)</bibRefCitation>
proposed a node-based definition for
<taxonomicName id="4C474D3D96190F17DF03FD2FFDD3FD65" box="[1083,1244,732,758]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="2" pageNumber="3" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
containing “the last common ancestor of
<emphasis id="B933EAAC96190F17DA87FCF0FBFFFC8F" box="[447,752,771,796]" italics="true" pageId="2" pageNumber="3">Anomocephalus africanus</emphasis>
,
<taxonomicName id="4C474D3D96190F17D9C7FCF0FD2AFC8F" box="[767,1061,771,796]" class="Reptilia" family="Galechiridae" genus="Patranomodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="nyaphulii">
<emphasis id="B933EAAC96190F17D9C7FCF0FD2AFC8F" box="[767,1061,771,796]" italics="true" pageId="2" pageNumber="3">Patranomodon nyaphulii</emphasis>
</taxonomicName>
,
<taxonomicName id="4C474D3D96190F17DF0CFCF0FDFCFC8F" box="[1076,1267,771,796]" class="Reptilia" family="Galeopsidae" genus="Galeops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="whaitsi">
<emphasis id="B933EAAC96190F17DF0CFCF0FDFCFC8F" box="[1076,1267,771,796]" italics="true" pageId="2" pageNumber="3">Galeops whaitsi</emphasis>
</taxonomicName>
,
<taxonomicName id="4C474D3D96190F17DE39FCF0F813FCD0" class="Reptilia" family="Eodicynodontidae" genus="Eodicynodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="oosthuizeni">
<emphasis id="B933EAAC96190F17DE39FCF0F813FCD0" italics="true" pageId="2" pageNumber="3">Eodicynodon oosthuizeni</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C474D3D96190F17DA64FCD9FB6AFCD0" box="[348,613,810,835]" class="Reptilia" family="Dicynodontidae" genus="Dicynodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="lacerticeps">
<emphasis id="B933EAAC96190F17DA64FCD9FB6AFCD0" box="[348,613,810,835]" italics="true" pageId="2" pageNumber="3">Dicynodon lacerticeps</emphasis>
</taxonomicName>
, and all its descendants.” We advocate a stem-based definition for this taxon instead, utilizing exemplars from all of the other major therapsid groups (
<taxonomicName id="4C474D3D96190F17DF7CFCA2F83DFC02" authority="Biarmosuchia, Gorgonopsia" authorityName="Biarmosuchia, Gorgonopsia" class="Reptilia" family="Anteosauridae" genus="Dinocephalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">Dinocephalia, Biarmosuchia, Gorgonopsia</taxonomicName>
, and Eutheriodontia) as external specifiers. Although relationships between the major therapsid taxa remain highly contentious (Rubidge &amp; Sidor 2001;
<bibRefCitation id="EFD64B4F96190F17D800FC6DFAC6FC2B" author="Kemp" box="[824,969,926,952]" pageId="2" pageNumber="18" refString="Kemp, T. S. (2006) The origin and early radiation of the therapsid mammal-like reptiles: a palaeobiological hypothesis. Journal of Evolutionary Biology, 19, 1231 - 1247." type="journal article" year="2006">Kemp 2006</bibRefCitation>
), the groups themselves have remained remarkably stable, and current phylogenies (
<bibRefCitation id="EFD64B4F96190F17D99AFC37FAE6FC4D" author="Hopson" box="[674,1001,964,990]" pageId="2" pageNumber="17" refString="Hopson, J. A. &amp; Barghusen, H. R. (1986) An analysis of therapsid relationships. In: Hotton, N., III, MacLean, P., Roth, J. &amp; Roth, E. (Eds.), The Ecology and Biology of the Mammal-Like Reptiles. Smithsonian Institution Press, Washington, DC, pp. 83 - 106." type="book chapter" year="1986">Hopson &amp; Barghusen 1986</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17D8CEFC37FD75FC4D" author="Sidor" box="[1014,1146,964,990]" pageId="2" pageNumber="20" refString="Sidor, C. A. (2000) Evolutionary Trends and Relationships Within the Synapsida. Unpublished PhD thesis, University of Chicago." type="book" year="2000">Sidor 2000</bibRefCitation>
) support their reciprocal monophyly. Use of a stem-based definition for the major therapsid clade
<taxonomicName id="4C474D3D96190F17DF37FC18FDBBFB96" box="[1039,1204,1003,1029]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="2" pageNumber="3" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
allows for greatest inclusivity in the likely event that a therapsid basal to
<emphasis id="B933EAAC96190F17D80DFBE1FAF8FBB8" box="[821,1015,1042,1067]" italics="true" pageId="2" pageNumber="3">Anomocephalus</emphasis>
with anomodont characteristics is discovered.
<bibRefCitation id="EFD64B4F96190F17DA1CFBCAF8C8FBC0" author="Battail" box="[292,455,1081,1107]" pageId="2" pageNumber="16" refString="Battail, B. (2000) A comparison of Late Permian Gondwanan and Laurasian amniote faunas. Journal of African Earth Sciences, 31, 165 - 174." type="journal article" year="2000">Battail (2000)</bibRefCitation>
suggested that the problematic Chinese taxon
<taxonomicName id="4C474D3D96190F17D8DFFBCAFD6CFBC1" box="[999,1123,1081,1106]" class="Reptilia" family="Biseridensidae" genus="Biseridens" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B933EAAC96190F17D8DFFBCAFD6CFBC1" box="[999,1123,1081,1106]" italics="true" pageId="2" pageNumber="3">Biseridens</emphasis>
</taxonomicName>
, originally described as an “eotitanosuchian” (
<bibRefCitation id="EFD64B4F96190F17DA40FBACFB44FBEA" author="Li" box="[376,587,1119,1145]" pageId="2" pageNumber="18" refString="Li, J. - L. &amp; Cheng, Z. - W. (1997) First discovery of eotitanosuchian (Therapsida, Synapsida) of China. Vertebrata PalAsiatica, 35, 268 - 282." type="journal article" year="1997">Li &amp; Cheng 1997</bibRefCitation>
), might represent just such a therapsid. Usage of a node-based definition for
<taxonomicName id="4C474D3D96190F17DBF9FB75F86CFB33" box="[193,355,1158,1184]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="2" pageNumber="3" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
ultimately would have destabilizing results for therapsid taxonomy, necessitating either the creation of new clade names for each node outward from
<emphasis id="B933EAAC96190F17D86FFB5EFD18FB55" box="[855,1047,1197,1222]" italics="true" pageId="2" pageNumber="3">Anomocephalus</emphasis>
to refer to all anomodont-like therapsids, or redefinition of
<taxonomicName id="4C474D3D96190F17DAD4FB20FB82FB7E" box="[492,653,1235,1261]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="2" pageNumber="3" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
with reference to each new taxon.
</paragraph>
<paragraph id="8BF836BE96190F17DBFEFB0AFBCDFA46" blockId="2.[151,1437,152,1493]" pageId="2" pageNumber="3">
<emphasis id="B933EAAC96190F17DBFEFB0AF8A0FA80" bold="true" box="[198,431,1273,1299]" pageId="2" pageNumber="3">Revised diagnosis:</emphasis>
Only three phylogenetic analyses (
<bibRefCitation id="EFD64B4F96190F17D86DFB09FD44FA87" author="Modesto" box="[853,1099,1274,1300]" pageId="2" pageNumber="19" refString="Modesto, S. P., Rubidge, B. S. &amp; Welman, J. (1999) The most basal anomodont therapsid and the primacy of Gondwana in the evolution of the anomodonts. Proceedings of the Royal Society of London. Series B: Biological Sciences, 266, 331 - 337." type="journal article" year="1999">
Modesto
<emphasis id="B933EAAC96190F17D8FFFB09FD0CFA80" box="[967,1027,1274,1299]" italics="true" pageId="2" pageNumber="3">et al.</emphasis>
1999
</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17DF61FB09FC08FA87" author="Frobisch" box="[1113,1287,1274,1300]" pageId="2" pageNumber="17" refString="Frobisch, J. (2007) The cranial anatomy of Kombuisia frerensis Hotton (Synapsida, Dicynodontia) and a new phylogeny of anomodont therapsids. Zoological Journal of the Linnean Society, 150, 117 - 144." type="journal article" year="2007">Fröbisch 2007</bibRefCitation>
;
<bibRefCitation id="EFD64B4F96190F17DE2DFB09F812FAA8" author="Frobisch" pageId="2" pageNumber="17" refString="Frobisch, J. &amp; Reisz, R. R. (2008) A new species of Emydops (Synapsida, Anomodontia) and a discussion of dental variability and pathology in dicynodonts. Journal of Vertebrate Paleontology, 28, 770 - 787." type="journal article" year="2008">Fröbisch &amp; Reisz 2008</bibRefCitation>
) have included
<emphasis id="B933EAAC96190F17DAE3FAD2FA01FAA9" box="[475,782,1313,1338]" italics="true" pageId="2" pageNumber="3">Anomocephalus africanus</emphasis>
, the most basal anomodont recognized to date, and we use these papers as sources of autapomorphies for
<taxonomicName id="4C474D3D96190F17D9DFFAB4FA83FAF2" box="[743,908,1351,1377]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="2" pageNumber="3" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
. The most consistently cited autapomorphies of
<taxonomicName id="4C474D3D96190F17DB80FA9DF855FA1B" box="[184,346,1390,1416]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomodontia" pageId="2" pageNumber="3" phylum="Chordata" rank="order">Anomodontia</taxonomicName>
are: (1) serrations absent on marginal teeth; (2) mandibular fenestra present; (3) zygomatic arch bowed dorsally. A complete list of autapomorphies from each paper for this and all subsequent clades discussed herein can be found in the Appendix.
</paragraph>
</subSubSection>
</treatment>
</document>